7 Americana A Journal of Entomology. Volume XLI (New Series) for 1961 PUBLICATION COMMITTEE JAMES A. SLATER, EDITOR GEORGE S. TULLOCH JOHN HANSON PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY 1961 ENTOMOLOGICA AMERICANA Vol. XLI (N.S.), for 1961 CONTENTS A Revision of the Genus Copris Muller of the Western Hemisphere (Coleoptera, Scarabaeidae). Eric G. Matthews j Pages 1-137 VOL. XLI (NEW SERIES) A Journal of Entomology. PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY PUBLICATION COMMITTEE JAMES A. SLATER, EDITOR GEORGE S. TULLOCH JOHN HANSON Published for the Society by the Business Press Inc. N. Queen St. and McGovern Ave., Lancaster, Pa. Subscription $6.00 Per Year Date of Issue April 9, 1962 AmerigAna VOLUME XLI A REVISION OF THE GENUS COPRIS MULLER OF THE WESTERN HEMISPHERE (COLEOPTERA. SCARABAEIDAE) By Eric G. Matthews1 TABLE OF CONTENTS Page Introduction 2 Materials and Methods 6 Acknowledgements 7 Morphology 8 Microsculpture 8 Head 9 Mouthparts 10 Prothorax 10 Pterothorax 11 Elytra 12 Legs 12 Abdomen 13 Genitalia 13 Sexual Dimorphism 13 The Sexual Armament in Copris 14 Intraspecific Variation in Horn Height 15 Interspecific Variation in Horn Height 26 Biology 27 Taxonomy 33 Genus Copris Muller, 1764 35 Key to the Species of the Genus Copris in the Western Hemisphere Based on the Major Males 35 Key to the Species of the Genus Copris in the Western Hemisphere Based on the Males and Females 39 Group I. The minutns group " 43 Complex 1. The incertus complex 44 1 Dept, of Biology, U. of Puerto Rico, Rio Piedras, formerly at Cornell University, Ithaca, New York. 1 ENTOMOLOGICA AMERICANA Complex 2. The minutus complex 56 Group II. The fricator group 63 Complex 1. The armatus complex 64 Complex 2. The arizonensis complex 84 Complex 3. The remotus complex 87 Complex 4. The rebouchci complex 98 Complex 5. The fricator complex 107 Literature Cited 118 Plates 121 INTRODUCTION Copris Muller is a genus of principally Old World coprophagous scarabs represented in the New World primarily in North and Central America. The present work deals only with the adults of the Western Hemisphere species. The name Copris is fairly old; it was coined by Geoffroy (1764) to include all those species in the Linnaean genus Scarabaeus characterized by the absence of a visible scutellum. However, since Geoffroy ’s work has been discarded for nomenclatorial pur- poses*, the first valid user of the name was Muller (1764), who included an indication but no species. The first valid inclusion of species was by Fourcroy (1785), who included ten species, the first of which was lunaris L. Fabricius did not adopt the name Copris until 1798 in the Supplementum Entomologiae Systematicae. Thereafter the name was in wide use but the first valid type designation appears to be that of Curtis (1832), who designated Scarabaeus lunaris Linnaeus as the type of the genus. Following Olivier’s (1790) very broad concept of the genus (including even many scutellate dung beetles) numerous groups were separated off as independent genera and by the end of the first third of the 19th century the name Copris had become re- stricted to a fairly homogeneous assemblage. In 1837 Hope pro- posed the first of several further subdivisions, in which he was followed by Burmeister (1846) and Erichson (1847). These sub- divisions Lacordaire (1856) did not accept, preferring to go back to the broader concept. Nevertheless, the genera and subgenera proposed by Hope, Burmeister, and Erichson subsequently became largely accepted and it was the latter author who formulated the restricted concept of Copris which exists, in largely unmodified form, today, although several additional small genera have been separated off since. Recently Balthasar (1958) divided the genus into four subgenera (one of which is Litocopris Waterhouse, 1891). * Opinions and declarations rendered by the International Commission on Zoological Nomenclature, Vol. 4, Opinion 228, 1954. 2 ENTOMOLOGICA AMERICANA All the American species belong to the subgenus C opr is as under- stood by Balthasar. There have been very few attempts at any revisional study of the large genus Copris and these have been restricted to relatively small geographical areas. The American species of the genus have been investigated very perfunctorily from two geographical vantage points — the United States and Central Mexico — with no attempt, until now, at coordinating the results. Fortunately, the two areas are sufficiently distinct zoogeographically that a minimum of synonymization has resulted. All the literature consists of scat- tered new species descriptions with four exceptions : Harold (1869), while describing most of the Mexican species, presented a key to them with short discussions of means of distinguishing them ; Horn (1873) presented a key and descriptions of the four United States species known at the time; Bates (1887-1889), in the Biologia Centrali Americana, reviewed the Mexican and Central American species known to him; and Schaeffer (1906) presented a short but quite usable key to the United States species known at that time. In his review, Bates used only Harold’s names, even though he was looking at several undescribed species which he attempted to fit into Harold’s descriptions. As a result, the range extensions given by Bates for Harold’s species, repeated in all subsequent catalogues and lists, are largely erroneous. There followed some individual species descriptions and Gillet (1911), in the Coleoptero- rum Catalogus, lists 16 species and two “varieties” for the Western Hemisphere. Thereafter there were several more species descrip- tions. The Leng Catalogue and Supplement (1920-1948) list eight species for the United States and Canada, while Blackwelder (1944) lists 12 species and two varieties for America exclusive of these two countries, of which one species and one variety are listed as common to both areas. Subsequently Pereira and d’An- dretta (1955) synonymized one name, Matthews and Halffter (1959) described five new species and synonymized one name, and Matthews (1959) described one additional species and elevated one variety to species rank, presenting a key to the males of the Mexi- can species. There were until now, therefore, 24 species and one variety known from the Western Hemisphere. The present work describes one new species and three new subspecies, elevates the one re- maining variety to species rank, lowers two species to subspecies level, synonymizes two names and resurrects one, bringing the total to 23 species and five subspecies known for this hemisphere at present. 3 ENTOMOLOGICA AMERICANA In the world as a whole the genus at present contains ap- proximately 160 described species which are distributed about as follows among the zoogeographical regions (with some species counted twice) : Ethiopian — 77, Oriental — 46, Palaearctic — 27, Ne- artic — 16, Neotropical — 8. It does not occur in Madagascar or in Australia. The northern limits of the genus in the Eastern Hemi- sphere appear to coincide approximately with the 50th parallel (Kolbe, 1905), except that one species occurs very locally in southern England. Of the Coprini, this is by far the most boreal of the genera, all the others being rather strictly tropical (except Synapsis) . In the Western Hemisphere (map, fig. 1) Copris is represented in the United States east of the 100th meridian, in all of Mexico and U. S. territory immediately bordering on Mexico (except Cali- fornia), and in all of Central America to Panama. In South America it is represented by a single species from North and Cen- tral America occurring in the mountains of Colombia and Ecua- dor. It is absent from the Antilles and the Galapagos Islands. One species has been introduced by man into Hawaii. The American forms are distributed by countries as follows : Canada: /. fricator (F.). United States: arizonensis Sch., f. fricator (F.), /. cartwrighti Rob., gopheri Hubb., kowdeni M. and H., incertus Say, inemargina- tus Blatch., 1. lecontei n. sp., minutus Dru., r. remotus Lee. Mexico : arizonensis Sch., armatus Har., boucardi Har., costari- censis dolichocerus n. subsp., halffteri Matt., incertus Say, k. klugi Har., k. sierrensis n. subsp., laeviceps Har., 1. lecontei n. sp., 1. isth- miensis n. subsp., lugubris Boh., megasoma M. and TI., mexicanus M. and H., moechus Lee., rebouchei Har., r. remotus Lee. r. dicyrtus M. and II., sallei Har. Guatemala: aspericollis Gill., boucardi Har., costaricensis do- lichocerus n. subsp., laeviceps Har., lugubris Boh. Belize : laeviceps Har., lugubris Boh. El Salvador: boucardi Har., lugubris Boh. Honduras : laeviceps Har., lugubris Boh. Nicaragua: lugubris Boh. Costa Rica: c. costaricensis Gahan, incertus Say, laeviceps Har., lugubris Boh., subpunctatus Gill. Panama : c. costaricensis Gahan, lugubris Boh., subpunctatus Gill. Colombia : incertus Say. Ecuador : incertus Say. 4 ENTOMOLOGICA AMERICANA Fig. 1. The apparent distribution, based on material in collec- tions, of the genus C opr is in the Western Hemisphere. 5 SMITHSONIAN Ann , _ MsmuTioN APR 27 ENTOMOLOGICA AMERICANA MATERIALS AND METHODS The present study was based on the collections of several institutions and individuals and was supplemented by field ex- cursions into certain critical areas from which adequate material was found to be lacking. The methods used were those standard to taxonomic procedure. An attempt was made to investigate cryptic characters for use in classification by examining one specimen of both sexes of each species in minute detail, dissecting it to its minimum component parts and examining all interior as well as exterior surfaces. In this manner the possible taxonomic significance of the mouthparts, antennae, wing venation, dorsal abdominal surface, metendosternite, male genitalia and female spermathecum was investigated. The results were disappointingly negative; although some characters, such as the male genital capsule, reflected species differences to a slight extent, these could be much more easily seen using ex- ternal features. Consequently, the use of cryptic characters was abandoned and the study was confined to those external ‘ 1 classical ’ 5 characters familiar to investigators for centuries. In the descriptions, characters cited in the group and complex descriptions are not repeated under the species. In the present classification the use of the male armament has been avoided completely in arriving at a natural system. However, the male armament, when developed, provides one of the easiest ways of identifying a species quickly; for this purpose facies illustrations have been provided on plates IV-VII and the first key to the species is based largely on male secondary sexual ornamentation. It is recommended that, if a series includes some major males, the first key be used for determination. However, all forms of both sexes should key through the second key. With regard to the geographical localities cited on labels, some judicious elimination has proved necessary. If a locality written on a label appeared to be unreliable for any reason, it was disregarded. A certain amount of difficulty was experienced in locating many Mexican and Central American localities on a map. It would greatly faciliate the work of future investigators if collectors in these areas would confine themselves to citing localities which are listed in the Index to the Map of Hispanic America, 1:1,000,000 (1943, 1944). In Central America, the Department in which a town is located is essential information. In the “material examined” section, localities for which the department or state was given but which could not be found on any map are preceded by a question mark. Additional pertinent 6 ENTOMOLOGICA AMERICANA information not provided on the label, snch as the altitude, was added to the citation in brackets. Information added to the Champion localities was obtained from Champion (1907). In the case of certain groups of insects, such as the present one, much useful biological information can be gleaned from the data on labels if the collector includes a word about how the insect was collected. The only collector in this group who has consistently done this is the late F. Nevermann of Costa Rica. All illustrations are by the author. The facies illustrations (plates IV— VII) showing the male armament were drawn by eye with the aid of proportional dividers. These illustrations are meant to show only the overall aspect of the forebody of a major male of each species and a general indication of the distribution of simple and complex punctures. Such details as the distribution of setae and the proportions of the mouthparts are only roughly indicated and not meant to be exact. The mouthparts and genitalia illustrated were mounted on slides and projected on paper with a standard projector. All other illustrations except the nest diagrams were done with the aid of an ocular grid and cross-lined paper. The nest diagrams were redrawn from field sketches and measurements. A total of 3,400 specimens was examined, including the holo- types of 14 names. Many of the Mexican species were described by the Baron de Harold. Harold’s types are presumed to be in the Paris Museum, but I was unable to obtain verification from that institution. Fortunately, Harold’s descriptions, although short, are excellent and little doubt remains in my mind as to which of his names belongs to which species. ACKNOWLEDGEMENTS I wish to thank the curators of the collections at the following institutions for the loan or examination of material : American Museum of Natural History (AMNH), British Museum (Natural History) (BM), California Academy of Sciences (CAS), Carnegie Museum (CM), Cornell University (CU), Defensa Agricola (Mexico) (DA), Department of Agriculture (Canada) (CNC), Musee Royal d’Histoire Naturelle (Brussels) (MRHN), Museum of Comparative Zoology (MCZ), Naturhistoriska Ricksmuseum (Stockholm) (NR), Instituto Rockefeller (Mexico) (IR), Phila- delphia Academy of Sciences (PAS), United States National Museum (USNM), University of Kansas (UKs), University of Michigan (UMich). ENTOMOLOGICA AMERICANA In addition, the following individuals generously made avail- able material from their private collections : 0. L. Cartwright (Washington) (OLC), G. Halffter (Mexico) (GH), H. F. Howden (Ottawa) (HH). I am particularly indebted to the following scarab specialists for invaluable advice and encouragement: V. Balthasar (Prague), 0. L. Cartwright (Washington), G. Halffter (Mexico), H. F. Howden (Ottawa), A. Martinez (Buenos Aires), F. S. Pereira (Sao Paulo). Mr. R. A. Crowson of Glasgow kindly made comparisons of specimens with a Fabrician type in the Hunterian Collection. Drs. S. E. Neff and 0. S. Flint, formerly at Cornell University, provided much help in the field work in Central America and North Carolina respectively. MORPHOLOGY The morphology of the adult Scarabaeinae has been the subject of some investigations. Hardenberg (1907) presented an excellent discussion of the mouthparts of Pinotus carolinus (L.), Co'pris fricator (F.), Canthon pilularius (L.), and several other Coprini, plus many other scarabs. Mohr (1930) compared the external morphology of Canthon pilularius (L.) with that of an aphodiine and a geotrnpine. Plalffter (1952) discussed aspects of the ex- ternal morphology of Phanaeus quadridens Say. Pereira and Martinez (1956) presented illustrations of the mouthparts and genitalia of many Canthonini and Matthews and Halffter (1959) discussed the external morphology of a species of Copris. For purposes of comparison, frequent mention is made below of some of these papers and of one on a melolonthine ( Amphimallon majalis Razoumowski) by Butt (1944) because of the detail in which the morphology of this species is there treated. The following discussion is confined to those features of the external morphology of adult Copris which are of taxonomic significance, i.e. which may serve to distinguish the genus from others closely related or which vary among the species. Microsculpture The interpunctural surfaces are generally quite smooth in the American species except in fricator and howdeni, where the surfaces show a very fine shagreening. 8 ENTOMOLOGICA AMERICANA The punctures are of two basic types which in turn vary importantly among the species groups and complexes : 1. Simple punctures. This term refers to simple surface de- pressions which have no raised areas within them and which usually have indistinct edges. They are found on the dorsal head surface and on the convex portions of the pronotum and elytra in all species. 2. Complex punctures. These are of varying type but always show a raised area within them and sharp edges. The raised area may or may not bear a seta ; in the former case the puncture is said to be setigerous or umbilical and setigerous ; in the latter it is said to be simply umbilical or, if very large, annular. The following areas show round setigerous punctures with the seta more or less erect (fig. 23) : along the posterior margin of the inner, smooth area of the proepimeron, on the sternellum, the mesosternum, the mesepimeron, occasionally the median lobe of the metasternum and the median coxae, and the abdominal sterna. All other areas with setigerous punctures show a reniform type (fig. 21), which is a setigerous puncture one side of which has been pushed in with the result that the seta appears to issue from the side of the puncture and is recumbent, being directed away from the pushed-in edge. The following areas normally show round or oval umbilical punctures which are not setigerous (figs. 20, 22) : the dorsal surface of the genae, the frontal area behind the horn, all depressions on the pronotum and the pygidial surface. Finally, some species complexes show a remarkable modification of the inside surface of the punctures, which is distinctly granu- late (when seen at above 90 x magnification) (figs. 20, 21). This type of surface is seen inside all complex punctures, of whatever type, which are large enough to show the inner surface, on both the dorsal and ventral body surfaces. Punctures of this type are said to be granular and occur throughout the minutus species group, recurring in the fricator group only to a lesser extent in the rebouchei complex. Head The cephalic taxonomic characters which are not self-explana- tory are the following : Posterior oblique carina.- — This is a carina on the dorsal surface which runs from just behind the eye obliquely inward towards the base of the horn. When present, it is usually very sharp immediately near the eye and then stops abruptly, being continued 9 ENTOMOLOGICA AMERICANA inward and forward by a raised ridge of varying sharpness (fig. 25, poe). Transverse occipital groove. — Running very near the edge of the dorsal portion of the occipital margin is a groove bearing short, closely set setae (figs. 24, 25, tog). This groove may be complete across the entire occipital margin, or it may be divided into three approximately equal parts with the lateral ones displaced slightly forward (fig. 25). Occasionally the median section is missing. Transverse occipital carina. — This is a carina of varying sharp- ness which runs in front of and closely parallels the transverse occipital groove (fig. 24, toe). It is found only in the incertus complex. Immediately behind the front margin of the head dorsally is a row of seta tufts which are most conspicuous medially but which extend for varying distances laterally and posteriorly, sometimes extending onto the genae. The extent of this row of tufts ap- parently varies significantly among the species, but this character was not used in the classification because of the difficulty of seeing the setae in worn specimens. The mouth cavity is rectangular in shape and is margined with a very broad, smooth margin which is anteriorly drawn out into a point. The shape of the mouth cavity differs somewhat in different coprine genera. Mouthparts The mouthparts of Copris and all other Scarabaeinae are quite remarkable for their modification of a basically primitive type towards very highly sensory and membranous appendages (plate II). A good description of the mouthparts of Copris fricator (F.) (= C. anaglypticus Say) is given by Hardenberg (1907). Since the mouthparts do not appear to provide useful taxonomic characters at the species level in this genus, they will not be further discussed here. Prothorax The prothoracic taxonomic characters which are not self-ex- planatory are the following: Lateral pronotal carina. — This is a carina which runs from an anterior point on the pronotal lateral margin upwards and back- wards to pass just under the lateral fossa, after which it dis- appears. It is absent in the minutus group. 10 ENTOMOLOGICA AMERICANA Prosternal-proepisternal suture (fig. 26. pp). — This is the suture between the anterior portion of the prosternum and the pleuron. In the incertus complex it appears carinate because of a deep depression of the pleural surface immediately next to it. Pleural elements. — The interpretation of the lower surface of the prothorax adopted here differs radically from that of many coleopterists (who call the entire lower surface the sternum) and that of Butt for Amphimallon, but agrees with that of Mohr and Ilalffter. Butt considers the lateral portions of the lower surface as parts of the pronotum, which is therefore believed to join directly with the sternum with a resultant complete inflection of the pleural elements, which Butt considers to be represented only by a tiny sclerite fused to the dorsolateral wall of the prono- tum on the inside. In my opinion, it is more logical to assume merely that the sutures delimiting the pleural sclerites, seen in the primitive Coleoptera, have become completely fused and effaced in order to reinforce the prothorax. This would mean that most of the under surface lateral to the sternum in scarabs is made up of pleural elements fused together. On this pleural area in Copris there is seen a distinct oblique carina running from the coxal cavity outward (fig. 26, pc). This carina is absent in many coprines and other scarabs and may be assumed to be a secondary modification evolved to further rein- force the prothorax. For the purposes of convenience, the area anterior to this carina is called the proepisternum, and that pos- terior to it is called the proepimeron. However, it should be understood that these areas do not in all probability correspond to the true morphological sclerites called by these names. The longitudinal proepimeral carina. — The proepimeron in Copris and a few closely allied genera bears another characteristic carina which arises perpendicularly from the posterior proepimeral margin and quickly fades out (fig. 26, lpc). This carina serves to partly delimit a densely, setigerously punctate outer area of the proepimeron from an almost smooth inner one (except in C. minutus). Pterothorax Very few taxonomic characters have been found in the ptero- thorax and all are self-explanatory. The proportions and shapes of the sclerites do not differ among the American species of Copris . 11 ENTOMOLOGICA AMERICANA Elytra There are ten distinct elytral striae in the genus Copris, of which the ninth is practically always present only for its pos- terior half and lies very close to the tenth. The eighth stria is of great taxonomic importance and may be present merely as a short segment near the middle of the sides, in which case it is said to be obsolescent, as a line more or less interrupted posteriorly at the hind angles, in which case it is said to be incomplete, or as a complete, uninterrupted line. Legs The general shape and proportion of the legs in Copris may be seen in fig. 26. There is one strong spur terminally on the anterior tibia, called the forespur, the shape of which is of great taxonomic significance. The fore coxae are elongated cylinders very deeply sunk into the prothorax (fig. 27). They are rigidly pivoted at the two opposite ends of this cylinder such that the only possible move- ment is a rotation. The outer pivot consists of a knob formed by an invagination of the pronotal surface, called the lateral fossa of the pronotum. The median coxae are oriented parallel to the body axis and widely separated. Their ventral (visible) surfaces are longi- tudinally obtusely carinate. The area exterior to this carina is called the outer face of the median coxa. The arrangement of setae on the middle and hind tibiae was found to be of taxonomic significance with regard to two sets of setae, as follows : Ventral seta tufts, also called distal seta tufts (fig. 49, st). — These terms refer to a row of tufts on the lower surface of the tibiae. The tibia is seen to be quadrate in cross section with a row of teeth and setae running down each ridge forming the corners. The ventral seta tufts are situated along the middle of the ventral face between the usual ridge setae and are usually confined to the distal end. Supplementary setae (fig. 50, ss). — These are confined to the incertus complex and consist of an oblique row of very few, in- distinct setae near the distal end of the dorsal tibial surface. The tarsi are unmodified and always present. Each segment is somewhat expanded distally and gradually diminishes in size apically. There is some variation in the comparative width of 12 ENTOMOLOGICA AMERICANA the tarsal segments in different species, but this character was not used in the taxonomy. The tarsal claws are small, parallel, and equally developed, with a small plate-like empodium at their base. Abdomen The only taxonomic characters on the abdomen involve the pygidium, which may be incompletely margined ventrally. The visible sternites are six in number and do not merge together along the ventral mid-line. The tergites of segments VII and VIII are of particular interest because they make up the ventral portion of the stridula- tory apparatus. The dorsal portion of tergite VIII (the pro- pygidium) is heavily sclerotized and bears a deep longitudinal groove medially. This groove firmly holds the down-turned median edges of the elytra in repose. The actual stridulatory surface is not on this segment, however, but on the preceding one, where it consists of a finely sclerotized median area bearing some extremely fine transverse ridges. These ridges rub against some similar ridges or teeth on the elytral ribs when the abdomen is moved against the inside elytral surface, the groove on tergite VIII serving to keep the two parts in line. Both sexes of every species of Copris I have examined show this apparatus and all those which I have collected in the field could be induced to squeak. For a discussion of stridulation in the lamellicornia the reader is referred to Arrow (1904). Genitalia On the whole, the male genitalia in this genus are not of any use in determining the species, at least as far as their gross aspect is concerned. The possibility that the internal sac may bear taxo- nomic characters was not examined. SEXUAL DIMORPHISM The phenomenon of sexual dimorphism in the lamellicorn beetles has been the subject of much speculation but, unfortunately, little organized investigation. Darwin (1871) used the scarabs as examples for his celebrated theory of sexual selection. Since then the diversity and possible uses of the horns or enlarged mandibles in the lamellicorns have been discussed in numerous works. It is 13 ENTOMOLOGICA AMERICANA not the purpose of this discussion to review this voluminous litera- ture; for a recent survey the reader is referred to Arrow (1951). For some of the theoretical implications of horn allometry in the scarabs see Huxley (1932) and Paulian (1935). An excellent recent quantitative survey of horn allometry in a single dynastine species is presented by Bowden (1959). However, no one has examined the nature of the variation in horn size among related species of a single genus. It is the purpose of this survey to examine intra- and interspecific variation in horn size and general qualitative differences in armament among all the American species of Copris. The Sexual Armament in Copris Nearly all the males of the American species display armament of a single type : there is one horn arising from the middle of the frontoclypeal suture on the head and the pronotum bears four forwardly directed prominences in a transverse row (plates IV- VII). The median two of these prominences are variable in shape (blunt, acute, or truncated) and sometimes merge into a single process. The lateral pronotal prominences are always strongly compressed and acute when developed. The females typically bear a transverse crest on the fronto- clypeal suture of the head. This crest is low and truncate apically, transversely oval and slightly excavate in dorsal view, and its sides converge apically, are parallel, or diverge, depending on the species or degree of development. The pronotum of the female does not bear any corniform prominences but does bear a median transverse carina (sometimes interrupted medially) and a small tumosity to either side of this carina. In the species descriptions female armament of this type is simply described as “ normal for the genus. ’ ’ Following the established custom among scarabaeidists I have called specimens (male or female) showing very little or no development in sexual armament “minor” individuals, and those showing strong development “major”. It should on no account be inferred that these are clear-cut categories, but rather that they represent the two extremes of a spectrum of variation. Alternately I have called these specimens poorly developed and well developed, respectively. The following exceptions to the above descriptions of armament are seen in the American species : 14 ENTOMOLOGICA AMERICANA In the minutus group, the female head horn (or crest) is narrow, high, and rather corniform but still apically truncate in the incertus complex, completely corniform (apically acute) in the minutus complex. In the latter complex, in fact, it is very difficult to distinguish between the sexes, since the pronotal arma- ment is often scarcely developed. The incertus complex is further distinguished by the presence of a corniform tubercle on the frons of the male behind the head horn. In the species halffteri Matthews an extraordinary reversal has taken place : the male has acquired female armament. This phe- nomenon must not be confused with the similarity to females shown by very minor males of many species. In halffteri the males I have seen are well developed and show strong armament, but this is always of an exaggerated female type (fig. 77) which few females of any species ever achieve, because they are seldom so developed. This species is apparently very localized and shows strong affinities with a common and widespread species ( C . rebouchei Harold), from which it evidently evolved. It is interesting that this form, which is most certainly a separate species, is completely sympatric with its parent species, both having been found in the same pile of cow dung. Intraspecific Variation in Horn Height The measurements used in the following analysis represent the height of the male head horn from the edge of the clypeus to the tip of the horn in direct front view. This is compared with the maximum length of the hind femur in ventral view. The measure- ments were carried out with an ocular micrometer at 9 x magnifi- cation, the units used being one micrometer unit or .097 mm in this case. Since only ratios are dealt with, the measurements were not converted to millimeters. For calculating curvilinear regres- sion lines, a constant factor was subtracted from each measurement of horn height to insure that the horn itself, and not also the height of the frons, was being taken into consideration. This factor was taken as the height from the clypeal edge to the tip of the frontal protuberance in the least developed specimens (those with no horn as such) of the species involved. The mathematical interpretations of the curves obtained were taken to be either the “simple heterogony” formula of Huxley (1932) (y = bxk, where y represents the magnitude of the differen- tially growing organ — in this case, the horn, x represents the magni- tude of the animal or some reference structure — the femur, and b 15 Table I. Regression of Male Cephalic Horn Height (y) on Length of Hind Femur (x) in 14 Species of Copris. 1 unit = .097 mm ENTOMOLOGICA AMERICAN A «H © rt Zj U 2 ® rcJ rQ *jH O > 2 s as £.2 S coiococorHcMoorHLorHcooo OCO CM r- I rH OXMHffi CM O b- OS 00 rH CO l-H r— I r— I Ncoo^cooaooN OH^Hoaoooooi lOrHOOOOOOSCOOSOO rH 05 lO O rH H H CO CO OS CM OS rH O OS O OS 00 os SO O pi I ^ IK (McoMt>THO]NOTj7 tL o' o' of oT o' io c\T rH o' Tta0©©^(»©OI>HWHHN05C0WHQ0 o' rH CM* tg CM cm" rH CO O* 00 OO OS o" rH* CM* rH OS* o’ CM COrHrHCOrHrHrHrHrHrHrHrHLOLOLOLOLOCOCO bC c$ be o K be o II be o b- CM O OS OS rH CM* CO ^ r- 1 1— I H LO Os LO 00 I | CM CO LO CO CO K CM rH 00 OS OS CM ^ o io o oo ^ LO rH CO l> be bCi-H CO cs co os' t>! 00 o o o CM I I o b- X K X X X rH io co OS CO LO OS ^ 0- co os os as os H Tfl OS rH CO CO II II II II II II II K K K !b h K K be be w o o X X GO co O co co’ LO II II r^s CM ^ OS 00 rH CO CM LO CM CM | O O rH I j*t GO LO LO ■> GO h cm "r be l l?Si Hr1 CO OS rH CO . °°. LO o' ^ rH LO rH CM CM II II II II II r-5 r“% r^ be bcw o o I I X ><1 oo o 00 CM co in’ II II os OS LO CO LO* rH £ ?3 Si o* 16 Average of a except those Average : Total in parentheses: 4.76 .825 656 ENTOMOLOGICA AMERICANA and k are constants) or a simple linear relationship (y = ax + b') such as that found by Bowden (1959), whichever seemed to fit best. These different interpretations are of no real significance, in my opinion, the linear curve probably being merely the top part of a power function curve. In the case of C. moechus, it seems that both a power function and a linear curve fit the data (fig. 4 and table I), with the break occurring near the middle. It may be objected that the material measured here was taken from museum collections and therefore not collected under proper statistical sampling procedure. To this it should be pointed out that we are concerned here with the positions of curves only, not with their slope or the positions of their origins of deviation (the common mean of the two variables) . It should, theoretically, make no difference where along a common curve a sample is taken; the subsequent plotting of these samples should reveal the position of the curve. One of the main purposes of this study is to determine whether significant differences can be found in horn development patterns within a single species and to analyze the nature of these differences if they occur. When plotted on a graph, the data points represent- ing horn height to femoral length ratios for an intraspecific group- ing, such as a population or geographical race, could differ from those representing another grouping, or the rest of the species, in two quite different respects : 1 ) they could fall along only a section of a common species curve, that is to say, a race may fall near the bottom of the curve, another near the top, etc. or 2) much more significantly, one or more populations may fall along a recognizably different curve, thus showing a different allometric relationship with at least a different value of 1). Individual populations could not be examined in this respect since they were almost never represented in sufficient numbers. With regard to geographical groupings, however, significant dif- ferences were found within five of the 14 species examined. In minutus (Drury) (fig. 2) specimens from a relatively small area of the range, represented by Mobile and Montgomery, Ala. and Clarksville, Fla. (in the Florida panhandle), are seen to fall almost entirely along the extreme upper portion of the curve (open circles) with little overlap with the rest of the species (dots). Mysteriously, two specimens from this area fall near the bottom of the species curve. It will be noted, however, that all specimens from this area give every appearance of falling along the common species curve. Visually these specimens are conspicuous for their great size and horn development. 17 30 28 26 24 22 20 18 16 (4 12 10 8 6 ] fem< this ENTOMOLOGICA AMERICANA minutus o specimens from length of hind femur I unit =.097 mm * ■ - L. i l l I . i I 22 24 26 28 30 32 34 36 38 ig. 2. C. minutus (Drury). Plot of horn height (y) against *al length (x) ; k = 9.40 for random 25% sample (N = 53). For nd the following figures refer to text and table I for explana- of the symbols used. 18 ENTOMOLOGICA AMERICANA In fricator (F.), on the other hand (fig. 3), specimens from a wide geographical area in the southwestern portion of the range (open circles) are all grouped in the lower part of the common curve. Visually these specimens also stand out, but in this case for their hornless condition and small size. Here, however, there is a suggestion that these specimens fall along a slightly different curve situated higher than the common species curve. fricator o specimens from Ks., Fig. 3. C. fricator (F.), k = 15.09 for x = 42 to 50. Interrupted lines estimated. 19 ENTOMOLOGICA AMERICANA The previous two examples of geographical variation come under category (1) above. The geographical race falls along a special part of the common species curve, with little or no evidence for any different allometric relationship. The most interesting and significant type of variation is the one in which a geographical grouping falls along a totally cliferent curve from the one which characterizes the rest of the species. This is unmistakably seen in three species of the genus (figs. 6-8). In the first ( klugi Harold), when the data from specimens origi- nating in the Sierra Madre Occidental are plotted, they are seen to fall along a line which is significantly to the right of those from the rest of the species, orignating in the Transverse Volcanic Range. The degree of significant separation may be judged by noting the extent of overlap of the 95% confidence intervals shown. Visually this difference is expressed by the northern specimens possessing consistently shorter horns for their size, in comparison with the southern form, but this difference is so slight as to be all but undetectable to the eye. This is in sharp contrast to the visually conspicuous but much less important difference in cate- gory (1) above. This shift in the position of the relationship curve in different geographical groupings is much more strongly accentuated in two additional species : in costaricensis Gahan, comparing specimens from Chiapas with those from Costa Rica (fig. 7), and in lecontei n. sp., comparing specimens from north and south of Cabo Corrientes on the west coast of Mexico (fig. 8). For each of these three species there are therefore two separate relationship curves (table I). In the last three examples the differences in allometric rela- tionship are reflected in morphological differences only in the first species (klugi). Here the northern specimens consistently show a faint sclerotized diagonal band on the male parameres; this band is absent in the southern form. In the other two species I could detect practically no morphogical differences between the geo- graphical groupings thus separated by their horn relationships. Hence, this procedure gives us a taxonomic tool of some sensitivity in the horned scarabs. It should further be noted that even if different geographical groups or different species fall along dif- ferent curves the slopes of these curves are approximately the same, allowing for sampling error and bearing in mind the very small samples in some cases. 20 70 65 60 55 50 45 40 35 30 25 20 15 10 height of cephalic horn I unit =. 097mm VOLUME XLI moec hus length of hind femur I unit = .097 mm 1 1 i i 1 1 1 1 1 44 46 48 50 52 54 56 58 60 Fig. 4. C. moechus Lee., k = 20.13 for x = 44 to 52 ; a = 2.63 for = 52 to 58. 21 ENTOMOLOGICA AMERICANA • arizonensis Fig*. 5. C. arizonensis Schaeff, C. armatus Har., and C. sub- punctatus Gillet, a = 3.88, 6.29, and 4.40 respectively. 22 VOLUME XLI Fig. 6. C. klugi Har., & = 4.14 for Transverse Volcanic Range, 5.86 for Sierra Madre Occidental. Interrupted lines indicate 95% confidence intervals for regression lines. 23 ENTOMOLOGICA AMERICANA costaricensis Fig. 7. C. costaricensis Gahan, a = 4.36 for Costa Rica, 9.99 for Chiapas. Interrupted lines indicate 95% confidence intervals for regression lines. 24 VOLUME XLI Fig. 8. C. lecontei n. sp., a = 3.99 for Arizona to Nayarit, 3.08 for Oaxaca and Colima. Interrupted lines indicate 95% confidence intervals for regression lines. 25 ENTOMOLOGICA AMERICANA Interspecific Variation in Horn Height It is not our purpose here to discuss the interesting theoretical implications of the species curve distributions. It should merely be noted in passing that the “rule of Lameere and Smith” (Huxley, 1932, pp. 212-216) is not at all adhered to in Copris (contrary to what Paulian [1935] states) but that each species falls along a curve (or two curves) of its own and that these curves are re- Pig. 9. Plot of horn height against femoral length ; regression lines for all species measured. Interrupted lines are of relatively low reliability. Numbers refer to species as listed in table I. markably parallel (fig. 9). This is expressed mathematically in the relatively constant value of a (averaging 4.76) in the linear formula y = ax + b' (table I). The points of origin of deviations are nearly isometric (fig. 10). This shows that there is a constant average pro- portion of horn-to-body size in all the species regardless of size and a more or less constant degree of allometry. It seems probable that the allometric nature of the horn size relationship is of selective advantage to the species, perhaps to maintain some sort of domi- nance hierarchy in random assemblages at food sources for the purposes of pairing off. 26 VOLUME XLI There are no detectable consistent qualitative differences in horn shape or design between species to suggest in any way that the horn could have any species-recognition value. There are differences in horn design and these are very useful to the taxon- omist, but these differences do not correlate with relationship or geographical distribution in any way (e.g., closely related sym- patric species do not necessarily differ more strongly in horn design). 30 39 40 43 30 33 60 65 70 Fig. 10. Regression of mean height of head horn on mean length of hind femur for all species. Solid line represents regres- sion of y on x; interrupted line represents the line of constant proportionality (isometry). Further explanation in text. Num- bers refer to species as listed in table I. BIOLOGY Notes on the nesting habits of five American species have been published. These will be briefly summarized below together with notes on the results of my own field observations on four species. The available data indicate that the majority of the species of the genus feed on mammal dung, the only known exception being that of C. gopheri Hubbard, which feeds on the dung of the 27 ENTOMOLOGICA AMERICANA Florida Gopher Tortoise. Aside from this, there is no evidence to indicate that any preference is shown for the dung of any particular species or type of feeder. There are scattered records of individuals occurring at carrion (these are mentioned in the sys- tematic section under the particular species involved), but it must be borne in mind that these could be attracted to the gut contents of the carcass. There are a very few records of individuals coming to traps baited with fermenting malt, mushrooms, decaying meat, and fruit. Insofar as I have been able to determine, the nidification be- havior of the American species differs only in detail from that of Copris hispanus (L.) in Europe, as discussed by Fabre (1918), Siyazov (1914), and Lengerken (1954). It appears, therefore, that biological differences between species generally do not reside in the nature of the food consumed or in the basic aspects of nidification. We may therefore expect that such differences are to be found in ecological tolerances to such factors as humidity, temperature, and so forth, so that the genus may be able to exploit the available dung food in all possible en- vironments. One example of this may be provided by the habitat preferences shown by the two closely related species incertus Say and lugubris Boheman. The former is typical of the humid tropical forest fauna of the more elevated portions of the Veracruz region, exemplified by Jalapa and Fortin de las Flores, whereas the latter, in this region and in Oaxaca, is restricted to the open areas — the clearings in the forest and the sandy areas immediately along the sea coast (Gonzalo Halffter, personal communication). There follow brief accounts of the nidification of seven American species the habits of which are at least partly known. Copris fricator (Fabricius) From my own observations in the Blue Ridge Mountains in May, 1959, the following outline of the life history of this species emerges. Nidification is preceded by a period of adult feeding in the spring. Feeding always involves the digging of a small, shallow individual burrow about 4 cm. long beneath or beside a pile of cow dung. The burrow is filled with dung before feeding, the beetle grabbing small “armfulls” of dung and backing into the hole. Each individual of both sexes digs its own burrow in- dependently. Nidification begins towards the end of May in North Carolina and usually involves a pair of beetles of each sex acting in co- 28 VOLUME XLI operation, although the female may begin the nest alone. Of 13 nests I uncovered in various stages of construction and provisioning, seven contained a male as well as the female. Of six later nests uncovered, containing completed brood ovoids, a male was present in only one, showing that the male usually departs after construction and provisioning is completed. The completed nest is a large oval underground chamber located under the dung pile and with the measurements indicated in table II. The floor of the chamber is 5-12 cm. below the ground surface and is connected to the surface by a short passageway just wide enough to admit one beetle ; this passageway is left open to the outside (fig. 82). Initially the nest is nearly filled by one large “cake” of dung which is “tended” by the female or pair wandering repeatedly over its surface. After an unknown period, the female begins cutting and shaping the brood ovoids from the dung cake (figs. 83-86). These measure about 2.5 cm. in diameter with one slightly longer vertical axis at the top of which is located a small spherical cavity containing one egg. The female evidently lays the egg and completes the ovoid before starting on the next one. The total number of ovoids seen in completed field nests varies from three to five. The larva on hatching begins to consume the inside of the ovoid, storing excrement in the characteristic coprine “hump” (an outpouching of the mid-gut). Emergence of adults in captivity from these ovoids completed in late May took place between 14 and 25 July. The continuous presence of the female in the nest was proved necessary to maintain the smooth, un- blemished contours of the outside of the ovoids. Ovoids kept apart from the female soon became heavily overgrown with fungi and molds. Lindquist (1933) indicates that the beetles overwinter singly as adults in very deep vertical burrows devoid of dung. Ritcher (1945) described the third-stage larva of this species (under the name tullius Olmer). Copris remotus Leconte Lindquist (1935) gave an excellent account of the nesting habits of this species near Uvalde, Texas. Some of his numerical data are summarized in table II. The points in which the nidification of this species appears to differ from that of C. fricator just discussed are as follows. The female apparently digs and provisions the nest alone ; provisioning, with cow dung, takes one or two days. Each brood ovoid takes two 29 ENTOMOLOGICA AMERICANA be be *g a r2 o a O *a o =4-1 o .2 bJDr^ © -2 *o '•§ 2 a 03 'co o 03 3 a o > b-p O M 02 a «H S o £ e=* 03 O <£> o ; a >t3 03 O .O 5'§:s :so 03 03 o >.s +J O 03 SfS.g g.ll > ® o a. © O 03 > 02 § O O O ^ ^ ^ O DO — « n3 03 03 a bn a be a s H-S be ° g a «-H o CO x TjH CO* X to o -+J a 0 o CM CO CO X co co X ^* to o 0 IO TjH X T* X tO tO +1 o t- CO CO CO cm X CM CO I tr- ee CO ©- * Q°. 05 OO* CM X to cm' ©=• CO 03 03 £ to CO e=>. tO . ©q* g I a 00 -r-1 es* r— I CM J2; e». tq CO to X X to t— t- X X 00 05 oo CO CM X X to 05 1—1 l— co t- co X to X t- to’ to CM* CO X X to t— ©=>• e»* t- X X CO to CO co X co Tji X co I- e=. —i . nd a •m *02 03 O r 1 u J > « 03 » v 03 03 •i 30 VOLUME XLI to three days to construct and there may be up to eight ovoids per nest. The female plugs the ingress passageway with soil. Under laboratory conditions embryonic development takes 5-10 days, the first instar 2-4 days, the second 2-5 days, the third 13-17 days, the pupa 12-37 days, for a total of 38-69 (average 49) days to complete development from hatching to adulthood. Eggs are obtained from 20 March to 10 October. A reared female lived 644 days in captivity. Several females laid eggs over two seasons. Lindquist was able to get up to 12 nests from one female, with a total of 41 eggs. The normal appears to be 3-4 broods annually. Copris aspericollis Gillet I uncovered three burrows of this species under cow dung near Guatemala City on 22 July 1958. Two consisted merely of short burrows containing one female each and no dung. The third consisted of a chamber containing a dung cake and a male and female pair. No ingress tunnel could be detected, suggesting that it had been blocked with soil. Evidently, nidification in this species involves the participation of both sexes. Copris minutus (Drury) The only available data on this common species is provided by Ritcher (1945), who found larvae in balls of dung associated with one or two adults in a brood chamber, the number of balls being two or three. The nests are dug several inches deep in the soil under cow dung. Measurements are given in table II. The adults are usually found in the chambers with the ovoids even after the larvae have pupated. Third-stage larvae were found in April and June, pupae in July. Ritcher described the full-grown larva. I found one nest of this species in the Blue Ridge Mountains of Virginia on 4 July, 1960, containing four brood ovoids and one female beetle. Copris gopheri Hubbard Hubbard’s (1894) account is still the only one we have on this interesting species. He found nests in large numbers four or five inches below the floor of the nest chamber of the Gopher Tortoise near Cresent City, Florida. The brood ovoids are apparently made of tortoise dung. An important difference and apparently 31 ENTOMOLOGICA AMERICANA unique feature of this species is that the larva, when pupating, constructs a cocoon of excrement around itself. This is apparently necessary because of the friable nature of the tortoise dung. The study of the biology of this species is made extremely difficult by the enormous depth of the tortoise burrows. Copris lugubris Boheman I was able to uncover six burrows of this species in Nicaragua on 27 and 28 July, 1958. The important features of the nests are diagrammed in figs. 87 and 88 and measurements are given in table II. Only the preliminary, or dung-cake, stage was seen. im- portant points are that the ingress tunnel is blocked with loose soil, there is usually a cavity under the dung cake (a unique feature), and the male always accompanies the female in this stage of nest construction (digging and provisioning). Copris incertus Say An excellent account of this species in New Zealand, where it was introduced, is given by Thomas (1960). His account is similar to that given above for C. fricator, but the following points are worth noting. On one occasion, three days after dung was exposed to beetles six burrows were dug up, four containing a female each and two containing a male and female pair. The dung cake remains in- tact for several weeks in the field. The egg cavity is made and the egg laid directly in the dung cake before the brood ovoid is cut and shaped around it. The particular process of oviposition has not been observed in the other species. Two to seven ovoids are made per nest. Development from egg to adult takes 57-70 (average 62) days in captivity. The larva and pupa are figured. It is to be noted that there is no cavity under the dung mass as is seen in the very closely related C. lugubris, but that the spaces are over and around the mass, as in the other species seen. Additional biological data taken from the labels attached to the collected material used in this revision are presented in the systematic section under the particular species involved. 32 VOLUME XLI TAXONOMY The genus C opr is Muller is completely isolated taxonomically from any other New World genus of Coprini and can easily be distinguished by the characters mentioned in the diagnosis. In the Old World, however, it shows close affinities with a number of genera, among which we may mention particularly Pseudo - pedaria Felsche (Africa), Coptodactyla Burmeister and Arrowia- nella Paulian (Australia). The customary grouping of coprine genera into “ subtribes” (ending in the termination -ides) after Gillet (1911) is in my opinion so artificial that it should be aban- doned. The genus C opr is has been recently divided into four subgenera (Balthasar, 1958), all the American species falling into the sub- genus Copris s. str. A study of the American species, however, reveals a sharp division into two groups which would merit sub- generic distinction if the genus were purely American. As it is, it is more probable that this division represents a double invasion of North America by two ancestral Asian species, since it is not reflected in the Old World fauna. It is impossible, therefore, to erect formal categories for these two subdivisions ; they have been treated as “groups” in the present work, each being named after the earliest described included species. Each of these groups are further divisible into subgroups, here called “complexes”, of species which fall naturally together. There are seven such complexes, each containing a few species, some of which are extremely closely related and most of which are distributed in geographical ‘ ‘ chains ’ ’ (allopatrically) . Figure 11 summarizes the relationships of the species as in- terpreted here. In the fricator group, the armatus complex is considered the most primitive as it contains large, morphologically unmodified species (with the full complement of elytral striae, simple anterolateral angles, typical puncturation, etc.). It will be noted that the geographical distribution of the groups suggests that the center of evolution for the genus in the New World is the northern Mexican highland region and that species become more evolved as they radiate out from this center. There is a further suggestion, derived from the close relationships between the species within the two groups, that the genus is at present rather actively speciating in the New World and may there- fore be of rather recent immigration, probably from Asia via the Bering bridge. Certain Chinese species I have seen (undetermined) bear a striking similarity to incertus Say. 33 Northern Mexico ENTOMOEOGICA AMERICANA Fig. 11. Proposed relationships of the American species of C opr is. 34 VOLUME XLI The fossil species Copris pristinus Pierce (1946), of the Upper Pleistocene (La Brea tar pits, California), is definitely a Copris and appears to be a true extinct species very close to C. armatus Harold but much larger, being in a size class with the giant C. megasoma Matthews and Halffter, which it also closely resembles. The latter two species are from central Mexico. I cannot place C. pristinus with greater accuracy at the moment, as I have not seen the original material. Palaeocopris labreae Pierce, from the same deposits, cannot be placed anywhere at present and appears to be a composite of two genera. Genus Copris Muller, 1764 Copris Muller, 1764, Fauna insectorum Fridrichsdalina, p. xi. Type, by subsequent designation, Scarabaeus lunaris Lin- naeus (Curtis, 1832, British Entomology, pi. 414). Diagnosis. Head semicircular in outline, very strongly flattened, and completely margined above, its dorsal surface punctate, never wrinkled. Thorax below with a distinct oblique propleural carina and a longitudinal proepimeral carina (fig. 26, lpc, pc), a short transverse mesosternum with a median impunctate area, an obtusely angular meso-metasternal suture, and a parallel-sided median metasternal lobe devoid of long setae. Elytra with ten striae, of which at least eight are complete (including the one along 'the lateral carina). Abdomen with six visible sternites, all of which are distinct along the mid-line, not being fused together or over- lapping. Fore tibia with four outer teeth or expansions, the proxi- mal one very small. Hind tibia with a very prominent transverse carina close to the middle on its outer face. Key to the Species of the Genus Copris in the Western Hemisphere Based on the Major Males* 1. Male armed like a female, with a well developed, typical female head horn which is transverse, quadrate, and apically ex- cavate (fig. 77) ; anterior margin of prosternum with a salient, bilobate median process (fig. 55). Guerrero half ter i Matthews. Male armed with a conical or slender horn ; anterior margin of prosternum with a median process varying in shape but never bilobate, or without any median process 2. * The following species, the males of which are hornless, are not included in this key: howdeni Matthews and Halffter, in- emarginatus Blatchley, and laeviceps Harold. C. sailed Harold, of which I have seen only females, is also not included. 35 ENTOMOLOGICA AMERICANA 2. Median pronotal prominences, when present, transversely truncated; lateral pronotal carina absent; lateral pronotal inner margin evenly curved; pygidial margin incomplete, its edge effaced vent rally 3. Median pronotal prominences rounded or acute ; lateral pro- notal carina present; lateral pronotal margin sinuate or slightly angulate anteriorly 6. 3. Head with a horn-like tubercle behind the horn; forespur expanded distally, truncate (fig. 33) ; a tuft of long setae present on hind edge of median and posterior trochanters 4. Head with only one horn-like prominence; forespnr acute or bluntly rounded; trochanters without setae 5. 4. Head horn arising approximately from middle of head surface ; tubercle behind it erect or sometimes bent (never inclined) forward (fig. 64) ; dorsal edges of median pronotal promi- nences, when seen from the front, sloping down laterally at an angle of about 45° from horizontal. S. Mexico and Central America lugubris Boheman. Head horn arising from before middle of head ; tubercle behind horn inclined forward (fig. 63) ; dorsal edges of median pronotal prominences, when seen from the front, almost horizontal. Tamaulipas to Ecuador, Hawaii, New Zealand incertus Say 5. Pronotum and elytral striae coarsely punctate ; clypeus con- trastingly more finely punctate than rest of head ; a segment of anterior portion of 9th elytral stria present and coarsely punctate ; proepimeron fully punctate. Eastern U. S. minutus (Drury) Entire body very feebly punctate ; elytral striae partially impunctate ; 9th elytral stria present only posteriorly ; proepimeron partly impunctate inwardly. Florida gopheri Hubbard 6. Anterolateral angles of pronotum with the point made salient by an inward curve or emargination of the margin im- mediately behind it; 8th elytral stria nearly always com- plete. Smaller beetles (10-18 mm in length) 7. Anterolateral angles of pronotum subquadrate, obtusely angled, or broadly rounded, occasionally acute in very highly developed specimens, never with a sharp inward curve of the margin immediately behind them. Larger beetles (14-29 mm in length) 13. 7. Pronotum uniformly very densely punctate, the punctures separated by a distance roughly equal to their diameter 36 VOLUME XLI (fig-. 66) ; elytral interstriae distinctly and profusely punctate ; head horn almost straight, slightly transversely dilated at apex in well developed individuals. Central and Northeast U. S., Southeast Canada f. fricator (Fabricius) Pronotum not uniformly very densely punctate, the punctures separated by a distance equal to far more than their diameter on at least part of pronotum; elytral interstriae very finely punctate (appearing impunctate to the unaided eye) or quite impunctate 8. 8. Median dorsal sulcus of pronotum and pronotal disc im- punctate; median pronotal prominences laminar, well sep- arated, with a deep depression between them, and parallel (fig. 76) ; pygidial margin incomplete, its inner border completely effaced ventrally (fig. 58) ; anterior prosternal margin with an acute, minute median tooth (fig. 54). Central Mexico rebouchei Harold. Median dorsal sulcus of pronotum punctate or, if impunctate, then anterior prosternal margin is without an acute median tooth and inner border of pygidial margin is completed ventrally by a close-set row of punctures (fig. 59) 9. 9. Pygidial margin incomplete, its inner border completely effaced ventrally ; median pronotal prominences closely approxi- mated, parallel in direction, in developed individuals tend- ing to become merged into a single bifurcate process re- ceiving the head horn (fig. 80); anterior margin' of pronotum forming a minute, downwardly directed median point. Chiapas and Central America costaricensis Gahan Pygidial margin nearly always complete ; median pronotal prominences never merging and otherwise not as above 10. 10. Male with well developed, divergent median pronotal promi- nences, but with either no trace of any lateral prominences (fig. 79), or these represented by tubercles. Tamaulipas remotus dicyrtus Matthews and Halffter. Male, if bearing well developed median pronotal prominences, also bears well developed, laminar lateral prominces 11. 11. Clypeal teeth obtuse, relatively approximated, the margin be- tween them excised at a distinct angle without any median notch ; hind angles of head quadrate ; median dorsal sulcus of pronotum not contrasting with rest of disc in punctura- tion ; disc usually moderately punctate (fig. 75). S. Arizona and W. Coast of Mexico to Tehuantepec lecontei n. sp. Clypeal teeth acute and remote, very small, the margin between them not appreciably excised, with or without a 37 ENTOMOLOGICA AMERICANA very shallow median notch ; hind angles of head acute ; median dorsal sulcus of pronotum coarsely umbilico- punctate in sharp contrast to the impnnctate or very finely punctate elevated portions of disc 12. 12. Median pronotal prominces remote and slightly divergent (fig. 78) ; forespur abruptly bent in very near apex, where it forms a dull point (fig. 43). S. Texas, N. Nuevo Leon r. remotus Leconte. Median pronotal prominences very approximated, their outer edges converging forward (fig. 67) ; forespur curved in- ward and tapering near apex to a sharp point (fig. 44). Miehoacan mexicanus Matthews and Halffter. 13. Clypeal emargination very shallow, wide, and arcuate, without a median notch ; forespur curved inward and tapering to a sharp point (fig. 39) ; median pronotal prominences closely approximated to form a single bifurcate process in well developed individuals (fig. 68) ; entire surface of pronotum regularly and usually densely punctate; 8th elytral stria complete. S. Arizona and New Mexico, W. Texas Chihuahua arizonensis Schaeffer. Clypeal emargination broadly triangular, with a median notch, or absent ; forespur rounded at the end or coming to a blunt point, curving downward a little but not inward appreciably ; 8th elytral stria incomplete, disintegrating posteriorly or largely effaced 14. 14. Median prominences of pronotum with the apices well sep- arated, their outside edges divergent in well developed in- dividuals 15. Median prominences of pronotum closely approximated, their outside edges evenly convergent anteriorly 18. 15. Anterior face of pronotum sparsely to moderately punctate ; clypeus devoid of teeth 16. Anterior face of pronotum coarsely granulate; clypeus ob- tusely bidentate 17. 16. Median pronotal prominences with the apices obliquely flat- tened, their lower surfaces flat or slightly concave ; clypeus entire, without any median notch (fig. 70). Miehoacan megasoma Matthews and Halffter. Median pronotal prominences conical, round in cross section; clypeus with a median notch (fig. 69). Central Mexico armatus Harold. 17. Elytral striae appearing distinctly punctate to the unaided eye; apices of median pronotal prominences conical, round in cross section (fig. 74). Guatemala, aspericollis Gillet. 38 VOLUME XLI Elytral striae appearing impnnctate to the unaided eye, the punctures obsolescent; apices of median pronotal promi- nences pyramidal, their lower surfaces flattened or slightly excavate (fig. 72.). Costa Rica, Panama suhp'imctatus Gillet. 18. Anterior face of pronotum sparsely to moderately punctate (fig. 71). Central Mexico klugi Harold. Anterior face of pronotum granulate or asperate, or if partly punctate, then very densely so, the punctures with raised edges, imparting an asperate appearance to the surface 19. 19. Apices of median pronotal prominences pyramidal, abruptly and sharply turned upward when seen from sides (fig. 73) ; head horn evenly tapering when seen from front. Chiapas and Central America boucardi Harold. Apices of median pronotal prominences blunt or conical, directed forward or evenly curving slightly upward when seen from sides (fig. 81) ; head horn, when seen from front or rear, expanded distally into transverse knob. Chihuahua and Durango moechus Leconte. Key to the Species of the Genus Copris in the Western Hemisphere Based on the Males and Females 1. Lateral pronotal carina absent ; lateral pronotal margin evenly curved ; pygidial margin incomplete, the inner border effaced vent rally 2. Lateral pronotal carina present ; lateral pronotal margin sinuate or slightly angulate; pygidial margin complete or not 6. 2. Median and posterior trochanters with a tuft of long setae on the hind edge. Total length 13.5-19 mm 3. Median and posterior trochanters without setae. Total length 8-13 mm 4. 3. J1 2 3. (Forespur spatulate.) Occipital margin usually with a sharp transverse carina closely paralleling the marginal setigerous groove (fig. 24, toe). J. (Forespur tapering.) Anterior portion of pronotum without any transverse carina; very developed specimens with two low transverse gibbosities not followed by any intensification of the punctures. S. Mexico and Central America lugubris Bolieman. (Forespur spatulate.) Occipital margin usually with a dull transverse carina closely paralleling the marginal ENTOMOLOGICA AMERICANA setigerous groove. J. (Forespur tapering.) Anterior por- tion of pronotnm, even in the least developed specimens, with two indistinct transverse carinae which are rounded and, especially in developed specimens, sinuate, with the disc immediately behind them abruptly and contrastingly densely punctate. Tamaulipas to Ecuador, Hawaii, New Zealand incertus Say. 4. Lateral pronotal margin finely serrate just behind anterolateral angle ; prosternal-proepisternal suture distinctly carinate ; pronotum and head completely devoid of gibbosities in both sexes. S. Mexico and Central America laeviceps Harold. Lateral pronotal margin not serrate ; prosternal-proepisternal suture not carinate ; pronotum sculptured or not ; head usually with a horn-like process 5. 5. Pronotum and elytral striae coarsely punctate; a punctate segment of 9th elytral stria present at base of elytron; proepimeron entirely, evenly punctate ; clypeus contrast- ingly more finely punctate than rest of head; forespur linear, rounded at the end (fig. 6). Eastern U. S. minutus (Drury). Entire body very finely punctate ; elytral striae partly im- punctate; proepimeron largely impunctate inside posterior longitudinal proepimeral carina; forespur cresent-shaped, curving outward (fig. 35). Florida gopheri Hubbard. 6. Anterolateral angles of pronotum with the point made salient by an inward curve or indentation of lateral margin im- mediately behind them (sometimes indistinct), the margin often sharply sinuate anteriorly; 8th elytral stria nearly always complete. Total length 10-18 mm 7. Anterolateral angles of pronotum subquadrate, obtusely angled, or broadly rounded, never with a sharp inward curve of margin immediately behind them. Total length 14-29 mm 17. 7. Pronotum uniformly very densely punctate, without smooth areas except at the very apices of prominences, if any 8. Pronotum with impunctate areas, or large areas with finer or sparser punctures than others 10. 8. Clypeal margin entire, slightly sinuate at most; elytral in- tervals appearing smooth, convex. Florida inemarginatus Blatchley. Clypeus distinctly emarginate ; elytral intervals appearing distinctly punctate to the unaided eye, moderately convex or flat; elytral striae crenulate, the punctures obsolescent 9. 40 VOLUME XLI 9. Elytral interstriae moderately punctate, convex, the punctures separated by a distance equal to 1-3 times their diameter. Central and N. E. U. S fricator (Fabricius). Elytral interstriae more densely punctate, flat, the punctures separated by a distance about equal to their diameter ; both sexes always completely devoid of pronotal prominences. Florida howdeni Matthews and Halffter. 10. Pygidial margin incomplete, its inner border completely effaced vent rally (fig. 58) 11. Pygidial margin complete, its inner border entirely en- graved, or incomplete but with median portion of inner border represented by close-set punctures (fig. 59) 13. 11. Median longitudinal sulcus of pronotum coarsely punctate and deeply impressed; anterior margin of pronotum forming a minute, downwardly directed median tooth ; median coxae with some coarse punctures on outer faces. Chiapas and Central America costaricensis Gahan. Median longitudinal sulcus of pronotum shallow and im- punctate or finely punctate, the punctures never umbilical ; anterior margin of pronotum not forming any median tooth 12. 12. Anterior margin of prosternum with a minute, acute median tooth (fig. 54) ; basal part of disc and median longitudinal sulcus of pronotum quite impunctate. Central Mexico rebouchei Harold. Anterior margin of prosternum without any median tooth (fig. 53) or with a very low broad lobe which may be trun- cate or bidentate apically; basal part of disc and median longitudinal sulcus of pronotum finely punctate in northern specimens. S. Arizona and W. Coast of Mexico to Te- huantepec lecontei n. sp. 13. Anterior margin of prosternum with a salient median process which is bilobed at the apex (fig. 55) ; base and median longitudinal sulcus of pronotum quite impunctate ; a deep depression between median pronotal prominences in both sexes ; $ with a typical female head horn. Guerrero halffteri Matthews. Anterior margin of prosternum without a salient, bilobed median process 14. 14. Forespur curved inward near the apex and tapering to a sharp point (figs. 40, 44) 15. Forespur straight and bluntly rounded at the apex, or abruptly bent inward very near apex and narrowing to a dull point (figs. 43, 45) 16. 41 ENTOMOLOGICA AMERICANA 15. Hind angles of head quadrate ; with a broad triangular median emargination of clypeal margin; base of pronotum and median longitudinal sulcus finely punctate or im- punctate, the puncturation gradually intensifying an- teriorly. S. Arizona and W. Coast of Mexico to Tehuan- tepec lecontei n. sp. Hind angles of head acute; J' with a very shallow, barely perceptible emargination and two remote, minute teeth on clypeal margin; base of pronotum impunctate except for median longitudinal sulcus, which is coarsely umbilico- punctate. Michoacan mexicanus Matthews and Halffter. 16. Eighth elytral stria incomplete, effaced or at least partly dis- integrating posteriorly; anterior pronotal margin with a minute, acute median tooth. Veracruz and Chiapas sallei Harold. Eighth elytral stria complete ; anterior pronotal margin with- out any median tooth; $ with a very shallow, arcuate median emargination of clypeal margin ; base of pronotum impunctate or finely punctate, becoming abruptly coarsely punctate anteriorly in the depressions. Texas, Nuevo Leon, Tamaulipas remotus Leconte. 17. Eighth elytral stria complete ; pronotal puncturation rather dense, umbilical, usually becoming sparser posteriorly, never asperate or granulate on the anterior face ; § head horn usually widely expanded distally; with a very shallow, arcuate clypeal emargination and curved, acute forespurs (when not worn). S. Arizona and New Mexico, W. Texas, Chihuahua arizonensis Schaeffer. Eighth elytral stria incomplete, disintegrating posteriorly (sometimes almost complete) ; pronotal puncturation much less dense on disc ; 5 head horn either parallel-sided or narrowing apically; forespurs of both sexes straight and with blunt apices ; J1 clypeal emargination angular or absent 18. 18. Elytral striae appearing impunctate to the unaided eye, the punctures obsolescent, quite effaced on the 8th stria. Costa Rica, Panama subpunctatus Gillet. Elytral striae appearing distinctly punctate to the unaided eye 19. 19. Clypeal margin entire, devoid of teeth or emargination; ventral surfaces of median and posterior femora impunctate. Total length 28-30 mm. Michoacan megasoma Matthews and Halffter. 42 VOLUME XLI Clypeal margin with at least a small median notch, often also with two broad, low teeth or expansions ; ventral surfaces of median and posterior femora with at least a few punctures distally. Size smaller, total length 14—25 mm 20. 20. Clypeal margin very broadly, shallowly emarginate, with two very small teeth (often worn) ; anterior face of prono- tum entirely punctate, the punctures large, umbilical, and often dense, not accompanied by pronounced asperation. 2- Anterior face of pronotum with at least a few umbilical punctures along anterior margin interspersed among the dense rugosities or granules of the surface. Central Mexico to Chihuahua, in the mountains klugi Harold. Clypeal margin with a small, acute median notch and no marginal expansions, or anterior face of pronotum granular, asperate, or both punctate and asperate. J. Anterior face of pronotum uniformly densely granular or sparsely to moderately asperate 21. 21. Clypeal margin with a small acute median notch and no salient teeth. Anterior face of pronotum punctate, with no rugosities. J. Anterior margin of prosternum usually with a quadrate median lobe ; anterior face of pronotum sparsely to moderately asperate, never granulate. Central Mexico armatus Harold. Clypeal margin with two low, indistinct expansions; anterior face of pronotum granulate, usually densely so 22. 22. Complete portion of 8th elytral stria fully punctate like the other striae ; anterolateral angles of pronotum angulate, the angle very obtuse. Chihuahua, Durango moechus Leconte. Complete portion of 8th elytral stria more feebly punctate than striae I-IV ; anterolateral angles of pronotum broadly rounded (in undeveloped specimens). Chiapas and Central America 23. 23. Basal part of pronotal disc impunctate, shiny. Chiapas, Guatemala, El Salvador boucardi Harold. Basal part of pronotal disc profusely but shallowly punctate, dull. Guatemala aspericollis Gillet. Group I. The minutus group. Outer face of apical maxillary palpal segment convex. Lateral pronotal carina absent. Lateral pronotal margin evenly curved. 43 ENTOMOLOGICA AMERICANA Male median pronotal prominences, when present, broadly trans- versely truncated. Complex punctures with minutely granular texture. Sternellum tending to be longitudinally carinate. Pygid- ial margin always incomplete. Male genital parameres slender, tapering to a fairly acute apex. Individuals are frequently in- completely pigmented. Five species. Complex 1. The incertus complex. Lateral pronotal margin finely and irregularly serrate just behind the anterolateral angles. Prosternal-proepisternal suture strongly carinate. Three closely related species of tropical Mexico, Central America, and northeastern South America : incertus Say, lugubris Boheman and laeviceps Harold. C opr is incertus Say Copris incerta Say, 1835, Boston Journ. Nat. Hist., 1 : 175 [type : Mexico ; Museum of Comparative Zoology] ; 1859, Com- plete Writings, ed. Le Conte, II : 649 ; Harold, 1869, Ann. Soc. Ent. France, ser. 4, IX: 494 (key); Bates, 1887, Biol. Cent.- Amer., Coleop. II, 2, p. 55 (distr.) ; Heyne and Taschenberg, 1908, Die exotischen Kafer, p. 64 (descr.) ; Pereira and d’An- dretta, 1955, Pap. av. Dep. Zool. Seer. Agric., S. Paulo, XII : 261-263 (descr. and syn.) ; Matthews, 1959, Ciencia XIX (6-7) : 135 (key and distr.) ; Thomas, 1960, N.Z. Journ. Sin., Ill (1) : 8-14 (biol. and distr.). Copris procidua Say, 1835, Boston Journ. Nat. Hist. I : 176 [type: Mexico; Museum of Comparative Zoology] ; 1859, Com- plete Writings, ed. Le Conte, II : 649 ; Harold, 1859, Ann. Soc. Ent. France, ser. 4, IX: 495 (key) ; 1880, Stettiner Ent. Zeit. XLI : 27 (distr.) ; Blanchard, 1885, Trans. American Ent. Soc. XIII: 171 (distr.); Bates, 1887, Biol. Cent.-Amer., Coleop. II, 2, p. 54 (distr.) ; Schaeffer, 1906, Trans. American Ent. Soc. XXXII: 255 (key) ; Williams, 1929, Proc. Hawaiian Ent. Soc. VII (2) : 210, 227, 237 (distr.) ; Pereira and d’Andretta, 1955, Pap. av. Dep. Zool. Seer. Agric., S. Paulo, XII: 260-263 (distr. and syn.) ; Matthews, 1959, Ciencia XIX (6-7) : 135 (key and distr. ) . 44 VOLUME XLI Description of Male. Head. — Armed or not. Clypens bi- dentate, clypeal teeth not at all prominent, margin between them broadly, angularly emarginate with a slightly deeper Y-shaped median notch not cutting through margin. Posterior angles of genae subquadrate or slightly obtuse. Upper surface of head rather densely punctate, the punctures simple except for those behind horn (between eyes), which are coarse and granular. Posterior oblique carina absent. Occipital margin with transverse setigerous groove intact and paralleled by an incomplete, dull carina, which is sometimes absent. Demarcation between gula and submentum usually broadly V-shaped, sometimes rounded. Thorax. — Pronotum armed or not. Anterolateral angles obtuse, lateral margin just behind them irregularly serrate. Lateral margin evenly arcuate. Anterior margin of pronotum not forming any median point or angle. Median longitudinal sulcus deep, coarsely punctured. Puncturation of pronotum as follows : sparsely and very finely punctate, or impunctate, on disc, becoming a little more densely and deeply punctate on anterolateral lobes and anterior declivities, the punctures here simple ; grossly umbilico-punctate, the punctures granular, along entire submargin (sparser laterally and anteriorly), along dorsal longitudinal sulcus, and in lateral fossae. Anterior prosternal margin with a rounded median tooth ; sternellum concave with an indistinct median longitudinal carina, moderately punctate. Median lobe of metasternum with no coarse punctures; median longitudinal groove complete. Elytra. — 8th stria obsolescent, effaced at base and for median third of elytral length, present intact only for a short length near base and behind posterior angle ; 9th stria arising from 10th about halfway down elytron; 10th complete or disintegrating temporarily for median third of elytral length. Striae moderately punctate, the punctures not deep, round, little wider than striae, separated by a distance equal to more than their diameter. Interstriae slightly convex, sparsely and very finely punctate, appearing smooth. Abdomen. — Pygidium punctate only clorsally, the punctures granular ; pygidium incompletely margined, inner edge of margin being effaced ventrally. Anterior legs. — Ventral surface of femur with coarse setigerous punctures on pos- terior longitudinal half, finely punctate on anterior. Forespur expanded and truncated distally. Middle legs. — Coxa with some indistinct punctures along median carina. Trochanter with a tuft of long setae arising from posterior margin. Ventral surface of femur very finely punctate, appearing smooth, except for a few setigerous punctures at distal end in some specimens. Tibia below with one distal seta tuft, above with 1-3 supplementary setae distally. Posterior legs. — Trochanter with a tuft of long setae 45 ENTOMOLOGICA AMERICANA arising from posterior margin. Ventral surface of femur without coarse punctures or with several coarse punctures at distal end. Tibia below with one distal seta tuft, above with 0-3 supplementary setae. Total length. — 13.5—18.5 mm. Male armament. The least developed male seen possessed neither a head horn nor any pronotal prominences. More developed speci- mens bear a slightly curved, tapering clypeal horn which is situated forward of the middle of the head (the more developed the horn the more anteriorly it is located), and an acute tubercle just behind it which is strongly inclined forward, almost recumbent (fig. 63). The median pronotal prominences are very low, approximated, and broadly, transversely truncated. The upper edges of the truncated faces, when seen from the front, are practically on a horizontal plane. The lateral pronotal prominences are low, conical, and directed upwards. The males never achieve a very high degree of secondary sexual development (compared to lugubris). Description of Female. Similar to male, but differing in armament and in the following features : clypeal teeth more promi- nent, rounded, approximated, the notch between them deeper, more rounded; forespur not expanded distally, tapering to a blunt point ; pronotum anteriorly more densely punctate ; median and posterior tibiae with more seta tufts distally below (1-3) and more supplementary setae above (2-4). Total length. — 13.5-18.5 mm. Female armament. Head sometimes devoid of a horn in small specimens, with only an acute transverse cariniform process, but usually bearing a narrow transverse horn which is truncated and excavate apically. Median pronotal prominences quite different from the male’s, indistinctly transversely carinate, the carinae vis- ible even in undeveloped specimens close to the anterior margin. In large specimens these carinae are sinuate, their anterior faces moderately punctate, the disc behind them abruptly and con- trastingly densely punctate. Lateral pronotal prominences absent. Distribution. Fig. 12. This species has a remarkably extended distribution which appears to be made up of isolated populations. It is represented in numerous localities in the tropical forest of eastern Mexico north of Veracruz and in Yucatan at altitudes of 150-1360 m. (500-4500 ft.). It does not occur along the coast itself where it is replaced by the closely related C. lugubris Boh. To the south the species disappears completely (in collections) to show up again among some specimens collected by Nevermann on the lower reaches of the Reventazon in Costa Rica, where it appears to occur in association with C. laevicegs Harold. Thence it dis- appears again and we do not find it until we reach the Cordilleras of Colombia and Ecuador, where it occurs at altitudes of 1050-1800 46 VOLUME XLI m. (3500-5800 ft.). Blanchard and Horn mention this species (under the name prociduus) as occurring in Texas, but I have seen no reliable records to support this and in view of the nature of its habitat in Mexico I consider it highly unlikely that it occurs in the United States (other than in Hawaii), except perhaps as an occasional stray. Blanchard further mentions having seen a speci- men from Guatemala. In addition to the localities cited below, Pereira and d’Andretta Fig. 12. Distribution of the incertus complex, I. Base map reproduced by permission of the University of Chicago. give five additional Colombian, six additional Ecuadorean, and five additional Hawaiian localities which certainly refer to this species. This species has been introduced into some Pacific islands in order to reduce the amount of exposed cow dung, which serves as a breeding medium for several species of flies. It has become successfully established in Hawaii, Western Samoa, and New Zea- land (Thomas, 1960). 47 ENTOMOLOGICA AMERICANA Biology. Recorded from cow dung in Mexico and New Zealand and cow and horse dung in Hawaii. Nevermann in Costa Rica found it under human dung, in dried wood, and in undergrowth at night. There is one Hawaiian record of its occurrence on Opuntia cactus attacked by Fusarium disease. Nidification is discussed previously. Remarks. I have examined three specimens from the Say ma- terial in the Harris collection at the Museum of Comparative Zoology which agree with Say’s original description of incerta. All are females (Say’s description was that of a female, as he suspected), two of them simply bear one very small label each with the number “51” on it while the third bears three labels: an old handwritten one with ‘ ‘ incerta Say ’ ’ and ‘ ‘ Mexico ’ ’ written on it, a newer handwritten one (Harris’s?) with “type Mexico” on it, and likewise a very small one with the printed number “51”. The latter specimen is here designated as lectotype of incertus Say. This same collection also contains a single specimen bearing the name procidua. It is a male of the same species and agrees perfectly with Say’s original description. It also bears three labels saying “ procidua Say” and “Mexico”, “type Mexico”, and the number “53”. Since this is the only male of this species in the collection, it is here considered to be the holotype of prociduus Say. All these specimens belong to the same species and since the female was described first, this species (which has hitherto been called prociduus or incertus var. prociduus ) must now bear the name incertus — engendering considerable nomenclatorial confusion, be- cause the name incertus has consistently been applied to a closely related, very abundant species here considered as lugubris Boheman. The separation of this species from the closely related lugubris Boheman and laeviceps Harold is discussed under the remarks pertaining to those two species. Material Examined. 99 males, 106 females, including lectotype. UNITED STATES: Hawaii: Mapulchu, Molokai; Maui. MEXICO: Hidalgo: Chapulhuacan ; Puebla: Necaxa; San Diego ; Mesa de San Diego ; Villa Juarez ; San Luis Potosi: El Salto ; ITuichihuyan, 20 mi N Tamazunchale ; Tamazunchale ; Tamaulipas: 3 mi NW Acuna; Quintero; Veracruz: Banderilla; Barranca de Metlac; Cordoba; Fortin; Jalapa; Laguna de Tamihaua, 20 km N Tuxpan; Martinez de la Torre; Orizaba. Yucatan: Uxmal. COSTA RICA: Cartago: Turrialba, 800 m (Schild), 1 $ (USNM) ; 31 May 1951 (O. L. Cartwright), 1 (USNM). Limon: Hamburgfarm, Reventazon, Plain of Limon, 21 Jul. 1931, 27 Jul. 1935, 21 May 1936 (F. Nevermann), 4 6 $2 (USNM). 48 VOLUME XLI COLOMBIA: Cundinamarca: Fusagasuga, 1800 m, 9 Sep. 1942 (F. J. Otoya), 6 2 ?? (USNM) ; 8 May 1946 (E. A. Chapin), 3 eft?, 2?? (USNM). ECUADOR: Guayas: Naranjal (F. Campos R.), 1 2 (USNM) ; Los Rios: 21 Apr. 1938 (W. MacIntyre), 1 £ (OLC) ; location un- determined: Paramba, 3500 ft, Mar. 1897 (Rosenberg), 1 £ (BM) ; Lita, 1 g (BM) ; San Rafael (F. Campos R.), 1 J1 (USNM). Copris lugubris Boheman Copris lugubris Boheman, 1858, Eugenies Resa, Coleop., p. 42 [type : Galapagos I. ; Naturhistoriska Ricksmuseum, Stockholm] ; Waterhouse, 1877, Proc. Zool. Soc. London V : 82 ; Linell, 1898, Proc. United States Nat. Mus. XXI : 258 ; Felsche, 1901, Deutsche Ent. Zeitschr., p. 145; Mutchler, 1925, Zoologica V(20): 237; Van Dyke, 1953, Coleoptera of the Galapagos Islands, p. 122; Pereira and d’Andretta, 1955, Pap. av. Dep. Zool. Seer. Agric. S. Paulo, XII: 261 (synon.). Description of Male. Head. — Armed. Clypeus bidentate, clypeal teeth relatively prominent, margin between them shallowly, angularly emarginate without any median notch. Posterior angles of genae subquadrate or obtuse. Upper surface of head largely im- punctate basally, becoming moderately punctate on flattened por- tions and horn, punctures simple ; in small specimens there are some granular punctures between the eyes. Posterior oblique carina obsolescent. Occipital margin with transverse setigerous groove complete, closely preceded by a sharp transverse carina which may be complete or interrupted at middle. Demarcation between gula and submentum a deep arc, usually with a median V-shaped area. Thorax. — Pronotum armed or not. Anterolateral angles obtuse, the lateral margin just behind them irregularly serrate. Lateral margin rather evenly arcuate without any pro- nounced angulation. Anterior margin of pronotum not forming any median point or angulation. Median longitudinal sulcus deep, coarsely punctured. Puncturation of pronotum as follows : sparsely and finely punctate on disc becoming a little more densely and deeply punctate on anterolateral surfaces and anterior declivity, the punctures simple here ; grossly punctate, the punctures um- bilical and granular, only along hind submargin (and occasionally along fore submargin), along the dorsal sulcus, and in the lateral fossae. Anterior prosternal margin with a prominent, rounded 49 ENTOMOLOGICA AMERICANA median tooth • sternellum somewhat concave but with an indistinct median longitudinal carina, moderately punctate. Median lobe of metasternum very finely punctate, with no coarse punctures; median longitudinal groove very indistinct anteriorly. Elytra. — 8th stria obsolescent, effaced at base and median third of elytron, present integrally only for a short length near base and apically; 9th stria arising at anterior third of elytral length and continuing distally; 10th stria complete. Striae indistinctly punctate, the punctures not deep, round, little wider than stria, separated by a distance equal to more than their diameter over most of strial length. Interstriae very slightly convex, sparsely and very finely punctate, appearing smooth. Abdomen. — Pygidium moderately punctate dorsally, the punctures granular, becoming sparser ven- trally; margin incomplete, effaced ventrally. Anterior legs. — Ven- tral surface of femur entirely punctate, the punctures coarser and setigerous on posterior longitudinal half. Forespur rather straight, expanded and truncate at apex (fig. 33). Middle legs. — Coxa faintly punctate along median carina. Trochanter with a single tuft of long setae arising from posterior edge. Ventral surface of femur sparsely and very finely punctate, usually with a few setig- erous punctures distally. Tibia below with a single distal seta tuft, above without supplementary setae. Posterior legs. — Tro- chanter with a single tuft of long setae arising from posterior edge. Ventral surface of femur very sparsely and finely punctate. Tibia below with a single distal seta tuft, above usually without supplementary setae apically, sometimes with one or two. Total length. — 13.5-18 mm. Male armament. The least developed male seen possessed a head horn, albeit a very low one, on the posterior part of the clypeus and a barely evident tubercle close behind it. The pronotum showed traces of median prominences in the form of two approxi- mated transverse tumosities and not a trace of any lateral promi- nences. With further development, the head horn elongates, be- coming gently arcuate and slightly dilated apically in the most developed specimens. The horn is situated at about the middle of the head. Behind it is a prominent tubercle which is either erect or bent forward, but never inclined forward (fig. 64). The median pronotal prominences are broadly transversely truncate, closely approximate, their outer edges converging anteriorly in dorsal view. The upper edges of the truncated faces, when seen from the front, are very strongly sloping down to the sides at about a 45° angle from the horizontal or more. The lateral prominences are laminate, directed forward and upward, and parallel to the longitudinal axis when viewed from above. 50 VOLUME XLI Description of Female. Similar to the male, but differing in armament and in the following features : clypeal teeth more promi- nent and approximated, with a V-shaped median notch in margin between them. Forespur not expanded and truncate distally, nar- rowing slightly to a rounded apex. Pronotum anteriorly more densely punctate. Hind tibia above with 2-4 supplementary setae distally (fig. 50). Total length. — 14-19 mm. Female armament. Many females are completely unarmed, with only a short transverse carina on the head. The head horn, Fig. 13. Distribution of the incertus complex, II. Base map reproduced by permission of the University of Chicago. when developed, is much less transverse than is usual in the genus and is often quite high and narrow; it is excavate on the posterior surface. The median pronotal prominences in developed specimens are like those of a minor male, that is, low rounded tumosities ; the lateral ones are absent. There is never an abrupt intensification of the dorsal pronotal puncturation just behind the median tumosities. 51 ENTOMOLOGrlCA AMERICANA Distribution. Fig. 13. Widely distributed throughout South- ern Mexico and Central America to the Panama Canal at all altitudes up to 1500 m. (4500 ft.). In the northern part of its range it extends up along the coastline on both sides of Mexico, occurring along the Gulf Coast in the sandy area just behind the sea beach. A single specimen was taken surprisingly far inland in northern Coahuila and might be a stray. It has been recorded from Colombia, Ecuador and Hawaii, but I have seen no reliably labelled specimens from these areas and I believe all these records refer to the closely related C. incertus Say. Biology. Some aspects of nidification have been discussed previously. The collected specimens bear the following data : at light, under cow dung, in a banana trap, in dead calf, on avocado. I have collected it under burro dung and human faeces. It appears to be most frequently collected at light. In Central America it is active throughout the year. In Central Mexico it is one of the only two species of coprophages occasionally encountered above ground during the dry season (G. Halffter, oral communication). Remarks. This species has long gone under the name incertus Say. In the remarks under the species now bearing that name I pointed out that Say’s types in the Museum of Comparative Zoology show that he did not have any specimens of this species. I have examined the type of C. lugubris Bolieman in the Naturhistoriska Ricksmuseum, Stockholm, and have found it to be this species. C. lugubris is therefore its valid name. C. lugubris may be told from incertus, to which it is closely related, by a number of characters in both sexes. Specific distinc- tion is shown beyond doubt by the differences in the male secondary sexual characters, which never show intermediacy between the two forms. The female also shows very distinct differences in pronotal sculpturing (compare descriptions of female armament) and, be- cause of the frequent occurrence of minor males in both species, it is actually often easier to separate the two species on the basis of the females. Additional, less reliable, differences are seen in the male genital parameres (which bear a minute dorsal hook at the apex in incertus (fig. 61) but not in lugubris) and, to a lesser extent, in the occipital margin of the head, which in lugubris usually bears a sharp transverse carina, whereas in incertus this carina is usually dull or absent altogether. Material Examined. 303 males, 360 females. MEXICO. Campeche: Ciudad del Carmen; Chiapas: Comitan; El Vergel (Volcan Soconusco) ; Escuintla; ?Finca “La Isla”; La Esperanza (Tapachula region) ; Las Cruces; Ocosingo; Pacific slope Cordilleras; Palenque; ?San Jose; ?Sta. Julia; Suchiate; Tapa- 52 VOLUME XLI chula; Tuxtla Gutierrez; Coahuila: 10 mi. S. Allende; Colima: Colima; Volcan de Colima; Distrito Federal: Mexico; Guerrero: Acapulco; Acuitlapan (nr. Cacahuamilpa) ; Balsas; Cacahuamilpa; Chilpancingo ; 22 mi. N. Chilpancingo ; El Margues, 28 km. W. Acapulco; Iguala; 3 mi. N. Mexcala; Taxco; Teloloapan; Jalisco: Chapala; Purificacion; Mexico: Rio San Jeronimo, 10 km. NE. Cacahuamilpa; Tenancingo; ?Tenayuca; Tonatico; Michoacan: San Jose Puma; Morelos: Cuautla; Cuernavaca; 10 mi. S. Cuernavaca; Jojutla; fOaxtepec; Yautepec; Nayarit: Tepic; 17 mi. NW. Tepic; Oaxaca: Amapa; Ixcatlan (W. of Tuxtepec) ; Oaxaca; 25 km. S. Oaxaca; Panixtlahuaca ; Soyaltepec (N. of Tuxtepec) ; Tehuan- tepec; ?Valerio Trujano; Quintana Boo: Bacalar; Sinaloa: Es- cuinapa; Mazatlan; Sonora: Rio Mayo; Tabasco: Teapa; Villa- hermosa; Veracruz: Atoyac ; Coatepec; Cotaxtla; La Gloria, Cardel ; Lake Catemaco ; Minatitlan ; Papantla ; Presidio ; Puente Nacional; Pureza, 5th RR. station Veracruz to Jalapa; San Andres Tuxtla ; San Carlos ; San Martin, San Andres Tuxtla ; Tecolutla ; Tlapacoyan; Tres Zapotes; Veracruz; Yucatan: Chichen Itza. GUATEMALA: Alta Vera Paz: Cacao Trece Aguas; Coban; San Miguel Tucuru; Baja Vera Paz: Rabinal; San Jeronimo; Chi- quimula: Chiquimula; Guatemala: Guatemala City; San Jose de Pinula; Izabal: Cayuga; Los Amates; Jutiapa: Laguna Atesca- tempa; Peten: La Libertad; fPacomon; San Marcos: San Marcos; Suchitepequez: ?Finca El Cipres; Moca; Variedades; Zacapa: Za- capa; location undetermined: Jocobo. BELIZE : Belize ; Benque Viejo ; San Antonio ; Punta Gorcla. EL SALVADOR: La Libertad: Santa Tecla; La Union: La Union; San Salvador: San Salvador; San Vicente: 49 Km. E. San Salvador ; Sonsonate: Izalco ; Sonsonate. HONDURAS: Atlantida: La Ceiba; Tela; Copdn: Copan; Mo- razdn: Esc. Agr. Pan. Zamorano; Santa Barbara: Choloma; Quim- istan; Tegucigalpa: Tegucigalpa; Yoro: Progreso; location unde- termined: Carmelina; Ruatan I. NICARAGUA: Chontales ; Jinotega: Jinotega; Madriz: So- moto ; Managua: Managua ; 8 mi N. Managua ; San Antonio. COSTA RICA : Alajuela: Greeia ; Guanacaste : Las Canas ; Gua- nacaste; Piedra Negra; Santa Elena; Heredia: Santa Clara; Pun- tar enas: Las Loras nr. Puntarenas; Orotina; San Lucas I., Gulf of Nicoya; San Mateo; San Jose: Santa Ana; San Jose. PANAMA: Canal: Alhajuelo; Chiriqui: David; Potrerillos; Volcan de Chiriqui; Panama: La Chorrera; Chepo; Taboga I.; location undetermined: La Joya. 53 ENTOMOLOGICA AMERICANA Copris laeviceps Harold Copris laeviceps Harold, 1869, Ann. Soc. Ent. France, ser. 4, IX: 498 [type: San Andres Tnxtla, Yer. ; Museum d’Histoire Naturelle, Paris] ; Bates, 1887, Biol. Cent.-Amer., Coleopt. II, 2, p. 54 (distr.) ; Matthews, 1959, Ciencia XIX (6-7) : 135 (key and distr.). Description of Male. Head. — Unarmed (with but a low coni- cal process on the clypeus. Clypeus bidentate, clypeal teeth promi- nent, angular, the margin between them broadly angulate with a slightly deeper median V-shaped notch not cutting through margin. Posterior angles of genae obtuse. Upper surface of head very finely punctate, appearing impunctate, except for a dense trans- verse band of coarse umbilical and granular punctures extending between eyes. Posterior oblique carina absent. Occipital margin with transverse setigerons groove complete ; marginal occipital carina paralleling setigerons groove very sharp and interrupted in middle. Demarcation between gula and submentum arcuate or subangulate. Thorax. — Pronotum unarmed. Anterolateral angles subquadrate. Lateral margin evenly curved, without angulations, indistinctly and irregularly serrate just behind anterolateral angles. Anterior margin of pronotum not forming any median point or angle. Median longitudinal sulcus impressed, complete, coarsely punctate. Puncturation of pronotum as follows : very finely punctate, appearing impunctate, except for the following areas, which are coarsely punctate, the punctures umbilical and granular : along entire submargin, on anterolateral lobes, in lateral fossae, in a narrow chain along entire median longitudinal sulcus, and form- ing two large patches just above lateral fossae on either side ; these patches are isolated from all other coarse punctures except those of lateral fossae. Anterior prosternal margin with a low median salience ; sternellum not very concave with a suggestion of a median longitudinal carina anteriorly, coarsely, closely punctate, the punc- tures granular and umbilical. Median lobe of metasternum with some coarse granular punctures anteriorly along sides; median longitudinal groove somewhat effaced anteriorly. Elytra. — 8th stria arising close to base but not from it, disintegrating halfway down elytron, then resuming intact after hind angle ; 9th stria arising from 10th anterior to the halfway point; 10th complete. Striae coarsely punctate, the punctures round, umbilical, separated by a distance equal to a little more than their diameter. Inter- striae slightly convex, very finely punctate, appearing smooth. When seen from behind, elytral surface is bent, forming an in- 54 VOLUME XLI distinct, obtuse ridge running from hind angle to end of median suture ; no striae cross this ridge. Abdomen. — Pygidium densely and coarsely punctate, the punctures granular ; pygidial margin in- complete, becoming effaced ventrally. Anterior legs. — Ventral sur- face of femur coarsely punctate, the punctures setigerous, on pos- terior longitudinal two thirds, very finely punctate on anterior third. Forespur slightly expanded distally, rounded and curved down at apex. Middle legs. — Coxa with a few coarse punctures on outer face anteriorly near median carina. Trochanter without setae. Ventral surface of femur very finely punctate with a few coarse punctures distally. Tibia below with 1-2 distal seta tufts, above with 1-2 supplementary setae. Posterior legs. — Trochanter with- out setae. Ventral surface of femur coarsely punctate over most of distal half. Tibia below with one distal tuft, above with 1-3 sup- plementary setae. Total length. — 11.5-12.5 mm. Description of Female. Differs from the male only in having a parallel-sided or tapering forespur (instead of a distally ex- panded one). In worn specimens this difference is obliterated. Total length. — 11-13 mm. Distribution. Fig. 12. Known from isolated localities from Jalapa, Ver., to the Reventazon River in Costa Rica. Appears to be an east coast lowland form and may occur throughout the “Mosquitia” coastal plain of Central America, a very poorly collected region. The localities given by Bates are all probably correctly attributed to this species. Biology. Nevermann records the following data on his spec- imens from the Reventazon River and Plain of Limon, Costa Rica : under horse dung, human dung, in the undergrowth at night in the primeval forest (“nachts im Urwald am Gebiisch”), in iguana car- cass, at light. Active throughout the year. Remarks. This species is very closely related to incertus Say and lugubris Boh., with which it shares the features characterizing the minutus group, plus several others such as the serrate lateral pronotal margin, broadened male forespur, and carinate prosternal- proepisternal suture characterizing the incertus complex. In ad- dition, it shows distinctly closer affinities with lugubris, with which it shares the type of male genital parameres and sharp transverse occipital marginal carina. However, all the specimens I have seen may be easily told from both incertus and lugubris by the lack of setae on the posterior margin of the median and posterior tro- chanters; it differs also in the pronotal puncturation, showing an oval patch of coarse punctures on the sides of the pronotal disc, and in the carinate nature of the posterior elytral angles. All specimens 55 ENTOMOLOGICA AMERICANA are small in size and totally devoid of cephalic and pronotal arm- ament. Material Examined. 39 males, 26 females. MEXICO. Chiapas: Tnxtla (0. W. Barret), lJ'(USNM); Veracruz: Jalapa (Hoege), 2 (BM) ; Presidio, E of Zongolica, 1200 m., Jul. 1952 (G. Halffter), 1 $ (GH) ; national record only: (Bock), 1 (CAS). Copris moechus Leconte Copris moecha Leconte, 1854, Proc. Acad. Nat. Sci. Philadel- phia VII : 222 [type : Camp 14* ; Museum of Comparative Zo- ology] ; Leconte, 1858, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, IV : 9-42 (distr.) ; Horn, 1873, Trans. American Ent. Soc. IV : 42-51 (key and descr.) ; Schaeffer, 1906, Trans. American Ent. Soc. XXXII: 255 (key). * I have not been able to determine the location of “Camp 14”. This refers to the United States and Mexican Boundary Commission explorations of 1850-1853 during which one member of the ex- peditions, Dr. Thos. H. Webb, made “large collections ... in the region between the Rio Grande and the Colorado River of California, chiefly in the valley of the Gila” (Leconte, 1858). Leconte in the 73 ENTOMOLOGICA AMERICANA Copris clavicornis Matthews and Halffter, 1959, Ciencia XVIII (9-10) : 191-194 [type : 100 km W of Sta. Barbara, Chih. ; American Museum of Natural History] ; Matthews, 1959, Ciencia XIX (6-7) : 136 (key and distr.). New synonymy. Description of Male. Head. — Armed. Clypeus with two low angular teeth and a U- or V-shaped notch between them, not cutting through margin. Upper surface of head densely, evenly punctate, the punctures simple, with a basal impunctate area between eyes and horn, and behind both. Posterior oblique carina reduced. Occipital margin with transverse setigerous groove interrupted into three sections, the latter ones displaced forward and partly over- lapping the median. Demarcation between gula and submentum arcuate. Thorax.- — Pronotum armed. Anterolateral angles obtuse, lateral margin curved inward slightly behind origin of lateral carina. Lateral carina sharp, issuing from margin. Anterior margin not forming any median point or angle. Dorsal median longitudinal sulcus complete and impressed, sparsely umbilico- punctate. Puncturation of pronotum very variable, as follows : basal part of disc shallowly punctate, the punctures usually simple, or impunctate ; rest of disc densely punctate, the punctures usually simple ; posterior and lateral submargins, lateral fossae, and de- pressions between median and lateral prominences coarsely um- bilico-punctate ; the anterior surfaces may be entirely asperate (as in the holotype of clavicornis) or more usually both asperate and punctate, the punctures simple or umbilical, the rugosities and punctures always very dense (one specimen seen was densely umbilico-punctate over the entire pronotum, without asperation). Anterior prosternal margin with a broad, low, rounded median lobe (no median tooth) ; sternellum moderately punctate. Median lobe of metasternum impunctate but with some faint punctiform impressions laterally, or umbilico-punctate anteriorly ; median longi- latter work places the type locality of C. moechus in Arizona, while Horn (1873) places it erroneously in Texas. However, a reading of the personal narrative of the leader of the Commission (Bartlett, 1854) reveals that two additional expeditions were carried out: one from the headquarters at the “Copper Mines” (Santa Rita del Cobre, Chih., now Santa Rita, N.M.) to Fronteras and Arispe, Sonora, the other from El Paso south to Chihuahua, Chih. and thence southeast to Saltillo, Coah., and Monterrey, N.L. Dr. Webb took part in both of these and it was almost certainly during the latter expedition, which passed right through the presently known range of moechus, that the type specimen was collected. This places the type locality in Chihuahua, probably near the capital, and this species cannot be considered as occurring within even the present boundaries of the United States. 74 VOLUME XLI tudinal impressed line complete. Elytra. — 8th stria incomplete, disintegrating- at posterior angle ; 9th stria arising about halfway down elytron; 10th stria complete. Striae closely punctate, the punctures transverse, separated by a distance equal to about their width. Interstriae slightly convex, very finely punctate, appearing smooth. Abdomen. — Pygidium completely margined, densely um- bilico-punctate. Anterior legs. — Ventral surface of femur densely and coarsely setigerous-punctate on posterior longitudinal half, finely punctate on anterior half. Forespur relatively straight and parallel-sided, bluntly rounded and somtimes slightly dilated apically. Middle legs. — Coxa finely punctate along median longi- tudinal carina, occasionally with a very few large setigerous punctures along outer edge. Ventral surface of femur finely punctate with a few coarser setigerous punctures distally. Tibia below with 1-2 distal median seta tufts. Posterior legs. — Ventral surface of femur finely punctate, more coarsely so distally, with or without stigerous punctures. Tibia below with 2-3 distal median seta tufts. Total length. — 15.5-20 mm. Male armament. In major males the head horn is elongated and slightly bent, dilated transversely at the apex, and the pronotal prominences are produced, the median ones closely approximated, acute, not at all divergent but not merging, and directed slightly upward, the lateral ones laminate and directed forward parallel to the long axis or slightly divergent (fig. 81). Description of Female. Similar to male but differing in arma- ment and in the following features: clypeal teeth rounded, more prominent, flanking a broader, U-shaped notch. Pronotum punctate on upper part of disc and sides below lateral carina, umbilico- punctate along posterior submargin and in lateral fossae, densely granulate or asperate elsewhere, especially on anterior declivity, which never shows any punctures or depressions. Median longi- tudinal sulcus impunctate and impressed only on superior part of disc. Both median and posterior tibiae with 2-3 distal seta tufts below. Total length. — 13-21 mm. Female armament. Normal for the genus. Distribution. Fig. 16. The mountains of Southern Chihuahua and Northern Durango at altitudes of 1520-2420 m. (5000-8000 ft.). Biology. Unknown. Remarks. This species is extraordinarily variable in the punc- turation of the anterior pronotal face in the male, which ranges all the way from being completely rugose (without punctures) to completely punctate. Of the more than 90 male specimens ex- amined, however, only one was completely punctate (without as- peration) . Possibility for confusion exists in attempting to separate the punctate males of this species from arizonensis Schaeffer and 75 ENTOMOLOGICA AMERICANA Fig. 16. Distribution of the armatus complex, II. Base map reproduced by permission of the University of Chicago. 76 VOLUME XLI klugi Harold, both of which occur in the same area. C. arizonensis may immediately be told by its very shallow, arcuate median clypeal emargination, acute forespur, and fully complete 8th elytral stria. With regard to the latter species, difficulty may be encountered, since klugi and moechus are very closely related. However, klugi is never rugose or asperate on the anterior pronotal surface, but usually relatively sparsely punctate (sometimes almost impunctate). C. moechus, when (very rarely) punctate exclusively here, is very densely so. There also appears to be a geographical or altitude separation between the two species; I have not seen both from the same locality. In Durango, the American Museum Expedition collected moechus exclusively at altitudes of up to 8000 ft. (2430 m.) and klugi exclusively at 8200 ft. (2500 m.) and higher. Upon recently examining the type of this name I was very surprised to find that it belonged to this Mexican species, which had just been redescribed under the name clavicornis Matthews and Halffter. The common Arizona species which has gone under the name moechus for a very long time is here described as new under the name lecontei n. sp. There can be little doubt that the specimen in the Leconte Collection labelled as the type of this species actually is such, since it bears the number “14” corresponding with the type locality given as “Camp 14”. Material Examined. 93 males (including liolotype) and 126 females. MEXICO: Chihuahua: Agua Caliente, Sta. Barbara Dist. ; Buena Vista; Catarinas; Cuevas, Matamoros Dist.; Gaborachic; 8 mi. S Gallego; 12 mi. W Gran Morelos; 10 mi. N Jimenez; Madera; Madera Chic ; 8 mi. W Matachic ; Naica ; Parral ; 15 mi. E Parral ; 2 mi. W Pedernales; Primavera; Salaices; San Jose Babicora ; Santa Barbara; Km. 36 Sta. Barbara-Ojito ; 63 mi. W Sta. Barbara; Valle de Olivos; Durango: Encino; Las Puentes; San Isidro, Cuencame Dist. Copris bo near di Harold Copris houcardi Harold, 1869, Ann. Soc. Ent. Prance, ser. 4, IX : 497-498 [type : Juquila, Oax. ; Museum d’Histoire Naturelle, Paris] ; Bates, 1887, Biol. Cent.-Amer., Coleop. II, 2, p. 54, 1889, Suppl., p. 387 (distr.) ; Matthews, 1959, Ciencia XIX (6-7) : 136 (key and distr.). 77 ENTOMOLOGICA AMERICANA Description of Male Head. — Armed. Clypeus with two barely perceptible teeth, the margin between them very shallowly emargin- ate with a slightly deeper median notch which is sometimes absent. Posterior angles of genae very acute. Upper surface of head closely punctate, the punctures simple, except for impunctate areas around eyes and behind horn. Posterior oblique carina developed, sharp. Occipital margin with transverse setigerous groove interrupted into three parts, the lateral sections displaced forward and partly overlapping median one. Demarcation between gula and submentum arcuate. Thorax. — Pronotum armed. Anterolateral angles subquadrate, margin behind them almost straight. Lateral carina sharp, not issuing from margin anteriorly. Anterior margin of pronotum not forming any median point or angle. Median longitudinal sulcus distinct, complete, umbilico- punctate at least anteriorly. Puncturation of pronotum as follows : very finely punctate, appearing smooth and shiny, over entire base ; lateral surfaces shallowly punctate or ridged; anterior declivity and inside surface of lateral prominences granulate or asperate; posterior submargin, dorsal sulcus anteriorly and also sometimes for its entire length, lateral fossae, and cavities between median and lateral prominences (even in minor males) umbilico-punctate. Anterior prosternal margin variable, either without any median tooth, with a rounded one, or bidentate ; sternellum moderately punctate. Median lobe of metasternum with a few punctiform impressions laterally and anteriorly ; median longitudinal im- pressed line usually complete. Elytra. — 8th stria incomplete, disintegrating about halfway down elytron; 9th stria arising about halfway down; 10th complete. Striae finely but distinctly punc- tate, the punctures round and separated by a distance equal to two or more times their diameter. Interstriae sparsely and very finely punctate, appearing smooth, and slightly convex. Abdo- men.— Pygidium completely margined, moderately umbilico-punc- tate, the punctures small. Anterior legs. — Ventral surface of femur coarsely setigerous-punctate on posterior (upper) longi- tudinal half, impunctate on anterior half. Forespur straight in dorsal view, narrowing apically, the apex broadly rounded. Middle legs. — Coxa impunctate. Femur below distinctly setigerous punc- tate only at distal end, impunctate elsewhere. Tibia below with 1-2 distal seta tufts. Posterior legs. — Ventral surface of femur entirely finely punctate, without setigerous punctures. Tibia with 0-3 seta tufts distally below. Total length. — 19-23.5 mm. Male armament. Minor and medium males have median pronotal prominences which are approximated but not extremely so. Not until well developed males are examined is the characteristic nature 78 VOLUME XLI of the median pronotal prominences seen. In these individuals, the median prominences are more or less pyramidal, very closely approximated, and directed sharply upwards (fig. 73). The head horn is gently curved, not bent, in some instances almost straight, and in other instances massive for most of its length when seen from the side, then abruptly narrowing apically. The lateral pronotal prominences are directed somewhat outward in dorsal view. Description of Female. Similar to male, but differing in armament and in the following features : clypeal teeth more promi- nent, rounded and approximated, with a shallow notch between them. Dorsal longitudinal sulcus of pronotum impressed only basally or entirely very faint. Puncturation of pronotum as follows : impunctate on disc, shallowly punctate on rest of base, anterior half of pronotum densely granulate ; umbilical punctures are present only along posterior submargin and in lateral fossae. Total length. — 19-21 mm. Female armament. Normal for the genus. Distribution. Fig. 15. Found on the volcanoes of northern Central America and Chiapas at altitudes of 1000-1200 m. (3300- 1000 ft.). The type locality is Juquila in southern Oaxaca and I have seen a female ascribable to this species from Omilteme, Guerrero (also cited by Bates), so it is probable that it occurs in Mexico south of the depression of the Rio Balsas as well. The British Museum has two male specimens labelled “ Venezuela”, but this record is certainly erroneous. Biology. Unknown. Remarks. This species is very closely related to aspericollis Gillet occurring in the same area, minor males and females of the two species being difficult to distinguish (see key). These two species provide a good example of how two sympatric, closely re- lated species may differ radically in male secondary sexual char- acters. The apices of the median pronotal prominences of boucardi are very closely approximated in developed males, those of asperi- collis being quite widely divergent (figs. 73, 74). Its separation from the superficially similar klugi Harold is discussed in the re- marks under the latter species. Material Examined. 13 males, 16 females. MEXICO: Chiapas: Ocosingo, 1200 m., Jim. -Sep. 1917 (M. del Toro), 1 (GIT) • Volcan de Tacana (Coffee belt), 30 Sep. 1956 (V. Aguilar), 1 2 (GH) ; 7500 ft., 4 Apr. 1939 (F. Brodkorb), 1 2 (USNM) ; Guerrero: Omilteme [W of Cliilpancingo] , 8000 ft., Jul. (H. H. Smith), 1 2 (BM). 79 ENTOMOLOGICA AMERICANA GUATEMALA: Alta Vera Paz: Senahu [1000 m.] (Paul Haase), 2 fg (USNM) ; Chimaltenango : Capetillo [Valley between volcanoes Agua and Fuego, 17 Apr. -12 May, 1879] (Champion), 1 ? (BM) ; national record only: (Salle) 1 $ (BM) ; 3 ££ (AMNLI, CAS). EL SALVADOR: Santa Ana: Monte Cristo, 7-9 May, 1958 (0. L. Cartwright), 5 JV?, 9 $? (USNM). C opr is aspericollis Gillet Copris aspericollis Gillet, 1910, Not. Leyden Mus. XXXII : 3 [type: Central America; Musee Royal d’Histoire Naturelle, Brussels] ; Matthews, 1959, Ciencia XIX (6-7) : 136 (key and distr. ) . Description of Male. Head, — Armed. Clypeus with two very low teeth, the margin between them very shallowly emarginate with a deeper median notch not cutting through the margin. Upper surface of head strongly and densely punctate, the punctures shallow and simple, disappearing at extreme base of head, posterior oblicpie carina distinct. Occipital margin with transverse seti- gerous groove complete but somewhat bent at middle of each side. Demarcation between gula and submentum arcuate. Thorax. — Pronotum armed. Anterolateral angles subquadrate in developed specimens, rounded in undeveloped ones, lateral margin slightly inwardly curved behind origin of lateral carina. Lateral carina sharp, almost issuing from margin anteriorly. Anterior margin of pronotum not forming any point or angle medially. Median longitudinal sulcus very faint, almost wanting. Puncturation of pronotum as follows: upper part of disc almost impunctate, the longitudinal sulcus impunctate, rest of upper surface and lateral surfaces punctate, the punctures simple ; anterior declivities densely and coarsely granulate or asperate ; posterior submargin and lateral fossae umbilico-punctate. Anterior prosternal margin with- out any median lobe or with a rounded low one ; sternellum densely punctate. Median lobe of metasternum with numerous punctiform impressions laterally and anteriorly ; median longitudinal im- pressed line complete. Elytra. — 8th stria incomplete, disintegrating posteriorly; 9th stria arising about a third of the way down the elytral length ; 10th complete. Striae on dorsal part of elytra distinctly punctate, the punctures slightly transverse and separated by a distance equal to once or twice their width ; striae 7, 8, and 9 less punctate but the punctures can still be made out. Interstriae very slightly convex, impunctate. Abdomen. — Pygidium moder- 80 VOLUME XLI ately umbilico-punctate, the punctures very small ; pygidial margin usually complete, sometimes effaced at apex. Anterior legs. — Ventral surface of femur coarsely setigerous-punctate on posterior (upper) longitudinal half, more finely punctate on anterior half. Forespur almost straight in dorsal view, tapering slightly to a rounded apex, curved down at apical third. Middle legs. — Coxa impunctate. Femur below finely punctate, with a few, usually setigerous, coarse punctures distally. Tibia below with 2-3 distal median seta tufts. Posterior legs. — Femur like that of middle legs. Tibia with 1-2, usually one, seta tuft distally on ventral surface. Total length. — 21-21.5 mm. Male armament. The head horn is not very tapering but comes to a rather abrupt end, and is noticeably bent in the middle (fig. 74). The median thoracic prominences are slightly divergent and conical, their lower surfaces not flattened, and directed hori- zontally in lateral view. The lateral prominences are laminate, rather acute, and directed forward, their dorsal edges parallel in dorsal view. Description of Female. Similar to male, but differing in armament and in the following features : clypeal teeth more prom- inent, rounded, and approximate, with a V-shaped notch between them. Pronotum more extensively granulate, the granules ex- tending from face over sides and anterior surface of disc, which takes on an asperate appearance. Base of disc finely punctate ; umbilical punctures are confined to posterior submargin and lateral fossae. Pygidium more finely punctate and always completely margined. Total length. — 21-23.5 mm. Female armament. Normal for the genus. Distribution. Known only from the area around Guatemala City at altitudes of 340-1700 m. (1100-5600 ft.). Biology. Feeds on cow dung. Some aspects of nidification are discussed on p. 31. Remarks. This species is poorly represented in collections, probably reflecting the generally uncollected nature of Central America rather than any rarity of the species. I collected it in a cow pasture on the road southeast from Guatemala City, indicating that it probably is abundant in central Guatemala, but I did not en- counter it again elsewhere in Central America. It is most closely related to boucardi Harold (see remarks under that species) and to subpunctatus Gillet, which it strongly resembles in habitus, being however much more strongly punctate than the latter species. It also differs from the latter in the shape of the male median pronotal prominences, which are conical in aspericollis and flattened at the apices in subpunctatus. 81 ENTOMOLOGICA AMERICANA Material Examined. Seven males, seven females. GUATEMALA : Escuintla: Escuintla [338 m., Feb. or Apr., 1881] (Champion), 1 J', 1 J (BM) ; Guatemala: 18 km. SE Guate- mala City, 610 m., 22 Jul., 1958 (Neff and Matthews), 1 3 52 (EGM) ; S. Jose Pinula [1500-2000 m.] May, 1924 (W. M. Mann), 1 ■<$ (USNM) ; national record only: 1 3 22 (USNM) ; 10 Jun. 1943, 1 J (AMNH) ; 9 Mar. 1924, 1 J (USNM) ; 15 May, 1924, 1 J (USNM) ; location undetermined: Azahar de Carboga (Undqr- wood), 1 J1, 1 2 (CM) ; national record only: 1897 (Pittier), 2 J'J' (USNM). PANAMA: Chiriqui: Potrerillos, 1 2 (CAS); Volcan de Chiriqui, 2500-4000 ft. [6-8 Jun. 1882] (Champion), l J (BM). Copris costaricensis dolichocerus n. subsp. Holotype: Volcan de Tacana, Chiapas, Mexico, 1 Oct. 1956 (V. Aguilar), J'; United States National Museum. Description. Differs from the typical subspecies only in the greater proportional length of the male cephalic horn. When the height of the male horn is plotted against the length of the hind femur on a graph (fig. 7), the specimens of this species are seen to fall into two lots and to follow different curves, corresponding to their geographical origin. Distribution. Fig. 17. Chiapas and Guatemala at 1500-2000 m. I have ascribed the two known Guatemalan females to this subspecies purely on the basis of geographical distribution, as the females are not separable from the Costa Rican subspecies. 97 ENTOMOLOGICA AMERICANA Biology. Unknown. Remarks. This form is recognized on the basis of the same character distinguishing the subspecies of klugi and lecontei. Dif- ferent horn-length allometric relationships in this genus suggest a fundamental (perhaps genetic) difference which is sometimes (but not in this case) reflected in other morphological characters. Material Examined. Three males, seven females. MEXICO: Chiapas: El Verjel, 6 Oct. 1939 (C. Bolivar), 3 ?$ (GH) ; Volcan de Tacana (Coffe Belt), 1 Oct. 1956 (Y. Aguilar), 3 JV?, 2 ?? (USNM, GH). GUATEMALA: Quezaltenango : San Isidro, 1500 m. [10-23 Sep. 1880] (Champion), 1 $? (BM) ; Quiche: Santa Cruz del Quiche, 13 Aug. 1947 (C. and P. Vaurie), 1 $ (AMNII). Complex 4. The rebouchei complex. Posterior angles of head quadrate. Anterolateral angles of pronotum acute, the margin behind them sharply sinuate. Median longitudinal sulcus of pronotum impunctate or very finely punctate. Forespur with the apex acute and curved inward in both sexes. Median coxae devoid of coarse punctures. 8th elytral stria com- plete. Pygidial margin complete or not. Complex punctures usually minutely granulate. Three species found at low to moderate altitudes in Central and Western Mexico and bordering United States territory: lecontei n. sp., rebouchei Harold, and halffteri Matthews. Copris lecontei lecontei new species Holotype: Huachuca Mts., Arizona, United States Na- tional Museum. Description of Male. Head. — Armed. Clypeus bidentate, the teeth not at all prominent, very obtuse, their inner edges meet- ing at a very broad angle forming the median emargination ; no median notch. Upper surface of head densely punctate except for area between eyes, which is sparsely and very finely punctate or impunctate; all punctures simple. Posterior oblique carina sharp. Occipital margin with transverse setigerous groove broken into three sections which do not overlap. Demarcation between gula and submentum arcuate. Thorax . — Pronotum armed. Antero- lateral angles with a salient point followed by a sharp inward curve of the margin, which is then angled out again at origin of lateral 98 VOLUME XLI carina. Lateral carina sharp and prominent. Anterior margin of pronotum not forming any median point or angle. Median longitudi- nal sulcus deeply impressed, with a few punctures or impunctate. Puncturation of pronotum as follows: densely punctate over all dorsal and lateral surfaces, the punctures smaller and less dense, or sometimes absent, on disc, becoming more sparsely punctate on anterior declivities ; punctures simple except for those of the follow- ing areas, which are coarse and umbilical : along entire submargin, a few along median longitudinal sulcus and median line of anterior face, in lateral fossae, and hollows between prominences. Anterior prosternal margin with a low, truncate median tooth ; sternellum moderately umbilico-punctate. Median lobe of metasternum with shallow umbilical punctures along sides and anteriorly; median longitudinal groove complete to anterior pit. Elytra. — 8th stria complete ; 9th arising about one third of way down elytron ; often a more anterior segment of 9th stria is present, not quite issuing from base or joining posterior section and not punctate ; 10th stria complete. Striae coarsely crenato-punctate, the punctures very little deeper than the striae, round, separated by a distance equal to their diameters or less. Interstriae very slightly convex, sparsely, finely punctate, appearing smooth, or quite impunctate. Abdomen. — Pygidium rather densely, coarsely umbilico-punctate, the margin complete or rarely incomplete, the inner edge of margin being effacted ventrally (fig. 59). Anterior legs. — Ventral surface of femur with coarse setigerous punctures on posterior longitudinal half, impunctate on anterior half. Porespur straight and parallel- sided to apical third, where it takes a sharp bend inwards and tapers to a sharp point. Middle legs. — Coxa impunctate. Ventral surface of femur very sparsely and finely punctate or impunctate except distally, where the punctures are coarse and setigerous. Tibia below with three distal seta tufts. Posterior legs. — Like middle legs, but tibia below with a row of 4—5 seta tufts. Total length. — 10-15.5 mm. Male armament. Medium specimens bear a horn which is prac- tically straight, the rearward curvature being barely perceptible (fig. 75). The horn of major specimens is gently arcuate and slightly expanded transversely at the apex. The pronotum bears four prominences as usual, the two median ones compressed, slightly upwardly directed, and approximated, but always with a punctate depression between them. Description of Female. Similar to male, but differing in ar- mament and in the following features : usually some umbilical punctures between eyes on upper surface of head ; puncturation on 99 ENTOMOLOGICA AMERICANA Fig. 18. Distribution of the rebouchei complex. Base map reproduced by permission of the University of Chicago. 100 VOLUME XLI anterior part of disc much denser and merging, the punctures umbilical only along submargins and in lateral fossae. Total length. — 11-15 mm. Female armament. Typical for the genus. Distribution. Fig. 18. Known from practically all the moun- tain ranges of southeast Arizona at altitudes of 2500 to 6000 ft. (760-1800 m.) but found most frequently at about 4000 ft. (1200 m.). There is one record each from the Animas Mts. of New Mexico and the Rio Mayo in Chihuahua and there are several records from the Sierra de Alamos in southern Sonora at altitudes of but a few hundred metres. Thence there are isolated records down the west coast of Mexico to Nayarit at altitudes of sea level to 1500 m. (4900 ft.). Biology. Unknown. In Arizona it is apparently most abund- ant in late summer. Remarks. The relationship of this species to rehouchei Harold is extremely close. This cannot be seen in the Arizona specimens, which are very different from rehouchei, being much more heavily punctate on the pronotum and the males bearing median pronotal prominences of a different type (figs. 75, 76). However, as we examine lecontei further south in Sinaloa and Nayarit we find it becomes increasingly less punctate and the median pronotal promi- nences become more laminate and remote, approaching both re- houchei and 1. isthmiensis in form. C. rehouchei possesses an acute tooth on the prosternal margin (fig. 54), whereas in lecontei there is at most a rounded expansion or truncate process here, and in rehouchei the pygidial margin is always quite effaced medially, whereas in lecontei it is nearly always either engraved for its entire length or continued medially by a close-set row of punctures (fig. 59). This species has been long familiar under the name moechus Leconte. A recent examination of the type of moechus revealed it to belong to a Mexican species hitherto bearing the name clavicornis Matthews and Halffter. As there were no synonyms available for this common species, it is here described as new and dedicated to the distinguished American systematist, J. L. Leconte. Material Examined. 244 males, 249 females. UNITED STATES : Arizona: Cochise Co. : Carr Cyn, Huachuca Mts. ; Lower Carr Cyn, Huachuca Mts. ; Chiricahua Mts. ; Huachuca Mts.; Mule Mts., Gilman Ranch, 8 mi. N Bisbee; Palmerly; 3 mi. E Portal; Ramsay Cn., Huachuca Mts.; San Bernardino Ranch; Texas Pass, Dragoon Mts. ; Webb ; SE end Whetstone Mts., Sands Ranch ; Gila Co. : Globe ; Base Pinal Mts. ; Rice ; Pima Co. : Babo- quivari Mts. ; Baboquivari Mts., Kitts Peak Rincon ; Baboquivari 101 ENTOMOLOGICA AMERICANA Mts., El Mirador Rcli., 4 mi. NW Sasabe ; Baboquivari Canyon, W. side Baboquivari Mts. ; Brown ’s Canyon, E side Baboquivari Mts. ; Elkhorn Ranch, E slope N end Baboquivari Mts. ; Sycamore Can- yon, N of Baboquivari Canyon, W side Baboquivari Mts. ; Con- tinental; Santa Catalina Mts.; Santa Rita Mts.; Santa Rita Mts., Old Parker Ranch ; Sierritas, Black Dike Prospect ; Tucson ; Tucson, St. Xavier Mission; Santa Cruz Co.: Elgin; Nogales; Patagonia; Ruby; 21 mi. SE Ruby; Sonoita R., Patagonia; 25 mi. E Sonoita; Tumacacori Nat’l. Mon.; Yanks Spring, 4 mi. SE Ruby, Pajaritos Mts.; Washington Mts., nr. Nogales; Yavapai Co.: Prescott; New Mexico: Hidalgo Co. : Animas Mts., Double Adobe Ranch. MEXICO: Chihuahua: Rio Mayo, Aug. (Gentry), 1 (CAS) ; Nayarit: Tepic, 28 Jul. 1953 (C. and P. Vaurie), 2 J'J1, 2 22 (AMNH) ; 27 Jul. 1954 (M. Cazier, W. Gertsch, Bradts), 1 2 (AMNH) ; Sinaloa: Culiacan, 250 ft., 21 Jul. 1959 (H. E. Evans), 2 2 2? (CU) ; Mazatlan, 15-17 Sep. 1918, 1 J1, 1 ? (USNM) ; 9 mi. N Mazatlan, 100 ft., 12-16 Jun. 1953 (R. K. Selander), 1 ?; El Venadio [4.5 mi. N Mazatlan]*, 16 Jun. 1918 (J. A. Kusche), 41 jy1, 35 (USNM, CAS) ; Sonora: Agua Marina nr. Alamos, 20 Jul. 1955 (R. and 0. Flint), 1 £ (EGM) ; Alamos, 29 Jul. 1940 (R. P. Allen), 1 (L.). Busck (1906) gives the following discussion of the larva (in turn the notes originated with Miss Murtfeldt) : “Feeds on rickweed (sic) ( Pilea pumila ), a succulent little plant of the nettle family, with adhesive, but not urticating leaves ; mining, twisting, and crumpling them. “Larva yellowish white, subcylindrical. “Head shining black, with the diversions defined by narrow white lines. Cervical shield broad, oblong, with fine white central line. The sutures are very deep, giving the larva a moniliform aspect. “These larvae are not confined to one mine, but may be seen wandering over the leaves and stems, cutting in between the two cuticles of a leaf and covering it with transparent spots of various sizes. They change to pupa under a fold of the leaf or between the wrinkles or not infrequently on the surface of the ground, pro- tected by a very slight dingy cocoon. There seems to be several broods in a season.” Type : Museum of Comparative Zoology. Type locality : Covington, Kentucky. Specimens examined: 90 20 §. Arizona: Madera Canyon, 5,600 feet, Santa Rita Mountains, Santa Cruz Co., 4 J', 2 J, Septem- ber 12, 1959 (R. W. Hodges), [CU, RWII] . Florida: Siesta Key, 21 ENTOMOLOGICA AMERICANA Sarasota Co., 27 4 J, January 28 through October 24 (C. P. Kimball), [CPK, RWH] ; Homestead, 1 April 15, 1959 (D. 0. Wolfenbarger), [CPK]. Georgia: Clarke Co., 5 April 5, 1924 (Richards), [ABK]. Illinois: Putnam Co., 12 3 J, April 24 through September 5 (M. 0. Glenn), [MOG] ; Arlington Hts., 1 J', August 12, 1949 (A. L. McElhose), [CNHM]. Kansas: Onaga, 1 c??, “8/5 02” [MCZ] . Kentucky: L. Sandy R., Ellicott Co., 1 May 3, 1936 (Annette F. Braun), [AFB], Minnesota: no local- ity, 1 J', August 15, 1903 [CU]. Missouri: Kirkwood, 4 1 J, June 1 through July 29 (Murtfeldt), [CU] ; no locality, 3 July 13 and 19 [USNM] ; Columbia, 2 [AMNH] . New York: Mon- roe Co. 4 J', July 30 through August 13, 1949 (C. P. Kimball), [CPK] ; Pelham, Westchester Co., 1 under rearing record B. 486, emerged August 16, 1913 (Annette F. Braun), [USNM] ; Conkle ’s Hoi., Hocking Co., 1 on Panicum clandestinum, emerged April 2, 1944 (Annette F. Braun), [AFB] ; Scioto Co., 1 2, 011 Panicum, emerged July 26, 1948 (Annette F. Braun), [AFB]. Virginia: Falls Church, 1 June 2, 1914 (Heinrich), [USNM]. This species is closest to C. pulchrimella but may be easily dis- tinguished from it by the absence of white markings on the thorax and the presence of broad silver-blue lines in the basal area of the forewdngs. Cosmopterix montisella Chambers (Figs. 27, 101, 141) Cosmopteryx montisella Chambers, 1875. Cincinnati Quart. Jour. Sci., 2 : 297. Chambers, 1878. Bull. U.S. Geol. Geog. Surv. Terr., 4:137. Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 535. McDunnough, 1939. Mem. S. California Acad. Sci., 2 : 63. 27 ENTOMOLOGICA AMERICANA Cosmopteryx monticella (sic) Busck, 1906. Proc. U. S. Natl. Mns., 30: 712. Cosmopteryx unicolorella Walsingham, 1889. Ins. Life, 1 : 291. new synonymy. Dyar, 1902 [1903] . List of the Lepidoptera of North America, Bull. U. S. Natl. Mils., 52:535. Busck, 1906. Proc. U. S. Natl. Mus, 30:710. McDunnough, 1939. Mem. S. California Acad. Sci., 2 : 63. Tongue shining buff. Labial palpi brown ; second segment with three white lines : one ventral, one internal, and one external ; third segment with an anterior and a posterior white line. Antennae brown ; pecten present ; ventral surface of scape white ; an anterior white line on scape, continued on shaft as a series of dots to one- half or farther ; apical four segments : three brown, one white ; or, two brown, two white ; or, one-half brown, three-and-one-half white. Apical four segments preceded by five brown, one white, one brown, two white, then brown ones. Face olive-brown with purple reflec- tions; vertex dark olive-brown with three white lines, one above each eye and one medial. Thorax dark olive-brown with continua- tion of three white lines on vertex. Forewings dark olive-brown basally and apically ; four white lines in basal area ; a yellow-orange fascia at one-half ; a silver-gray fascia bordering it inwardly ; out- wardly, yellow-orange fascia bordered by two silver-gray spots, one dorsal and one subcostal; subcostal spot offset apically from dorsal one ; a white costal streak above subcostal spot ; a broad white streak at apex of wing; a few silver-blue scales on dorsal margin midway between apex and dorsal spot. Hindwings fuscous. Ab- domen brown and buff dorsally, brown with apices of segments silver ventrally. Legs silver-gray with purple reflections basally, brown distally ; metathoracic tibiae with a dorsal line from base to one-half ; a white annulation at two-thirds and one at apex ; apices of tarsal segments buff to white. Male genitalia: (Fig. 27) R.W.H. slide no. 537. Female genitalia: (Fig. 101) R.W.H. slide no. 538. Alar expanse : 10-13 mm. Food plant : unknown. Type : montisella, Museum of Comparative Zoology ; unicolorella, British Museum (Natural History). Type locality : montisella, Spanish Bar, Colorado ; unicolorella, Siskiyou Co., California. Specimens examined : 32 17 J. Arizona : Madera Canyon, 4,880 and 5,600 feet, Santa Rita Mountains, Santa Cruz Co., 9 9J, July 9 through September 29, 1959 (R. W. Hodges), [CU, 28 VOLUME XLII RWH] ; same data except 4,400 feet, Pima Co., 1 2, October 6. 1959 [RWH] ; Santa Catalina Mountains, Pima Co., 1 August 1, 1938 (Bryant), [LACM] ; Palmerlee, 1 , J1 [USNM]. California: Mt. Shasta City, Siskiyou Co., 9 July 2-18, 1958 (J. Powell), [UCB, RWH] ; Davis Creek, Modoc Co., 1 J, July 11, 1957 (J. Powell), [RWH] ; Bear Creek, Shasta Co., 3 , under rearing record B. 1643, emerged July 28, 1938 (Annette F. Braun), [AFB]. Connecticut: East River, 1 July 31, 1908 (Chas. R. Ely). [USNM]. Kentucky: Cumberland Falls, 2 1 5, under rearing record B. 1315, emerged July 12, 1932 (Annette F. Braun), [AFB, RWH] ; Cumberland Valley, Letcher Co., 1 J', June 27, 1933 (Annette F. Braun), [AFB] ; Mammoth Cave, 1 $ , If, under rearing record B. 1315, emerged September 9 and 11, 1941 (Annette F. Braun), [AFB] ; Yahoo Creek, McCreary Co., 1 lCf, 1 J, under rearing record B. 1315, emerged July 5, 1935, collected June 18, 1935 (Annette F. Braun), [AFB]. Ohio: Beaver Pond, Adams Co., 1 5, under rearing record B. 1315, emerged August 15, 1927 (Annette F. Braun), [AFB], South Carolina: Clemson College, 1 J, leaf miner on legume, emerged August 27, 1933 (W. C. Nettles), [USNM]. C. lespedezae can be distinguished from C. attenuatella by the apex of the antennae being white in lespedezae, brown in atten- uatella. The markings place lespedezae closest to C. abdita ; how- ever, the characters in the keys to the male and female genitalia will suffice to separate the two species. Apparently, Walsingham did not have complete notes on lespedezae when he synonymized it with attenuatella because the differences in the antennal markings and the maculation of the forewings are more than adequate to separate lespedezae as a dis- crete species. Even though the distribution records are not com- plete, lespedezae appears to have a more northern range than does attenuatella. By Walsingham ’s indicating that attenuatella equaled lespedezae, an error in the identity of attenuatella occurred. Busck (1906) gave the characters of the type of lespedezae for attenuatella, and this definition was followed by subsequent workers. Mr. Bradley has made it possible for me to identify attenuatella by furnishing me with a sketch of the male genitalia. The type of genitalia is so characteristic of this species that there can be no doubt as to it identity. 36 VOLUME XLII Cosmopterix opulenta Braun (Figs. 37, 104, 148) Cosmopteryx opulenta Braun, 1919. Ent. News, 30:260. McDun- nough, 1939. Mem. S. California Acad. Sci., 2 : 63. The description is quoted from Braun’s paper (1919). “Palpi white, outer and inner surfaces each marked with a black longitudinal line. Antennae grayish brown, becoming darker towards apex; with a conspicuous white line on anterior surface near base ; last segment black, or sometimes merely black-tipped ; next three preceding segments white ; next three segments black ; followed by a white, then a black, then a white segment [pecten present]. Head and thorax grayish brown, with three longitudinal white lines. “Forewings brownish gray, or seal brown, with the basal half marked with five fine white longitudinal lines; one starting from base just within the costal edge diverges from the costa outwardly, extending about two-thirds through the basal brown area ; a second extends along the extreme costa from the basal fourth almost or quite to the yellow fascia, becoming broader outwardly; a third along middle of wing from base to a little beyond the costo-basal streak ; a fourth shorter streak below fold not attaining the base ; a fifth white streak dorso-basal. Just beyond middle of wing, a yellow fascia. Four patches of metallic scales ; the costal one of the inner pair limits the fascia inwardly, not touching the costa, and has a few black scales on its outer margin ; dorsal patch placed farther back and with black scales on its inner margin ; the yellow fascia extends between them and borders the inner side of the dorsal patch, sometimes almost to dorsal margin. Posterior pair of me- tallic patches almost opposite, attaining the margins, and limiting the yellow fascia outwardly, except in the middle of the wing where the fascia extends between and a little beyond them. Costal cilia immediately following the second costal metallic patch white ; occasionally the yellow of the fascia is almost confluent with this white patch. Remainder of apical portion of wing and cilia of the basal brown ground color, except for a long white line extend- ing from just beyond the yellow fascia to the tips of the apical cilia. Hind wings and cilia concolorous with fore wings. Legs gray streaked and banded with white. [Metathoracic tibiae with a longitudinal blue-white line extending from base to two-thirds or nearly to apex ; this line moving from ventral surface to dorsal surface; apex blue-white.] 37 ENTOMOLOGICA AMERICANA Male genitalia: (Fig. 37) R.W.H. slide no. 552. v Female genitalia: (Fig. 104) R.W.H. slide no. 539. Alar expanse : 7%-9 mm. Food plant: Ambrosia psylostachya DC. Brann (1919) gives the following description of the larval mine : ‘ ‘ The mines extend principally along the midrib, with irregular proections branching out on either side. The larva spins a cocoon on the densely pubes- cent under side of the leaf, constructed of silk, and the whitish pu- bescence of the leaf. ” Type : Annette F. Braun Collection. Type locality: Rivera, Los Angeles County, California. Specimens examined : 10 J', 2 J. Arizona : Madera Canyon, 4,400 feet, Santa Rita Mountains, Pima Co., 1 J', October 12, 1959 (R. W. Hodges), [RWH]. California: Rivera, Los Angeles Co., 2,c?, 1 under rearing record B. 589, emerged April 17, 1910 (Annette F. Braun), [AFB]. Oklahoma: Oklahoma City, 8 J*, July 27 through September 4 (D. R. Davis), [DRD, RWH]. C. opulenta can be separated from C. quadrilineella by the ab- sence of lateral thoracic lines. The presence of a single signum in the female genitalia serves to separate it from the other species of Cosmopterix. Specimens from Arizona and Oklahoma are decidedly darker brown than those from California. In part this may be a result of fading of the California specimens ; however, when the descrip- tion was made, the specimens were relatively fresh. Cosmopterix quadrilineella Chambers (Figs. 38, 94, 149) Cosmopteryx quadrilineella Chambers, 1878. Bull. U. S. Geol. Surv. Terr., 4: 95. Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 535. Busck, 1906. Proc. U. S. Natl. Mus., 30:710. McDunnough, 1939. Mem. S. California Acad. Sci., 2 : 63. Tongue white to light buff. Labial palpi gray-brown ; second segment with three white lines : one ventral, one on inner surface, and one on outer surface ; third segment with an anterior and a posterior white line ; apex white. Face buff ventrally becoming gray -brown dorsally; vertex gray -brown with three white lines, one over each eye and one medial. Antennae brown ; pecten present ; ventral surface of scape white ; an anterior white line continued on shaft to one-half ; apical two segments brown, pre- 38 VOLUME XLII ceded by two white, five brown, one white, four brown, one white, then brown segments. Thorax brown with continuation of three white lines on vertex. Forewings with basal third and apical area dark gray-brown ; four white lines in basal area ; a yellow fascia from one-third to two-thirds with a medial extension to three- fourths ; four silver patches in fascia ; outer costal silver patch fol- lowed by a white streak in cilia ; a white line from end of fascia to apex ; cilia gray -brown. Hindwings fuscous. Abdomen brown with purple reflections dorsally, becoming ochreous ventrally. Legs brown ; prothoracic legs with a white line from femur to apex of tarsus ; tibiae with three oblique stripes ; basal, medial, and apical ; tarsi brown with a dorsal white line ; metathoracic tibiae with a basal and a medial oblique white line ; apex and spurs white ; tarsi with a dorsal white line. Male genitalia: (Fig. 38) R.W.TI. slide no. 544. Female genitalia: (Fig. 94) R.W.TI. slide no. 848. Alar expanse : 7-9 mm. Food plant : unknown. Type : Museum of Comparative Zoology. Type locality: Bosque Co., Texas. Specimens examined : 11 (Cf, 21 §. Arizona : Madera Canyon, 4,880 and 5,600 feet, Santa Rita Mountains, Santa Cruz Co., 10 20J, July 10 through September 24, 1959 (R. W. Hodges), [CU, RWH] ; same date except, Pima Co., 4,400 feet, 1 J', 12, October 10 and 26, 1959 [RWH] . C. quadrilineella can be separated from the other species of Cosmopterix by the fascia surrounding both of the basal silver spots. Cosmopterix minutella Beutenmueller (Figs. 31, 150) Cosmopteryx minutella Beutenmueller, 1889. Ent. Americana, 5:10. Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 535. Busck, 1906. Proc. U. S. Natl. Mus., 30:711. McDunnough, 1939. Mem. S. Cali- fornia Acad. Sci., 2 : 63. Tongue shining pale buff-yellow. Labial palpi white, a dorsal brown line on second segment, an internal and an external brown line on third segment. Antennae olive-brown ; pecten present ; scape with ventral surface and an anterior line white ; anterior line continued on shaft to one-third or one-half ; apical four seg- ments white, preceded by three brown, one white, one brown, one 39 ENTOMOLOGICA AMERICANA white, and one brown segment. Face shining gray-broAvn; vertex olive-brown with three white lines, one above each eye and one medial. Thorax olive-brown with continuation of three white lines. Forewings with basal area olive-brown and with four white lines ; fascia pale yellow, bordered by four silver spots ; apical area buff-gray with a white line along dorsal margin. Hindwings pale fuscous. Abdomen ochreous dorsally, shining buff ventrally. Legs shining buff basally; metathoracic tibiae olive -brown with three white lines : one from base to one-third, one from base to three-fourths, and one from two-thirds to near apex; metathoracic tarsi with a brown annulation basally followed by a broad white annulation, then buff -brown and buff ; segments two through five brown ventrally, buff dorsally. Male genitalia: (Fig. 31) R.W.H. slide no. 893. Female genitalia : no specimens available. Alar expanse : 7-8 mm. Food plant: unknown. Type : United States National Museum. Type locality: Central Florida. Specimens examined : 6 ,J\ Florida : Archbold Biological Sta- tion, Highlands Co., 1 lCf , March 29, 1959 (R. W. Hodges), [RWH] ; Everglades, 1 J1, April 8-15 [USNM] ; Homestead, 3 lCf, May 1 and November 2, 1959 (D. 0. Wolfenbarger), [CPK] ; Lakeland, 1 under rearing record B. 2105, emerged July 10, 1946 (Annette F. Braun), [AFB] ; Ithaca, Tompkins Co., 1 a)s = love, d7ea = warmth (Figs. 3, 18, 51, 110) Type: Eriphia albalineella Chambers, 1878. Head smooth-scaled ; tongue scaled, well developed ; maxillary 59 ENTOMOLOGICA AMERICANA palpi folded over base of tongue ; labial palpi smooth-scaled, re- curved, not reaching apex, second segment longer than third, apex acute ; antennae two-thirds to three-fourths, pecten present, scape elongate and slightly swollen distally. Dorsal surface of meta- thoracic tibiae with long scales. Forewings lanceolate, apex acute; 12 veins present; lb furcate basally; 2 and 3 before end of cell ; 6, 7, and 8 stalked, 8 weak; 10 opposite 2; 11 from middle of cell. Hindwings linear; a tuft of scales on costal margin ; 8 veins present ; lb simple ; 1, 2, 3, and 4 evanescent ; 6 and 7 stalked, divergent. Male genitalia : valvae lobate, rounded apically ; costae separate, short, lobate ; tegumen heavily sclerotized ; uncus and socii absent ; arms of gnatlios separate, asymmetrical, left brachium longer than right one ; aedeagus short, cylindrical, a basal process from dorsal surface, ankylosed. Female genitalia : bursa copulatrix membranous ; two signa, cornucopia shaped ; lamella antevaginalis sclerotized, forming a plate in front of ostium ; apophyses moderately short. Eralea differs from Stagmatophora in the following characters : the second segment of the labial palpi of Eralea is longer than the third; in Stagmatophora the third segment is longer than the sec- ond. The second segment of the labial palpi of Eralea is not thickened ventrally ; that of Stagmatophora is thickened ventrally. The scales on the dorsal surface of the metathoracic tibiae of Eralea are long and thin ; those of Stagmatophora are normal. In the hindwings veins 1 through 5 are evanescent in Eralea ; these veins are well developed in Stagmatophora. In the male genitalia of Eralea the costae are symmetrical and not fused with the anellus ; those of Stagmatophora are asymmetrical and fused with the anellus. The female genitalia of the two genera are very similar, indicating their close relationship. Two species of Eralea are known from the southern United States, one from Florida and one from Texas and Arizona. Future collecting will probably extend the known ranges of these species. Key to the Species of Eralea based upon the Maculation 1. Vertex dark brown striata Vertex pale buff -white albalineella Key to the Species of Eralea based upon the Male Genitalia 1. Ventral margin of costa concave striata Ventral margin of costa almost straight albalineella 60 VOLUME XLII Eralea albalineella Chambers, new combination (Figs. 51, 110, 163) Eriphia (?) albalineella Chambers, 1878. Bull. U. S. Geol. Geog. Surv. Terr., 4 : 95. Eriphia albalineella , Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 540. Tanygona albalineella, Braun, 1923. Trans. American Ent. Soe., 49 : 115. McDunnough, 1939. Mem. S. California Acad. Sci., 2:63. Tongue white to pale gray. Labial palpi brown with a dorsal and a ventral buff line on second and third segments. Antennae buff, scape with a broad dorsoposterior brown line continued on shaft to one-third, a faint brown line on anterior surface of scape and basal part of shaft. Face and vertex buff-white, brown an- terior to and dorsad of eyes, brown along cervical margin. Thorax brown with three blue-white lines, two lateral and one medial. Forewings brown-black ; an oblique buff-white line from costal margin at base to dorsal margin at two thirds; a second starting from costal margin at one-half to middle of wing at three-fourths ; and a short one in middle of wing at five-sixths ; costal cilia buff ; apical cilia buff basally, brown distally ; dorsal cilia buff with some brown flecks. Ilindwings buff, cilia gray. Legs shining buff ; outer surface of metathoracie tibiae with a broad gray-brown streak from base to one-half, another from one-half to nine-tenths ; meta- thoracic tarsi gray-brown on dorsal surface of first segment and basal half of second. Male genitalia: (Fig. 51) R.W.H. slide no. 868. Female genitalia: (Fig. 110) R.W.II. slide no. 869. Alar expanse : 8-9 mm. Food plant : Unknown. Type : Museum of Comparative Zoology. Type locality: Bosque Co., Texas. Specimens examined : 1 J1, 2 §. Arizona : Chiricahua Mts. near Portal, Cochise Co., 1 J1, 1 J, July 5, 1939 (Annette F. Braun), [AFB]. Texas: [Bosque Co.], 1 ?, “3/3” [MCZ]. The buff-white line from the base of the forewings of albalineella is narrower than the one of E. striata. This character, in com- bination with the pale vertex, will separate albalineella from striata. 61 ENTOMOLOGICA AMERICANA Eralea striata, new species (Figs. 50, 165) Tongue white basally, gray distally. Labial palpi brown with a ventral white line on second and third segments. Antennae buff- white, scape brown except for an anterior white line, pecten brown, base of shaft pale brown. Face gray-brown with lavender reflec- tions ; two oblique white streaks, one from base of each antenna to base of tongue ; vertex brown. Thorax brown ; patagia white on inner and outer margins. Forewings buff-white to white ; a broad white band from base of wing to dorsal margin at one-half ; an oblique white line near base running from near costal margin to medial white band ; a row of brown scales at apex of wing ; cilia with pale brown flecks basally, buff -white distally. Hindwings pale fuscous. Legs buff-white ; prothoracic legs with an anterior brown line ; metathoracic tibiae with a brown streak from base to one-half, another from two-thirds to nine-tenths; metathoracic tibiae with brown on dorsal surface of first, second, and third segments. Male genitalia: (Fig. 50) R.W.H. slide no 925. Female genitalia : No specimens available. Alar expanse : 8 mm. Food plant : Unknown. Holotype : .J', Siesta Key, Sarasota, Co., Florida, May 13, 1960 (C. P. Kimball), [CU Type No. 3812]. Paratype: Oneco, Manatee Co., Florida, 1 May 19, 1953 (Paula Dillman), [CPK]. Melanocinclis, new genus .ueXav = black pigment, kwkH* = latticed gate (Figs. 8, 15, 57, 116) Type : Melanocinclis lineigera , sp. non. Head smooth-scaled; tongue scaled, well developed; maxillary palpi minute; labial palpi recurved, not reaching vertex, second segment longer than third and slightly thickened ventrally, third segment rough-scaled, apex acute ; antennae three-fourths, pecten present, simple, scape long and broad. Metathoracic tibiae with long hairs on dorsal surface. Forewings lanceolate, acuminate ; 10 veins present ; 2 and 4 absent; lb furcate basally; 3 before end of cell, not reaching margin ; 6, 7, and 8 stalked ; 9 from end of cell ; 10 before 3 ; 11 from middle of cell. Hindwings linear; a series of scales on costal 62 VOLUME XLII margin ; 4 veins present ; 2, 3, 5, and 6 absent ; lb simple. Male genitalia: valvae linear, expanded distally; costae sepa- rate, fused to anellus; anellus heavily sclerotized; aedeagus anky- losed, cornuti absent ; tegumen heavily sclerotized ; uncus and socii absent ; brachia broad basally, aciculate distally, subequal in length. Female genitalia : bursa copulatrix membranous ; signa absent ; lamellae vaginales forming an irregularly shaped tube ; apophyses long. Melanocinclis differs from Tanygona in the following characters : The third segment of the labial palpi is rough-scaled and shorter than the second segment ; that of Taviygona is smooth and longer than the second segment. Veins 2 and 4 of the forewing of Melanocinclis are absent, whereas these veins are present in Tany- gona. Vein 6 is absent in the hindwing of Melanocinclis ; it is present in Tanygona. In the male genitalia the brachia of Mela- nocinclis are very slender apically; those of Tanygona are shorter and stouter. The configuration of the lamellae vaginales of Mela- nocinclis is irregular; the lamellae vaginales of Tanygona form a straight tube. The paucity of specimens of the known species of Melanocinclis make it difficult to give an accurate statement as to the geographic range of the genus. M. lineigera occurs in Florida, and M. ni >■ grilineella occurs in Texas and Arizona. I am placing E. nigrilineela Chambers in Melanocinclis because the general facies agree very well with the type of the genus. Chambers (1878) was quite correct in questioning the generic placement of nigrilineella. Key to the Species of Melanocinclis BASED UPON THE MACULATION 1. Forewings dusted with tawny flecks lineigera Forewings dusted with black flecks nigrilineella Key to the Species of Melanocinclis BASED UPON THE MALE GENITALIA 1. Right brachium with a lateral process (Fig. 57) lineigera Right brachium without a lateral process (Fig. 56) nigrilineella 63 ENTOMOLOGICA AMERICANA Melanocinclis lineigera, new species (Figs. 57, 116, 168) Tongue white. Labial palpi white ; basal half of second seg- ment black, a few ventral black scales before apex ; third segment with a black annulation at two-fifths and one at four -fifths ; apex black. Antennae with pecten black ; scape white with a black saddle on posterior surface, apex white ; apical segment fuscous, preceded by one-and-one-half buff-white, one-and-one-half fuscous, one buff-white, one-half fuscous, one buff-white, one-and-one-half fuscous, two buff -white, four-and-one-half fuscous, etc. Face and vertex white, lower half of face dark gray with purple re- flections, a few black flecks above eyes. Thorax white medially, black laterally, patagia white with tawny flecks laterally. Fore- wings white overlaid with tawny flecks ; a broad oblique black band from base at costal margin to one-half at center ; a second black band starting at one-half on costa running to apex of wing below costa; black dorsad of fold from base of wing to one-fourth, some- times reaching margin ; cilia with a black medial line and a fuscous terminal line from apex to a point parallel with dorsal margin. Hindwings shining gray. Abdomen brown-black with ochreous scales dorsally, buff ventrally. Legs shining white ; metathoracic tibiae with an oblique black stripe at four-fifths, dorsal surfaces of spurs black, under certain light conditions ; metathoracic tarsi with a broad black annulation on middle of first segment, black at apex of second segment and at base of third segment. Male genitalia: (Fig. 57) R.W.H. slide no. 919. Female genitalia: (Fig. 116) R.W.H. slide no. 781. Alar expanse : 5%— 1 6% mm. Food plant : Firms caribaea Morelet. Holotype : 5, Archbold Biological Station, Highlands Co., Florida, March 31, 1959 (R. W. Hodges), [CU Type No. 3813]. Paratypes: same locality as type, 3 March 29 and April 4, 1959 [RWH] : Homestead, Florida, 2 February 9 and March 21, 1959 (D. 0. Wolfenbarger), [CPK] ; Lake City,' Florida, 1 12, reared ex Slash Pine cones, emerged February 7, 1951 (C. F. Spears), [USNM] ; Siesta Key, Sarasota Co., Florida, 5^, 5$, March 31 through May 3, 1960 (C. P. Kimball), [CPK, RWH]. Other specimens examined : Homestead, Florida, 1 J', 2 February 8 and April 16, 1959 (D. 0. Wolfenbarger), [CPK] ; Siesta Key, Sarasota Co., Florida, 3 2 J, February 25 through May 13 (C. P. Kimball), [CPK, RWH]. The tawny flecks on the forewings will separate lineigera from M. nigrilineella. 64 VOLUME XLII Melanocinclis nigrilineella Chambers, new combination. (Figs. 56, 167) Eripliia (?) nigrilineella Chambers, 1878. Bull. U. S. G-eol. Geog. Surv. Terr., 4: 96. Chambers, 1880. Jour. Cincinnati Soc. Nat. Hist., 2 : 204, f. 42. Eriphia nigrilineella , Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 540. Mc- Dunnough, 1939. Mem. S. California Acad. Sci., 2 : 64. Tongue white. Labial palpi white ; second segment black to two-thirds, a few black scales ventrally before apex ; third seg- ment with a black annulation at two-fifths and another at four- fifths. Antennae buff-white, pecten ochreous, no pattern of black and white on specimen available. Head white, lower two-thirds of face gray with purple reflections, vertex with lavender reflections. Thorax black medially, white laterally. Forewings white with a light dusting of black flecks; a pattern of two oblique black bands as in M. lineigera, but basal one interrupted medially ; black at end of fold ; cilia white, heavily dusted with black at apex, a few black flecks between apex and dorsal margin, no markings dorsad of wing margin. Ilindwings gray. Legs shining white ; metathoracic tibiae with an oblique black stripe at four-fifths, dorsal surfaces of spurs black, under certain light conditions ; metathoracic tarsi with a broad black annulation on middle of first segment, black at apex of second segment and at base of third segment. Male genitalia: (Fig. 56) K.W.H. slide no. 921. Female genitalia : no specimens available. Alar expanse : 7 mm. Food plant : unknown. Type : Museum of Comparative Zoology. Type locality : Bosque County, Texas. Specimens examined: 2 Arizona: Madera Canyon, 4880 feet, Santa Rita Mountains, Santa Cruz Co., 1 June 30, 1959 (R. W. Hodges), [RWH]. Texas: Bosque Co., 1 J', no date [MCZ] . M. nigrilineella can be separated from M. lineigera by the black dusting on the forewings. 65 ENTOMOLOGICA AMERICANA Eteobalea, new genus Eieos = true, patios = spotted (Figs. 2, 20, 53, 113) ! j Type: Gelechia sexnotella Chambers, 1878. Head smooth-scaled ; tongue scaled, moderately short ; maxillary palpi folded over base of tongue ; labial palpi recurved, reaching beyond vertex, first and second segments smooth dorsally, second segment tufted ventrally, third segment smooth, second and third segments subequal ; antennae three-fourths, pecten present, simple. Forewings broadly lanceolate, apex acute ; lb furcate basally, sometimes furcate distally ; 2 from two-thirds ; 2, 3, 4, and 5 equidis- tant, subparallel ; 5 connate with 6,7, and 8 ; 6, 7, and 8 stalked ; 10 slightly before 3 ; 11 from two-fifths of cell. Ilindwings lanceo- late, apex acute ; no tuft of scales on costal margin ; 8 veins present ; lb simple ; 2, 3, 4, and 5 equidistant, subparallel ; 6 and 7 stalked, divergent. Male genitalia: costae separate from valvae ; right costa smaller than left one, partially fused with aedeagus ; valvae lobate, slightly expanded distally ; aedeagus shaped like a long-necked vase without a well defined base, ankylosed; cornuti absent; tegumen heavily sclerotized ; tuba analis emerging between brachia ; tegumen emargi- nate apically ; left brachium larger than right one, apex enlarged, blunt. Female genitalia : lamellae vaginales small, heavily sclerotized, mainly developed anterior to ostium bursae with a narrow band encircling posterior part of ostium bursae; ductus bursae slender, lightly sclerotized ; corpus bursae lightly sclerotized ; signa present. Eteobalea differs from Stagmatophora (Figs. 58, 111) in the following characters : the second segment of the labial palpi is tufted ventrally in Eteobalea ; rough-scaled in Stagmatophora. The second and third segments of the labial palpi are subequal in Eteobalea ; the second segment is shorter than the third in Stagma- tophora. In the male genitalia of Eteobalea the left brachium is large, the apex is blunt and slightly enlarged, the right brachium is a blunt lobe ; in Stagmatophora the brachia are slender, subequal, and taper to the apices. McDunnough (1939) lists six species in Stagmatophora. Of these 8. iridella, S. sexnotella , and S . wyattella are transferred to Eteobalea ; S. niphochrysa is a synonym of iridella; and 8. ceano- thiella and S. gleditschiaeella are transferred to Periploca in the Walshiidae. In addition one new species is described, bringing the number of known species for our fauna to four. 66 VOLUME XLII Key to the Species of Eteobalea based upon the Maculation 1. Forewings black with white spots 2 Forewings brown or bronze 3 2. Hindwings light fnscous ; base of tongue white, base of fore- wings usually black sexnotella Hindwings dark fuscous ; base of tongue black ; base of fore- wings white wyattella 3. Base of second segment of labial palpi bronze-black enchrysa Base of second segment of labial palpi white iridella Key to the Species of Eteobalea based upon the Male Genitalia 1. Apex of left brachium four or more times width of narrowest part of brachium (Fig. 52) iridella Apex of left brachium less than three times width of narrow- est part of brachium 2 2. Apex of left brachium more than twice width of narrowest part of lobe (Fig. 55) wyattella Apex of left brachium less than twice width of narrowest part of lobe 3 3. Left brachium with a lobate medial projection (Fig. 54) enchrysa Left brachium without a medial projection (Fig. 53) sexnotella Key to the Species of Eteobalea based upon the Female Genitalia 1. Lamella postvaginalis subtriangular, lightly sclerotized; twTo, lightly sclerotized signa in form of triangular plates (Fig. 113) sexnotella Lamella postvaginalis not subtriangular, heavily sclerotized ; signa heavily sclerotized 2 2. Lamellae vaginales irregularly margined with constriction before anterior termination ; signa with irregular margins (Fig. 114) wyattella Genital plate and signa with smooth margins 3 3. Lamellae vaginales subreniform (on right side) ; signa one- third to one-half length of corpus bursae (Fig. 112) iridella Lamellae vaginales subquadrate ; signa one-seventh to one- eighth length of corpus bursae (Fig. 115) enchrysa 67 ENTOMOLOGICA AMERICANA Eteobalea sexnotella Chambers, new combination (Figs. 53, 113, 197) Gelechia sexnotella, Chambers, 1878. Bull. U. S. Geol. Geog. Surv. Terr., 4:88. Hagen, 1884. Papilio, 4:99. Riley, in Smith, 1891. List of the Lepidoptera of Boreal America, 102. Mompha sexnotella, Busck, 1902. Jour. New York Ent. Soc., 10: 97. Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 543. Stagmatophora sexnotella, Walsingham, 1907. Proc. U. S. Natl. Mus., 33:219. Barnes and McDunnough, 1917. Check list of the Lepidoptera of Boreal America, 153. McDunnough, 1939. Mem. S. California Acad. Sci., 2 : 64. Laverna sexnotella, Forbes, 1923. Lepidoptera of New York and neighboring state, Cornell Univ. Agric. Exp. Sta. Mem., 68 : 324. Tongue and maxillary palpi white. Labial palpi white with three black annulations : one at base of second segment, one at two-fifths and one at four-fifths on third segment. Face and vertex white, a fuscous line from base of each antenna to base of tongue, a row of black scales along posterior edge of eyes. Antennae : scape white on ventral surface and at apex, black on dorsal surface ; shaft black with fuscous annulations, becoming white on outer third. Thorax black. Forewings black with six shining white spots; basal white spot broken in area of fold. Hindwings light fuscous. Abdomen dirty yellow, last segment fuscous. Legs black ; metathoracic tibiae with three white stripes : one at one-fourth, one at two-thirds, and one apical; tarsal segments with white apices ; base of first tarsal segment white. Male genitalia: (Fig. 53) R.W.H. slide no. 830. Female genitalia: (Fig. 113) R.W.H. slide no. 831. Alar expanse : 8-14 mm. Food plant: Tricliostema dichotomum L., T. suffrutescens Kear- ney. The larva is a stem gall former. Type : Museum of Comparative Zoology. Type locality: Bosque County, Texas. Specimens examined: 75 J1, 31 2- Connecticut: East River, 1 August 6, 1907 (Chas. R. Ely), [USNM] ; same locality, 8 J', July 8-21 [MCZ]; Putnam, 1 ?, May 27, 1933 (A. B. Klots), [ABK]. Florida: Archbold Biological Station, Highlands Co., 2 J1, 4 March 27 through April 2, 1959 (R. W. Hodges), [RWH] ; same locality, 1 lCf, 1 2, July 15-31, 1948 (A. B. Klots), [ABK] ; Georgiana, 1 1 2? gall on Tricliostema dichotomum, emerged 68 VOLUME XLII July 11, 1882 (Wm. Winfield), [USNM] ; Longboat Key, Sara- sota, 1 5, gall on Trichostema suffrutescens, emerged January 31, 1945 (J. G. Needham), [CU] ; Oneco, 1 J1, May 28, 1953 (Paula Dillman), [CPK] ; Sarasota, 2 J*, 1 J, February 1945 (J. G. Need- ham), [CU] ; Siesta Key, Sarasota Co., 2 2, April 24 and May 4, 1960 (C. P. Kimball), [CPK]. Kansas: Onaga, 1 $ [MCZ]. Mary- land: Frederick, 3 J1, June 11, 1906 [MCZ] ; Hyattsville, 1 J', June (A. Bsk), [LACMl ; Washington, D. C., 2 J1, May 21, 1906 (Chas. P. Ely), [USNM] ; same locality, 1 J', 1 J, May and June (A. Bsk), [USNM, MCZ]. Massachusetts: Barnstable, 8 J', 3 2, June 5 through July 24 (C. P. Kimball), [CPK] ; Magnolia, 1 <$, June 3, 1908 [USNM] ; Martha’s Vineyard, 2 on corn, emerged September 15 through October 15, 1894 [USNM] ; Monroe, 1 1 2, from old cotton boll, emerged October 22 and 23, 1900 [USNM, RWH]. Maryland: Washington, 2 4 no date (A. Busck), [USNM]. Mississippi: Batesville, 1 <2, 1 2> 1-2-1915 [USNM] ; Greenwood, 1 2> June 16, 1915 (J. M. Langston), [USNM]. Texas: Beeville, 1 ,<$, issued 75 ENTOMOLOGICA AMERICANA from cotton bolls, November 21, 1895 [USNM] ; Beeville, 1 J', corn husks, emerged September 13, 1895 [USNM] ; Beeville, 2 2, April 25, 1896 [USNM, KWH] ; Columbia, 2 J, from rotten cotton bolls, emerged December 25, 1879 (E. A. Schwarz), [USNM] ; Corpus Christi, 1 September 21, 1943 (W. M. Gordon), [CU] ; Smith Point, 1 <^, from old bolls of cotton, emerged October 17, 1923 (L. J. Bottimer), [USNM] ; Smith Point, flower heads of Mesosphaerum rugoswm, emerged October 18, 1923 (L. J. Bottimer), [USNM] ; Welasco, 1 J, reared from flowers of castor bean, emerged August 30, 1952 (P. T. Riherd), [USNM]. The characters given in the keys will serve to distinguish be- tween rileyi and S. badia. Sathrobrota badia, new species (Pigs. 60, 118, 170) Pyroderces rileyi Busck, 1917. Jour. Agric. Res., 9 : pi. 8, f. D. ( misidentification. ) Tongue pale buff-white basally, yellow distally. Labial palpi white with tinge of buff ; second segment with a broad tawny band on outer surface at one-fifth, a tawny annulation at three-fifths, apex tawny ; third segment with three brown annulations : one postbasal, one medial, and one preapical. Antennae : pecten brown, scape buff-white with tawny dusting on dorsal surface, apex of scape pale, shaft white to buff-white ventrally and on distal half of each segment dorsally. Pace buff-white, vertex pale tawny. Thorax pale buff-white overlaid with tawny. Forewings heavily overlaid with tawny ; costa fuscous to one-half ; pattern composed of black tipped scales preceded or followed by buff-white scales ; a few black scales on costal margin at one-eighth ; a broken transverse row of black scales at one-fourth, not reaching costal or dorsal margin ; a subtriangular to quadrate patch of blackish scales at one-half in center of wing ; a subcostal series of black scales from three-fifths to four-fifths ; a row of dark brown scales along outer margin from tornus to apex of wing; a short brown cross at apex; cilia gray- brown. Hindwings fuscous. Abdomen buff-brown with lavender re- flections dorsally, cream-white with some gray ventrally. Meta- thoracic tibiae tawny on basal two-fifths of outer surface, an oblique white streak from middle tibial spur, outer surfaces of distal half of tibiae dark brown, apex pale buff -white, tibial spurs white or pale buff -white with black at middle ; tarsal segments dark brown basally 76 VOLUME XLII on dorsal surface, pale apically. Male genitalia: (Fig. 60) R.W.H. slide no. 1031. Female genitalia: (Fig. 118) R.W.H. slide no. 595. Alar expanse : 7%-ll% mm. Food plant : scavenger, recorded from the following materials : pine cones infested by Dioryctria (Lepidoptera : Pyralidae), rust infected cones of Pinus elliottii, Pinus palustris, pods of Cassia occidental is, peach mummy, mnmmy fossil of loquat, limes, grape- fruit, bananas, cabbage, blossoms of cocoannt, and elm leaves. Holotype : So. Florida, in pine cones infested by Dioryctria, emerged June 16, 1931 (J. K. Small), [USNM Type No. 66083]. Paratypes: same data as type, 3 J1, 1 2, June 16-18, 1931 [USNM, RWH] ; Santa Ana, Orange So., California, 27 J', 28 2, reared from peach mummy, collected March 26, 1943, emerged April 1-16, 1943 (H. H. Keifer), [CAS, RWH, JAP] ; Costa Mesa, California, 1 2? reared from peach mummy, collected November 10, 1942, emerged December 16, 1942 (Gammon), [CAS] ; Cocoanut Grove, Florida, 2 lCf, 4 2> from blossoms of cocoanut, May 16-23, 1916 (H. M. Matheson), [USNM, CU] ; Miami, Dade Co., Florida, 6 limes, May 1918 (G. F. Meznette), [USNM] ; Osceola Natl. Forest, Baker Co., Florida, 1 J', reared from rust infected cones of Pinus elliottii, June 28, 1957 (E. P. Merkel), [USNM] ; Palmetto, Florida, 1 2, cabbage, June 12, 1945 [USNM] ; Royal Palm State Park, Florida, 1 2, no date (F. M. Jones), [USNM] ; 3 miles S. Clarcona, Florida, bred from grape- fruit, emerged September 25-30, 1930 (F. H. Benjamin), [USNM] ; Siesta Key, Sarasota Co., Florida, 2 J', 5 2, December 1 through April 19 (C. P. Kimball), [CPK] ; Boothville, Louisiana, 5 2? ex bananas, March 8, 1945 [USNM]. 77 ENTOMOLOGICA AMERICANA Limnaecia Stainton (Figs. 1, 12, 63, 121) Type : Limnaecia phragmitella Stainton, 1851. Monobasic. Limnaecia Stainton, 1851. Suppl. Cat. Brit. Tineina, 4. Wocke, in Heinemann, 1877 [1876]. Die Schmetterling* Deutschlands nnd der Schweiz, 2 : 421. Staudinger and Rebel, 1901. Catalog der Lepidopteren des Palaearctischen Faunengebiets, pt. 2 : 187. Spuler, 1910. Schmetterlinge Europas, 2 : 384, f. 142. Forbes, 1923. Lepidoptera of New York and neighboring states, Cornell Univ. Agric. Exp. Sta. Mem., 68 : 324, f¥. 181, 194. Fletcher, 1928. Catalogue of Indian Insects, pt. 16 : 18. Fletcher, 1929. Mem. Dept. Agric. in India, Ent. Ser., 11 : 127. Diakonoff, 1954. Verh. Akad. Wet. Amsterdam, 50, no. 1: 74. Laverna Stainton, 1854. Insecta Britannica. Lepidoptera : Tineina, 238. (partim.) Limnoecia Meyrick, 1888. Trans. New Zealand Inst., 172. (emenda- tion). Busck, 1901. Proc. Ent. Soc. Washington, 4 : 421. Anybia Meyrick, 1928. Revised Handbook of British Lepidoptera, 652. (incorrect association). Fletcher, 1928. Catalogue of Indian Insects, pt. 16:19. (as synonym of Limnaecia.) Fletcher, 1929. Mem. Dept. Agric. in India, Ent. Ser., 11 : 17, 127. (as synonym of Limnaecia). Lymnaecia Dyar, 1902 [1903]. List of the Lepidoptera of North America, Bull. U. S. Natl. Mus., 52 : 537. McDunnough, 1939. Mem. S. California Acad. Sci., 2 : 64. The synonymy given by Fletcher (1928) and cited with doubt by Diakonoff (1954) is not repeated. Study of the type species involved should be made before the list is accepted. Head smooth-scaled; tongue scaled, well developed; maxillary palpi folded over base of tongue, slightly drooping; labial palpi recurved, reaching well beyond vertex, smooth-scaled, third segment longer than second, apex acute ; antennae two-thirds to four-fifths, pecten present, ciliate in male, simple in female. Forewings narrow, elongate, apex acute ; 12 veins present ; lb furcate basally ; 2 from two-thirds on cell ; 2, 3, and 4 equidistant ; 7 and 8 stalked; 10 from three-fourths on cell; 11 from two-fifths on cell. Hindwings narrow, elongate, apex acute ; 8 veins present ; 2 from three-fifths on cell ; 2, 3, 4, and 5 equidistant ; 5, 6, and 7 approximate ; 6 and 7 diverging. Male genitalia : valvae asymmetrical ; costal area separate or heavily sclerotized and connected to valva; aedeagus short, anky- 78 VOLUME XLII losed, basal process present ; arms of gnathos separate, left brachium larger than right one, heavily sclerotized. Female genitalia : bursa copulatrix membranous ; signa present ; lamellae vaginales forming a long heavily sclerotized tube ; apo- physes anterior es broad, heavily sclerotized ; apophyses posterior es sinuate distally. One species of Limnaecia is known to occur within the region considered by this paper; and it is reported by Meyrick (1928) to be found in Europe, North Africa, Australia, and New Zealand. He believed that it has been dispersed in the larval stage by being blown with the down of Typha , the food plant of the larva. Limnaecia phragmitella Stainton (Figs. 63, 121, 171) Limnaecia phragmitella Stainton, 1851. Suppl. Cat. Brit. Tineina, 4. Spuler, 1910. Schmetterlinge Europas, 2 : 384, pi. 89, f. 66. Forbes, 1923. Lepidoptera of New York and neighboring states, Cornell Univ. Agric. Exp. Sta. Mem., 68 : 324. Laverna phragmitella Stainton, 1854. Insecta Britannica. Lepi- doptera : Tineina, 238. Stainton, 1870. Natural History of the Tineina, 11 : 150-159, pi. 4. Limnoecia phragmitella , Meyrick, 1895. Handbook of British Lepi- doptera, 675. Busck, 1901. Proc. Ent. Soc. Washington, 4: 421. Lymnaecia phragmitella , Dyar, 1902 [1903]. List of the Lepi- doptera of North America, Bull. U. S. Natl. Mus., 52:537. Claassen, 1921. Typha Insects, Cornell Univ. Agric. Exp. Sta. Mem., 47 : 487. McDunnough, 1939. Mem. S. California Acad. Sci., 2 : 64. Tongue buff, apical one-third very lightly scaled. Labial palpi buff -white, a few pale brown scales on dorsolateral surface of first segment, second segment with a few pale brown scales subbasally and subapically, third segment with an internal and an external brown line. Antennae buff, ventral surface of scape with a few pale brown scales at base, scape with an anterior brown line con- tinued on shaft as a series of dots to one-half. Face and vertex buff-brown. Forewings buff-brown with a medial brown spot at one-third, another brown spot at two-thirds, four or five pale brown spots on costal margin on apical one-third, a faint pale brown spot on dorsal margin near vertex, a pale brown spot on fold at two- thirds, internal brown spots encircled or partially bordered by white scales, costal spots bordered internally by white scales, a gray 79 ENTOMOLOGICA AMERICANA streak from inner medial dot to apex complete or partially devel- oped, cilia shining buff. Hindwings gray-brown, cilia buff. Abdo- men buff dorsally, buff-white ventrally. Legs buff to buff-brown, prothoracic tibiae gray-brown with a dorsal buff line. Male genitalia: (Fig. 63) R.W.H. slide no. 597. Female genitalia: (Fig. 121) R.W.H. slide no. 3. Alar expanse : 15-20 mm. Food plant: Typha latifolia L. and T. angustifolia L. Claassen (1921) gives the following discussion of the habits of the immature stages : ‘ ‘ The larvae restrict their work to the head of the plant, except occasionally when they bore into the stem to transform. The young larvae feed on the tender styles of the pistillate flowers, but as these grow larger and become dry, the larvae move farther inward and eat the seeds of the plant. As cold weather approaches, they mi- grate still farther inward, and finally locate near the rachis of the flower spike, where they often eat away the basal part of the little stalks which bear the seeds. The larvae spin an abundance of silk with which they tie the down, or pappus, together, thus keeping it from being torn off or blown away. “The cat-tail heads which are infested by these larvae present a striking appearance. The silk spun by the larvae holds the downy material together and does not allow the seeds to escape, but the heads fluff out. . . . “The larvae overwinter in the half-grown stage in the head of the plant, the fluffy material of the fruiting spike being their pro- tection. “In the latter part of May or early June the larvae attain their full growth. Then, in the midst of the downy material, the larvae spin their thin, tough, white cocoons and transform to the pupal stage. Many of the larvae, leaving the heads, go down and bore into the stem of the cat-tail plant, forming burrows which they line with silk; and there they pupate.” Type: British Museum (Natural History). Type locality : ? England. Specimens examined : 100 J1, 108 J. Colorado : Watkins, Adams Co., 4 August 10, 1956 (J. G. Franclemont) , [JGF] ; McLean Bogs Re- serve, Tompkins Co., 1 lCf, 23 2> July 17-22 (J. G. Franclemont), [CU, JGF] ; Mattituck, Long Island, 1 110 date given [USNM] ; Orient, Long Island, 3 1 2? May 12 through June 16, 1934 and September 9, 1934 (Roy Latham), [CU] ; North Collins, 10 J1, 10 2, June 27 and 28, 1938 (W. T. M. Forbes), [CU] ; River- head, Long Island, 1 2? July 1, 1934 (Roy Latham), [CU] ; Roches- ter, 1 2, July 8, 1933 (A. B. Klots), [ABK]. Ohio: Cincinnati, 1 J', reared, emerged June 10, 1922 (Annette F. Braun), [CAS]. Pennsylvania : Germantown, 1 J', June 21 [MCZ] ; Hazelton, 5 June 8 through 24 (Dietz), [USNM, MCZ] ; Mt. Airy, 1 1 2, June 25 [USNM, MCZ] ; Oak Station, Alleghany Co., 3 5% Permanent I 5(50%) — 2(20%) 3(30%) Permanent II 3(38%) 2(24%) 3(38%) Temporary Type 1 Type 2 13(46%) 5(18%) 5(18%) 5(18%) 12(92%) 1(6%) 5(33%) 1(08%) 4(27%) 5(33%) Among the brachypters of permanent habitats, two, Scoloposte- tkus diffidens and Antillocoris minutus have a frequent production of macropters, and, in accordance, both may occur in subclimax 34 VOLUME XLIII Betula populifolia litter, and Antillocoris in fact sometimes is found sporadically in some cool meadows, then always macropterons. Southwood (1960) listed the habitat of Antillocoris minutus and A. pilosulus to be temporary. However, the habitat of the macrop- terous A. pilosulus in Florida and North Carolina (forest edge litter) does not agree with this placement. The frequency of Antillocoris in air trapping is undoubtedly the result of its small size which allows it to be lofted high by winds during the dispersal phase. The three species with a very low level of macroptery, Drymus crassus, Plinthisus americanus, and Ligyrocoris caricis are all found in climax or serclimax habitats. When the many rhyparochromines (74%) of temporary habitats are considered, in the sense of Southwood (Table 3, Temporary) it is seen that only a minority (46%) are completely maeropterous, which is again, seemingly unaccountable. But if one considers what is rarely mentioned by ecologists (Burges 1960), the time element involved in the succession rates, a quite different pattern emerges (see Ecology). Compared to the annual life of an insect the decades in which some xeroseres undergo succession (Blizzard 1931) is a very long time. Type 1 (Table 3) includes the rapid mesosere and bare ground (rotation arable, ruderal, etc.) succession habitats which rarely last more than a year or two. In complete correlation 92% of the rhyparochromines of such habitats are totally maeropterous and the only exception is the introduced Stygnocoris rusticus which may not be exploiting its natural habitat. Type 2 (Table 3) includes the slow progressing xerosere successions, and is again in close correlation with 94% of the species exhibiting brachyptery. The bionomics of the sole apparent exception, Kol- enetrus plenus , is not well understood. The three levels of brachy- ptery proportions also correspond to habitat permanence within Type 2 habitats. Those species with a large percentage of ma- cropters (2) are members of the Andropogon association which often constitutes a relatively short-lived habitat in the open xerosere series. In contrast, those species with rare macropters (4) are fre- quently found in sparse numbers in serclimax grassy or Vaccinium scrub bald habitats but extend ( in greater abundance ) into morainic lowland sites. The sole exception is Sisamnes in the Andropogon association whose range, however, is mostly western. Moreover the rareness of Sisamnes may stem from a consequent poor adaptation to the habitat mosaic of New England. Those species with intermedi- ate brachyptery are also found in serclimaxes, but are even consider- ably more numerous in the lowland temporary habitats. It appears then that a good correlation exists between the propor- tions of brachypters and the habitat permanency. Lack of exact 35 ENTOMOLOGICA AMERICANA habitat ages precludes a closer comparison, but it should be noted the proportion of brachypters in some species especially abundant in climax forest may be higher than some other species upon xeric field succession slopes, i.e., there is a complete overlap between per- manent and temporary habitat as defined by Southwood. The over- lap was explained as with water beetles (Jackson 1928) in terms of the total habitat and distribution range. Nevertheless since southern New England was once almost com- pletely denuded of forests (Bromley 1935), and the dry sites on outcrops and morainic slopes are quite discontinuous, a perfect mosaic of favorable habitats for different species is present. As Parshlej^ (1920b) said, on finding brachypterous Microvelia in pools on Mt. Greylock, Massachusetts, “The common occurrence of several species of this genus in such isolated situations indicates the importance in the economy of the race of the fully winged phase, which must appear in sufficient frequency to provide for a favorable rate of dispersal.” Relation of Dispersion to Habitat Despite the misgivings expressed about the lygaeid flight records (in southern England) there appears to be a good correlation be- tween temporary habitats and dispersal records, when one discounts the tribes which despite their abundance are rarely or never found at lights, especially the Gonianotini, Megalonotini and temperate Drvmini, Stygnocorini and Plinthisini and considers the Myodochini alone (Table 4). In excellent correlation all the myodochines of short-lived temporary (Ti) habitats have been collected at lights, including the marsh dwelling Pachybrachius albocinctus. Only a few dispersal records on the other hand are available for the species of the T2 xerosere habitats and these are mostly the inhabi- tants of the newest almost ruderal Andropogon type habitats, which species also have a high percentage of macropters. Even among the other tribes, the various existing dispersal records are of species of temporary habitats except with the spring-dispersing Eremocoris and Antillocoris and Ozophora which very frequently flies despite its permanent type habitat. Dispersal among the other tribes may be of mainly a diurnal type. The apparent difference in reaction to light among these tribes in eastern North America would be a significant area of research. Southwood (1962a) has grouped migratory variability into two categories, obligatory and facultative. In the first the species is obligatorily dimorphic, i.e., under genetic control; in the second, the migratory response is facultative, and triggered by certain en- vironmental conditions, such as crowding, or habitat changes. The 36 VOLUME XLIII latter is an especially ideal adaptation to temporary environments. In the second category Johnson’s teneral migration period may be modified to include other non-sexual periods (Johnson 1960b). It is difficult to apply this system to the present taxon for not enough is known about what promotes dispersal, nor was any work done upon the proportion of obligatorily dispersing individuals in a population of macropters. However, most of the fully macropter- ous species are probably of the facultative type. Among the bra- chypterous species, the type would depend on whether the ma- cropterous condition is genetically obligatory; or also requires an environmental stimulus, and so, is facultative. For this a fuller discussion of wing polymorphism is required. Wing Polymorphism Fully half (51%) of the Rhyparochrominae of New England have brachypterous forms (Table 4). As described in the previous discussion on dispersal the proportion of brachyptery was noted (Table 4) from 1 (macropterous) to 4 (nearly entirely brachypter- ous) . In nearly all species the brachypterous form is distinct with very few intergrades between it and the macropter. Several differ- ent types of brachyptery exist. In 15 (75%) of the brachypterous species the entire wing is shortened, and the membrane is reduced in extent and slightly overlaps. Trapezonotus arenarius (see Species Discussion) exhibits a sexual dimorphism in that the female shows a more plastic expression of brachyptery. Scolopostethus thorn soni shows a considerable non-sexual variation in degree of brachyptery (Butler 1923, Southwood 1961b). In Ligyrocoris syl- vestris, depictus, and caricis the membrane is not so reduced (sub- macropterous) but no macropterous forms were found. The five remaining species have quite different types of brachyp- tery. Antillocoris minutus is unusual in that it has three distinct types of wing development, the macropter, the “normal” brachyp- ter, with the wing reduced, and a subbrachypterous form with the corium truncated and the membrane lost (see Antillocoris) . Drymus crassus has rather coleopteroid hemelytra, and the hind wings are reduced. The corium is elongated and the membrane is narrowed and reduced although attaining the apex of the abdomen. In an occasional female the membrane was slightly shorter. In Plinthisus americanus, although the corium is also elongated and the elaval suture indistinct, the membrane is reduced but overlaps. In Carpilis consimilis the membrane is reduced to a very thin non- 37 ENTOMOLOGICA AMERICANA overlapping fringe. Finally in Sisamnes clavigera the membrane is altogether absent as in Antillocoris and the corium somewhat trun- cated. It is significant that Drymus crassus and Plinthisus with their coleopteroid type of brachyptery were rarely found outside of climax forests. The obvious question, as Butler (1923) asked, is why the varie- ties of hemelytral development? Indeed, the question may be re- stated to ask why does the development of the brachyptery extend beyond a flight polymorphism as in some aquatic Corixidae and No- tonectidae (Young 1961) ? A flight polymorphism of this type would be sufficient to insure that the favorable habitats remain populated especially if a dispersal flight is obligatory in teneral macropters as Johnson suggests. As Darwin (1859, pp. Ill, 346) long ago proposed, such wing reductions could result from natural selection to allow the most economical growth, when flight became of no selective value. It therefore follows that if a large scale dispersal away from a habitat is no longer a selective advantage, but instead only a certain low per- centage is required, a disruptive selection (Ross 1962) could oper- ate. This could bring about an evolutionary progression from little or no structural dimorphism to the most extreme, but economical di- morphism between the macropter and the brachypter. It would then also follow that only under long continuing selection would the extreme dimorphism result, and it then becomes relevant that the rhyparochromines with extreme brachypters are denizens of some of the most permanent habitats in eastern North America. Brinkhurst (1959) in this connection, considered the selective advantage of the brachypter of Gerris odontogaster (Zett.) to be at least 5% over the macropter to overcome a lethal condition. Hormone Control Southwood (1961b) in reasoning from Wigglesworth’s experi- mental work (1952) with Rhodnius has proposed that a brachypter- ous form is either a neotenic (juvenile) expression (methathetely) produced by an excess of juvenile hormone, or is paedogenetic and results from a depression of the juvenile hormone level leading to a last instar with adult characters (prothetely) . Since all the rhy- parochromines studied have five full instars, and are heavily sclero- tized, all then would represent some expression of metathetely. Southwood (1961b) proposed that cold temperatures promoted metathetely by lengthening the exposure of the last instar to juve- nile hormones, and cites evidence to show that in many species the brachypter is associated with northern distributions and with moun- tain populations. He considered most important as experimental evidence Brinkhurst ’s (1958, 1959) demonstration that the exposure 38 VOLUME XLIII of the eggs of Gerris odontogaster (Zett.) to warm temperatures produced macropterous adults. However, this cannot be considered as being metathetelous in the sense that the effect of the juvenile hor- mone is extended in the last instar. Southwood (1961b) further considered the reduction in ocelli size (Butler 1923) as another neotenic expression. Only Cnemodus among the New England rhyparochromines lacks ocelli. This, how- ever, occurs both in the macropter and the brachypter. In Plinthi- sus americanus the macropter has distinctly larger ocelli. There appears to be little difference in size in the other species. If the ocelli are important in flight behavior, their quoted reduction in the brachypters may also correlate with their reduced selective value. While, like much growth phenomena, the wing shortening is under hormonal control ( Wigglesworth 1952, 1954), it appears somewhat debatable to interpret all brachypters as simple general neoteny, since the rhyparochromine brachypters are very similar otherwise to the macropters and the wings are usually nearly iden- tical in structure and only reduced in size, i.e., it is not a wing pad in structure or an intermediate structure. Moreover, a complete gradation exists between wing muscle loss and very short wings, and an all or nothing tissue response in different species would produce a more abrupt and nymphoid wing distribution as in the Rhodnius seventh instar adult (Wigglesworth 1954). The close relation be- tween habitat permanence and brachyptery would suggest a selec- tion control rather than environmental cold control suggested by Southwood (1961b). The various and diverse modifications of the brachypterous wings, of which a few simple examples among the rhyparochromines have been given here, would further illustrate a selective and adap- tive procession, it may be more accurate and useful then to con- sider selection as utilizing a variable hormone threshold to effect a brachypterous condition. Given a cold habitat, selection may then possibly utilize the normally lengthened last stadia to operate a hormone balance or threshold mechanism. Latitudinal Relationship Since Southwood (1961b) suggests that the metathetelic macrop- ters would be more abundant at the southern limit of the species range, the distribution patterns should be considered. In the Rhyparochrominae such a relationship was not seen, and on the contrary, the brachypterous species with more northern distribu- tions were almost entirely brachypterous and the macropters oc- curred near the center of the species distribution, not at the southern limits, while the brachypterous species of more southern distribu- 39 ENTOMOLOGICA AMERICANA tion appeared to have an ample representation of macropters in Connecticut. Sisamnes clavigera is a possible exception as all speci- mens of this species ever collected in the northeastern United States have been brachypterous, but rearing under many conditions failed to yield the macropter. Despite both areas having a north temperate location, the gen- erally continental climate of northeastern North America produces very warm mid-summer temperatures in contrast to the moderate Atlantic climate of western Europe, and open land areas (Cloudsley- Thompson 1962) become very hot and dry. Since many brachypter- ous rhyparochromines favor such habitats, the actual development period, especially of the last instar, occurs under very hot field con- ditions. Yet these progeny are as brachypterous as those reared under cooler laboratory conditions. Moreover both slow-developing and rapid-developing species may be brachypterous or macropterous. There appears, nevertheless, to be a definite correlation between species of northern distributions and wing polymorphism. The in- cidence of brachypters among boreal species is 11 of 14 species (79%) ; and among intermediate ranges 4 of 13 species (31%) ; and among austral ranges 5 of 14 species (36%). This correlation, however, may stem more from the habitat per- manency of these particular boreal species than from their distribu- tion. Most of these boreal northern species are found in semiperma- nent or serclimax habitats, usually hot in summer. Indeed, this group may represent the element most closely adapted to New Eng- land climatic conditions, while most of the other species which have their main distribution centers elsewhere are adapted to temporary subclimax associations. This situation is common in subclimax asso- ciations (Ross 1962). Moreover the relationship in terms of total latitudinal range limits is the direct converse of what Southwood predicted — the northern species at their austral limits are nearly entirely brachyp- terous, and most southern and intermediate species macropterous near their boreal limits. From the standpoint of selection this pat- tern would be expected. Other Environmental Factors On more general grounds, if metathetely were based on hormone control such as suggested by Southwood many detrimental condi- tions should favor the production of the brachypters by lengthening the last instar through low food levels or through excessive cooling. If purely genetic considerations are involved, a rare gene combina- tion for macroptery would be more possible under heavy densities. Macroptery arising from a crowding stimulus would produce a 40 VOLUME XLIII similar result. Thus macroptery development and dispersal would occur at optimal times ; brachyptery and contraction at inoptimal conditions. In a permanent or semi-permanent environment, this situation is reasonable, for the brachypters would insure survival of a popu- lation ; and dispersal would occur when optimal conditions for colony establishment or ecesis may exist in other habitats. Laboratory Work First it should be emphasized that all the species were reared under laboratory conditions in environments considerably warmer or cooler than field conditions, and several species were deliberately reared under hot conditions in an effort to obtain the rare macrop- terous forms. Moreover, the stadia could be considerably acceler- ated or delayed by temperature and food conditions. The Heterop- tera moult only 5 (rarely 4) times and the stadia may be greatly varied by starvation, or, in a few species, a superabundance of food. If cool conditions can lengthen the stadia and increase the exposure to the juvenile hormone, which assumes that the action of juvenile hormone is not equally affected by cooling, then the lengthening of a late stadia may similarly extend the exposure to juvenile hor- mone, but, moreover, without a concomitant reduction in the rate of action of the hormone. It is then significant that in nearly all of many hundreds of laboratory cultures no change in the proportion of macropters' as compared to field populations was observed. The single exception was Carpilis consimilis where rearing under heavy density condi- tions did apparently promote the appearance of the macropters much as in the leaf hopper Nitaparavala (Kisimoto 1956). On the contrary the pattern usually appears to be under genetic control with little if any phenotypic plasticity, e.g., the penetrance of the gene was complete under the varied and different laboratory con- ditions. These results then do not support among rhyparochromines the juvenile hormone hypothesis of Southwood, and the production of the macropters appears to be usually genetically controlled, e.g., controlling the wing anlage thresholds to the action of the juvenile hormone without phenotypic variability. It is realized that other obscure triggering factors not present in the artificial laboratory conditions may be involved. It might be significant that whenever one of the rarer macropters of Plinthisus and Carpilis was found in the field it appeared among an unusually heavy population of the species. Moreover whenever a sparse popu- lation of a Type 3 brachypter was found it was usually composed 41 ENTOMOLOGICA AMERICANA almost entirely of brachypters. To what extent this results from lack of crowding, or the infrequency of chance rare gene combina- tion, is uncertain and certainly deserves intensive work. While brachyptery does not then appear to be controlled in the New England Rhyparochrominae by nutritional factors, there may be a relation between habitat productivity and brachyptery. There is a predominance of brachypterous rhyparochromines in xerosere (T2) habitats which frequently approach a serclimax in their per- manence. Now the reason for the slowness or failure of these habi- tats to develop to climax is attributed to poor overdrained soil which renders the biological productivity of these areas low. The vegetation of such sites is accordingly very sparse and low, and represents a low biomass productivity. Recalling that the evolution of brachyptery represents a selec- tion for metabolic economy, in contradiction with flight dimorphism per se, and that the rhyparochromine populations are frequently sparse in these severe habitats, it would appear that there would be strong selection for the most economical development to the adult in these habitats of nearly permanent but low productivity. Because so many of the brachypterous species have univoltine life cycles with obligatory diapauses, it was difficult to do crossmat- ings of the wing types and yield genetic proof for the pterygopoly- morphism, as done by Poisson (1924) and Brinkhurst (1958, 1959). It was possible to show that a few macropters will result from brachypter x brachypter matings in Trapezonotus arenarius , Pseu- docnemodus canadensis, Cnemodus mavortius, Cryphula trimacu- lata, Scolopostethus atlanticus, 8. thomsoni, and S. diffidens (see Species for details) . In Antillocoris minutus, as in Eremocoris ferns and Myodocha serripes fertilization occurred before the post hibernational flight by the macropters. In Antillocoris, a univoltine species, the progeny of random mated macropters were nearly all normal brachypters, and all the subbrachypters yielded only subbrachypterous progeny (see Antillocoris) . As Southwood (1962a) emphasized, dispersal patterns and brachyptery should be interpreted as methods of surviving an un- favorable period, and that “the type of habitat change has been of profound importance in determining the evolution of methods of surviving such environmental change by the Arthropoda. ’ ’ To dis- cuss this, the life histories need consideration. 42 VOLUME XLHI Seasonal Cycles In a cold temperate climate such as New England’s, the life cycles of the various rhyparochromines must be adapted to the sea- sonal fluctuations and synchronized to allow survival over the ad- verse winter and sometimes, the late summer periods. As summar- ized by Andrewartha (1952) and Lees (1955), it has long been known that arthropods adapt to seasonal ecological stress by going into a state of diapause, that is, a cessation or retardation of growth, metabolism, and reproductive activity, a state which is largely inde- pendent of the ambient temperatures. Quiescence, in contrast, is directly controlled by the ambient conditions and upon return to favorable conditions the insect will develop, etc., without delay. As Andrewartha (1952) and Bonnemaison (1945) have stressed, diapause functions chiefly as a timing mechanism to ensure that the insects will become active only during favorable periods and will not respond to brief variations during unfavorable conditions, such as an unusually warm period during autumn or spring. In this study only the general outlines of the seasonal cycles and diapause were traced and no attempt was made to establish in de- tail the optimum conditions for diapause development or the pre- cise conditions which initiate facultative diapause. It should be added that since many of the species have a considerable north-south distribution, the seasonal cycles and diapause may show some geo- graphical variation (Musaki 1961, de Wilde 1962). In a few spe- cies data from North Carolina and Florida could be included. The seasonal patterns given are characteristic of the New England Rhyparochrominae. The aspects of the seasonal cycles considered here (Table 4) include generations a year, the overwintering state, type of dia- pause, diapause intensity, and periods of nymphal development and oviposition. Details of the phenology, diapause conditions, photo- periodicity, etc., are given under the individual species. All the rhyparochromines were either univoltine or bivoltine, and in a few species, Pseudocnemodus canadensis and Peritrechus f rat emus, both conditions prevailed. As usual (Lees 1955) most of the univoltine species have obligatory diapause states. Diapause occurred only in the adult or egg state, and there were no examples of nymphal diapause as in Beduvius personatus (L.) (Readio 1931) . In both the egg or adult diapause the diapause was either obligatory or facultative. There were several types of facultative diapause. The most com- mon involved a photoperiodic response resulting either in retarding reproductive development (gonadotropic disassociation) or in the 43 ENTOMOLOGICA AMERICANA production of diapausing eggs. In several species which laid dia- pause eggs, the adults remained in reproductive diapause until an autumnal short day or cold stimulus induced oviposition. These species are designated with a ; — designates no summer dia- pause period. In all cases when evidence was gathered (see Species Discus- sions) the factor initiating a bivoltine facultative diapause was photo-period much as predicted by de Wilde (1962). The diapause intensity could be measured by the reaction of the diapausing adults or eggs to continuous warmth. In some, diapause was completed within two months (1) or was absent (0). In others, it was broken after two or three months in warmth (2) or was not completed in warmth before death and a cold exposure was obliga- tory (3). The numbers classify the species in this scheme on Table 4. In most species the diapause development was completed by pro- longed cold exposure (4°C.) in a cold room. Of the 38 species, 15 (40%) overwinter as eggs. This is an un- usual percentage as compared with the north European fauna where only three species of Stygnocoris and Ligyrocoris sylvestris, itself a myodochine, overwinter as eggs (Pfaler 1936, Southwood and Les- ton 1959). This may, as in the nocturnal response of myodochines and ozophorines to light, stem in part from a taxon difference as all of the cold adapted boreal myodochines (10 of 17 species) diapause as eggs. However Plinthisus americanus and both species of Dry- mus also diapause as eggs, yet all of the European species of Dry- mus and Plinthisus whose overwintering condition is known over- winter as adults (Pfaler 1936, Southwood and Leston 1959). It is perhaps significant that the two introduced rhyparochromines which have very successfully invaded the natural ( ? ) habitats in New England are the egg-overwintering species of Stygnocoris. Rather than a response to a more continental climate, the prevalence of egg overwintering may possibly reflect an arid subtropical origin of the Myodochini where estivation through dry periods might occur in the egg state, preadapting the eggs to winter diapause. Egg diapause intervenes in at least three different stages of embryonic development. (1) During early development before much development has occurred — most of the species. These all have strong diapauses except Perigenes constrictus, and Kolenetrus plenus. (2) During late anatrepsis at the “diapause” stage — Ligyrocoris diffusus, and (3) in late katatrepsis, with the embryo nearly fully developed — Stygnocoris rusticus and pedestris and some eggs of Drymus unus. It is then significant that Stygnocoris and Ligyrocoris diffusus, and especially Drymus, considering their mesic habitats, hatch much earlier than most other egg-diapausing 44 VOLUME XLIII species which must complete egg development during the spring. In the species Ligyrocoris diffusus and Drymus unus, an intense cold exposure of some adults in late autumn destroyed the diapause producing factor in the adults and nondiapause eggs were laid. Of the remaining 23 species which overwinter as an adult, 7 or 8 are univoltine (Antillocoris pilosulus appears to belong to this type). The remainder are bivoltine. The diapause intensity among these species varies considerably (see Table 4). The boreal Antillo- coris minutus and Xestocoris nitens have especially intense dispause states. The more austral species exhibit a weaker diapause, espe- cially Sisamnes clavigera, Heraeus plebejus, and Cnemodus mavor- tius. In four bivoltine species Myodocha serripes, Ptochiomera nodosa, Peritrechus fraternus, and Malezonotus fuscosus, the dia- pause state was not broken except under long daylight condition, and after 4-5 months (see Species Discussions). A progression then may be traced in cumulative adaptations to a cold environment, which allows us to group the various species into several seasonal cycle types. Species Groups Type 1. — this corresponds to the sommertypus of Pfaler (1936). These species are univoltine, with an obligative strong diapause and overwinter as an egg. There are two subgroups. The first lays eggs shortly after becoming adult, and the eggs are exposed to warm conditions, often for several months before winter begins, espe- cially the first two species. Ligyrocoris depictus, L. caricis, L. sylvestris, Carpilis consi- milis, and Plinthisus americanus. In the other subtype, the adults do not lay eggs until autumn and require a cold stimulus for its initiation. Stygnocoris rusticus, 8. pedestris, Drymus unus, D. crassus, and Kolenetrus plenus. Type 2. — these species have an obligative univoltine adult dia- pause. In the first three cold exposure was obligatory to complete diapause : Antillocoris minutus, Xestocoris nitens, and Peritrechus palude- maris. Scolopostethus diffidens, 8. atlanticus, Trapezonotus arenarius, and Cryphida trimaculata. In all the species except the early developing Trapezonotus, the new adults did not appear until after July 10 or later. There is a possibility that Peritrechus paludemaris, an austral species, has a facultative diapause determined in an early instar. Type 3. — these species are bivoltine with a facultative diapause as an egg. The first two species lay diapause eggs immediately, 45 ENTOMOLOGICA AMERICANA while the other three have a brief photoperiodic reproductive pause until autumn short day conditions : Pseudocnemodus canadensis , Sphaerobius insignis; Zeridoneus costalis, Perigenes constrictus, and Ligyrocoris diffusus. Type 4. — these species have a facultative bivoltine diapause in the adult stage. The first two become adult remarkably late in the year. The next two species did not break diapause except under long day conditions : Ptochiomera nodosa, Sisamnes clavigera, Myodocha serripes, Malezonotus fuscosus, Heraeus plebejus, P achybrachius basalis, Cnemodus mavortius , P eritrechus fraternus, Megalonotus chi- ragrus, Sphragisticus nebulosus, and Emblethis vicarius. Type 5. — here is included a species with a facultative univoltine (usually) life cycle and a weak diapause. Ozophora picturata. Type 6. — the following bivoltine species have no apparent dia- pause, and overwinter through quiescence. This corresponds to the mischtypus of Pfaler (1936). P achybrachius albocinctus, Eremocoris ferus, and Scoloposte- thus thomsoni. It is significant that only in the non-diapausing Eremocoris ferus were a few nymphs ever found overwintering. A final aspect of the seasonal cycles is the period of nymphal de- velopment and of oviposition. In part, of course, this corresponds closely to the univoltine-bivoltine divisions, but differs in detail and subdivides the other seasonal cycle categories already mentioned. In Type 1 and 3, which overwinter as eggs, the species can be dis- tinguished into early maturing (by late June) and late maturing groups (later, usually much later, than late June). These species directly utilized a spring fallen seed source without prior spring depletion by adults. Early Maturing. — (univoltine) — Ligyrocoris depict us, L. ca- ricis, Kolenetrus plenus; (bivoltine) — Ligyrocoris diffusus, Peri- genes constrictus, Zeridoneus costalis, Sphaerobius insignis, Pseu- docnemodus canadensis (part). Late Maturing. — (univoltine) — Ligyrocoris sylvestris, Carpilis consimilis, Plinthisus americanus, Stygnocoris rusticus, S. pedestris, Drymus unus, D. crassus; (bivoltine) — Pseudocnemodus canadensis (part). The next group includes those which overwinter as adults — types 2, 4, 5, and 6 — and thus have a spring feeding and oviposition period followed and overlapped by the nymphal feeding period. Here early maturing adults (by June 21), mid-summer adults (by July 15), and late maturing adults (after July 15) may be distin- 46 VOLUME XLIII guished. The bivoltine species have a second feeding period in late summer, and these feeding groups overlap broadly. In contrast to the egg dispause species there is little or no fall feeding. Early Maturing Adults. — (by June 21) — (univoltine) — Trapezonotus arenarius (part) ; (bivoltine) — Emblethis vicarius, Peritrechus fraternus , Megalonotus chiragrus, Pachybrachius ba- salts, Sphragisticus nebulosus. Mid-Summer Adults. — (by July 15) — (univoltine) — Antillo- coris minutus, Scolopostethus diffidens, S. atlanticus, Trapezonotus arenarius (part), Peritrechus paludemaris ; (bivoltine) — Malezono- tus fuscosus, Myodocha serripes, Heraeus plebejus, Cnemodus ma- vortius, Scolopostethus thomsoni, Eremocoris ferus. Late Summer Adults. — (after July 15) — (univoltine) — Xesto- coris nitens, Cryphula trimaculata ; (bivoltine) — Pachybrachius al- bocinctus, Sisamnes clavigera, Ptochiomera nodosa, Ozophora pic- turata. While some species simply become sexually mature in spring at different rates or have different temperature responses, in a few such as Myodocha serripes the photoperiod control continues to operate in the spring. In others, especially and significantly the woodland Drymini, but also Xestocoris nitens and Cryphula trima- culata, although the adults mate in early spring and oviposition can occur under short photoperiods, the nymphs appear in the field very late. It seems most probable that this is the result of the cool forest conditions throughout the spring which cause a very slow egg de- velopment. In Xestocoris and Cryphula a similar explanation may be warranted for the eggs of these species develop especially slowly. Relation to Distribution As might be expected some good correlation exists. All the spe- cies of Type 1 (univoltine egg diapause) except the Drymus spp. have boreal distributions. Among the species of Type 2 (univoltine, adult diapause) all but Peritrechus paludemaris and Cryphula tri- maculata have generally boreal ranges and Scolopostethus atlanticus is intermediate. In Type 3 (egg diapause, adult bivoltine) all spe- cies are boreal or intermediate in range. In Type 4 and 5 none of the species have boreal ranges, and those with the weakest diapause are among the austral element except Ptochiomera. The species of Type 6 (no diapause) appear unusual in that while one of the species is austral, the other two have ranges which extend far north. These latter two, Eremocoris ferus and Scolopos- tethus thomsoni, are cold adapted and perhaps like the arctic midge larvae that are merely quiescent (Lees 1955) no synchronization is 47 ENTOMOLOGICA AMERICANA required for survival. Indeed, these are entirely successful species. Relation to Systematics Lees (1955) said, that while some general trends can be ascer- tained in systematic relationships, in most short-lived insects there is little constancy in the stage of arrest. This, he emphasized, indi- cates that the diapause states had evolved independently, and their selective value is little affected by the stage in which it occurs. This last consideration does not appear to be entirely true, for all the cold-adapted Myodochini genera have evolved an egg diapause and many independently, it appears, but on the other hand, apparently none of the extensive Palearctic fauna of Gonianotini, Megalonotini, and Rhyparochromini have evolved an egg diapause. Such an egg diapause was found elsewhere only in Stygnocoris and in the Nearc- tic Drymus and Plinthisus americanus. Within the Myodochini all the members of the related New England Ligyrocoris group includ- ing Zeridoneus, Sphaerobius, and Perigenes diapause as an egg. This certainly must indicate a systematic relationship to diapause evolution. Relation to Ecology and Brachyptery As noted earlier, there is a direct correlation between habitat permanence and brachyptery, and it was suggested that this faunal element, since it is essentially composed of climax or serclimax spe- cies, was closely adapted to the New England climate. The life cycles further bear this out: most (77%) of the strictly univoltine species with an obligative diapause are largely brachypterous, indi- cating at least a semi-permanent environment. In converse, of the bivoltine species, only 29% are brachypterous. Actually overwintering as an egg also is a definite biological risk, requiring that a permanent habitat for the next year be present at the place of oviposition. There is again a correlation of 67% be- tween permanent habitats and egg diapause and the exceptions are all bivoltine species except the introduced Stygnocoris. All the brachypterous species which overwinter as eggs are found in perma- nent habitats. Most of the species, 12 of 13 (93%), with a low level of macropters (Type 3 and 4) are univoltine or mostly so ( Pseu - docnemodus) . Sisamnes again is the only exception. All the spe- cies with unusual types of brachyptery (see Brachyptery) are uni- voltine. As was discussed earlier, the serclimax or semipermanent vege- tation is usually sparse and poor, indicating a low biomass produc- tivity, and it was proposed that brachyptery may function as an economical adaptation. Also, this may conversely explain the fair 48 VOLUME XLIII abundance of macropters in similarly permanent, but seed-produc- tive woodlands. It therefore appears possible that univoltinism could represent a similar adaptation to a poor environment, and again a good relation exists. Of the seven species found on the dry sparse serclimax, all are univoltine. Malezonotus may represent an exception as the single station, although temporary, resembles a serclimax. An adaptation to low biomass productivity may be as important a fac- tor in the evolution of brachyptery as an adaptation to climax per- manent habitats. These poor open habitats may become very hot, as hot as an open ruderal site. In the northern open Andropogon association, in a more productive habitat, a bivoltine life cycle can be afforded as in Sphaerobhts. A univoltine life cycle may be prolonged over most of the sum- mer or the peak developmental period placed early or late in the summer. In Xestocoris, a boreal species, and Cryphula, an austral species, the univoltine generations do not appear until mid- July ; Ligyrocoris depict us and L. car ids become adult in late May and early June and Carpilis becomes adult from late June to late July. A bivoltine life cycle on the other hand must be dependent on a seed source early in the year, usually fallen seeds from the previous year, and also upon new fallen seeds in late summer. In Ligyrocoris diffusus, at least, an active migration period exists at the time of the early new adults to seek out food sources. This need correlates with the frequent occurrence of the bivoltine myodochines at lights, and the macropterous condition of many bivoltine rhyparochromines. It is then apparent that a strong relationship exists, as South- wood (1926a) predicted, among the ecological requirements, habi- tat permanency, migration, brachyptery, and the seasonal life his- tories. It is moreover apparent, that there are definite assemblages of species in the different habitats. But before we can consider these, other aspects of their general biology should be considered. Protective Coloration and Behavior This subject certainly forms an important part of the ecology of the rhyparochromines. However, it is difficult to generalize from human perceptions on the nature and function of color and move- ments (Cott 1940, Klopfer 1962) to the perceptive realm of verte- brate and invertebrate predators and also parasites. It is extremely difficult to observe predation on the ground layer, and caged preda- tors and prey are unlikely to represent the natural or open ecological situation (Cott 1940). 49 ENTOMOLOGICA AMERICANA TABLE 4 Summary of Various Biological Information Species Macroptery Habitat Dispersion Generations Overwintering Form Diapause Summer Diapause Diapause Intensity M. serripes 1 Ti LD 2 A F — 3 H. plebejus 1 T, L 2 A F - 1 Pa. loasalis 1 Tx LBA 2 A F - 2 Pa. albocinctus 1 P LA 2 A H - 0 C. mavortius 2 t2 L 2 A F - 1 P. canadensis 3 t2 B 1(2) E F - 3 S. insignis 2 t2 — 2 E F - 3 L. diffusus 1 T, LB 2 E F + 2 L. depictus 4 t2 — 1 E 0 - 3 L. sylvestris 4 T2 — 1 E 0 - 3 L. car ids 4 P — 1 E 0 - 3 Z. costalis 1 Ti L 2 E F .+ 3 P. constrictus 1 Tx LA 2 E F - 2 Pi. nodosa 9 LU Tx LA 2 A F + 1 3 S. clavigera 4 To — 2 A F + ! 1? C. consimilis 4 t2 — 1 E 0 - 3 K. plenus 1 To B 1 E 0 + 2 P. americanus 4 P - — - 1 E 0 - 3 A. minutus 2 P D 1 A 0 - 3 A. pilosulus 1 P AL 1 A 0 ? 3 C. trimaculata 3 To — 1 A 0 - 2 X. nitens 3 To — 1 A 0 - 3 0. picturata 1 P L 1(2) A F a 1 St. rusticus 2 Tx — 1 E 0 + 3 St. pedestris 1 Tx — 1 E 0 + 3 I), unus 1 P — 1 E 0 + 3 I). crassus 4 P - — 1 E 0 + ! 3 S. diffidens 3 P — 1 A 0 - 2 S. atlanticus 2 To — 1 A 0 - 2 S. thomsoni 2 To — 2 A H - 0 Er. ferns 1 P DB 2 A H - 0 P. f rat emus 1 Tx L 1(2) A F - 3 P. paludemaris 1 P • — 1 A 0 — 3 50 VOLUME XLIII b£ xn a> ft o cd o "53 xn o a o i-Q og ft 15% macropterous, 3 — 1 5-5% macropterous, 4 — <5% macropterous Habitat: P — permanent, Ti — brief temporary, T2 — persisting* tern.' porary Dispersion: L — lights, B — beach wash, A — airplane sampling, D — * diurnal Generations : number per year ; overwintering form, A — adult, E — ■ egg Diapause : F — facultative, 0 — obligative, H — quiescence Summer diapause : 4- ; diapause intensity, 3 — strong, 2 — moderate, 1 — weak, 0 — none The apparent colors, forms, and motions of the rhyparochro- mines vary remarkably, a variation which must stem from some se- lective basis, although the patterns can also reflect a systematic re- lationship especially in the coloration of the nymphal forms. Sig- nificantly the coloration pattern nearly always correlates with the field behavior of these animals. Moreover, there has been some im- pressive recent experimental work on the effectiveness of lepidop- teran mimicry (Brower 1958, Sheppard 1959) and on predation by jays and toads (Brower 1959, Brower and Westoff 1960). Isely (1938) was able to show the value of protective coloration in grass- hoppers against predators. Cott (1940) effectively amassed a great deal of evidence in support of protective coloration and behavior. It therefore would appear reasonable to consider the variable pro- tective patterns of the rhyparochromines as important biological factors and also important in evaluating the problem of several spe- cies existing in the same habitat. ENTOMOLOGICA AMERICANA Odors In any consideration of the meaning of the protective coloration, a paramount factor is the presence, as in most Heteroptera, of odor- ous or repellent glands in all the species, both nymphs and adults. In all the species and all instars the insects have a burning acrid taste which is quite independent of variations in the odors of the insects. The burning sensation of a minute first instar is quite re- markable. With few exceptions the odor of the rhyparochromines is the heavy “ buggy” odor so frequently associated with Pentato- midae. The exceptions, however, were remarkably different. In the lethaeines Xestocoris nitens , Cryphula trimaculata, and Antillocoris minutus, the odor was strongly reminscent of the peculiar Tapinoma ant odor. Eremocoris ferns and at least one population of Scolo- postethus thomsoni have a distinctly sweet fruity odor. It is note- worthy that the acrid taste remained constant although the odor varied. Significantly, from an evolutionary point of view, in Cry- pliula, Xestocoris , and Antillocoris both the nymphs and adults, with their quite independently derived scent glands, shared the same peculiar odors, while in Eremocoris and Scolopostethus the adults and nymphs had different odors. There is a distinct and remarkable type of scent gland in nymphs which possess the Y-suture described by Slater and Sweet (1960). The anterior gland between terga III and IV is white and apparently of a tubular nature, in contrast to the bright orange walls of the posterior two scent glands and the adult metathoracic glands. In groups without a Y-suture, the anterior gland when present, resembles the posterior two. The anterior gland is absent in the Gonianotini and in Antillocoris. The Y-suture is present in every instar except the first and thus allows the first and second instars to be distinguished in species with the Y-suture. It is appar- ent that the significance of these different scent glands should be pursued further. Remold (1962) has effectively and clearly shown that the scent gland secretion of various heteropterans not only repels, but acts as a temporary paralytic poison upon ants and carabid beetles. The ac- tion is that of a nerve poison, and may paralyze an ant ( Lasius ) for 15 minutes. He found that various lygaeids utilize all the known methods among the Heteroptera for distributing the secretion. Some spread it over their own bodies (nymphs of Nysius and Hene- taris), place the secretion with their legs or tarsi directly on the aggressors ( Lygaeus nymphs, Geocoris ), or release it slowly as a protective odor (Lygaeus adults), but all 17 rhyparochromines representing 6 tribes investigated by Remold spray the secretion 52 VOLUME XLIII directly at the aggressors. In the present study, none were seen to spread the secretion with their legs. For the most part, the species did not release their odor until handled. Indeed, some, for example Antillocoris, do not re- lease the odor until nearly crushed. Cnemodus, and especially Eremocoris ferns, release their odors very readily, but none release the odor a little at a time as do Spilostethus saxatilis (Scop.) and L. equestris (L.) (Remold 1962). With the effectiveness of the scent gland secretions against ants and beetles established, their significance against vertebrates should be followed up. Since, un- like many aposematic plant-living Heteroptera, most adult rhyparo- chromines are procryptic in coloration, it may be that the odor itself may at least function as a warning to mammal insectivores, notably shrews as Rothschild (1962) suggests. She also points out that since bird and lizard predators do not use smell to locate prey, the procryptic coloration may protect against birds, and the odor against the color blind, smell-sensitive mammals. She quotes an example of a coati-mundi which refused to eat procryptically colored pentatomids. The situation is rendered more complex by the nymphs fre- quently being colored quite differently from the adults, and each must be considered separately. First the general color character- istics should be mentioned. These however should be considered with care as they may show considerable systematic relationships especially in the first instars. Of course such resemblances may re- flect a biological similarity Among nearly all myodochines, despite the diverse appearance of the adults, the first instars are pale yellow with a narrow red band across the abdomen. This pattern is also shared by the ozo- phorine, Ozophora picturata. The first instars of the two Mega- lonotini have a yellow band on the first two segments of an other- wise red abdomen. In all known Gonianotini except Emblethis, the first instars are deep red but the later instars are deep black. In the remaining species the color of the first instar varies from red to pink. The color patterns may be grouped into three categories: (1) procryptic or concealing coloration, (2) ant mimicry, (3) aposematic or warning coloration. It is not clear which category some nymphs fall in, partly because white lateral spots on the mid-abdomen may also be interpreted as a disruptive coloration, and partly because the succeeding instars may progressively differ from earlier to late instars. For details see the individual species discussions. 53 ENTOMOLOGICA AMERICANA Procryptic Coloration In nearly all species the adults possess a very evident procryptic type coloration which lends to the subfamily a brown, somber, “sparrow-like” mien, which has contributed to our poor knowledge of the taxon. Nymphs exhibiting procryptic coloration include the black nymphs of the Megalonotini and most Gonianotini, Emblethis, Car- pilis, Perigenes, Ozophora, Pachybrachius, Zeridoneus, Ligyrocoris diffusus, and the last instar nymphs of Eremocoris ; the nymphs of Stygnocoris pedestris and Kolenetrus plenus have disruptive pro- cryptic color, and Myodocha nymphs are longitudinally striped. The species with procryptic coloration fall into several behavior groups. The Gonianotini, the Megalonotini, and Carpilis, Sisamnes and Kolenetrus of the Myodochini tend to hide in narrow crevices and run agilely and rapidly from one crevice to another, rather like a roach. The coloration of Emblethis closely matches its sandy habitat and it moves in short rapid bursts, stopping frequently, which renders it difficult to see. The other group includes the long-legged, rapid running adult myodochines of open places which are not ant mimics such as Li- gyrocoris, Zeridoneus, and Perigenes. The third group includes the species of rank habitats and loose litter, and which accordingly may have generalized legs such as Peritrechus fraternus, Stygno- coris spp., Drymus, Pachybrachius, Antillocoris, and also a few long-legged myodochines like Myodocha. Some species of rank habitats characteristically move slowly and deliberately, in contrast to the roach-like scurry of the gonianotines. These include Myo- docha, Heraeus, Antillocoris, Peritrechus paludemaris, and Plinthi- sus americanus. Ant Mimicry This may be a more controversial category to interpret, but it is especially marked in the nymphs of some species. In others, both nymphs and adults, the appearance of ant mimicry “grades” into procryptic disruptive coloration. Field observations, however, are invaluable here, for the ant mimics move with an exaggerated jerky hesitant gait much like ants. The species are listed in order of re- semblance to ants. Adidts. — Sphaerobius insignis, Cnemodus mavortius, Pseudoc- nemodus canadensis, Eremocoris ferus, Scolopostethus spp. Nymphs. — above, but only early instars of Eremocoris and late instars of Scolopostethus, Pachybrachius albocinctus, P. bilobatus, Ligyrocoris depictus, L. sylvestris, L. caricis, and up to fourth in- stars of L. diffusus. 54 VOLUME XLIII The most remarkable ant mimic is Sphaerobius insignis. Fre- quently this insect was overlooked in the field for it ran actively about and was very difficult to distinguish from a large Formica ant. This species is of considerable interest for it is polymorphic in color (see Sphaerobius) with few intermediates, and so resembles both black and red ants. The fifth instars, however, are all red, not black, and were the better mimics. Such dimorphism (Sheppard 1959) which is adapted to two models should greatly enhance the protection from mimicry resemblance. Nearly its equal as a mimic although much larger is Cnemodus mavortius. The adults, especially the brachypters, resemble huge black ants, the nymphs, red ants. In the other mimics the resemblance is not so striking, but fre- quently this is not necessary with many predators, as only a gen- eral resemblance is required (Klopfer 1962). To a certain extent the predominance of myodochine ant mimics may result from the predominance of these preadapted long-legged, rapid running forms in open dry habitats. These habitats are espe- cially favored by ants. The ‘‘mimicry’’ by Eremocoris and Scolopostethus is of a differ- ent sort. These species tend to form loose aggregates under litter in favorable biotopes, and frequently, disturbing the litter causes the whole group to ‘ ‘ boil up ’ ’ and actively run in all directions as when an ant nest is disturbed. A difficulty is that the ant resemblance, is not so obvious although the color pattern is very clearly marked in these genera. However, if this is a defense against bird predation, the constricted rapid moving aspect of the insects may form a simple sign stimulus so frequent in bird behavior (Klopfer 1962). The obvious question is, if this is mimicry, is it Batesian or Mul- lerian. It is realized that the two types may grade into each other (Huheey 1961). While it may be assumed that the Rhyparochro- minae may be distasteful because of their scent glands (Remold 1962), no conclusion can be drawn until the potential predators are determined, along with their reaction to the taste of the rhyparo- chromines. The difficulty with a Mullerian interpretation might be the absence of clearly marked warning coloration schemes among other non-ant mimic rhyparochromines. If it is Batesian the abun- dance levels as compared to the ants is often remarkably high which would allow, say, some bird predator to see through it. The final aspect, which may suggest a combined Batesian-Mullerian explana- tion, lies in the remarkable tendency for the nymphs to be ant mimics, the adults procryptic. Moreover, some of the nymphs ap- pear to be aposematic. The possibility suggests itself that the nymphs may form a Mullerian assemblage with ants, the adults, a 55 ENTOMOLOGICA AMERICANA Batesian assemblage. The different position of the scent glands in the nymphs and adults may be significant here. The red aposematic coloration would here, of course, be mean- ingful only to a bird predator, as the mammals are color-blind and most arthropods would not perceive red, and in which case the in- sect may appear black. These are all nymphs. The brightest and most conspicuous are the striking red nymphs of Drymus spp. and Stygnocoris rusticus. The nymphs of Ptochiomera and Sisamnes are similar, a bright contrasting red and white. Finally, the most generalized rhyparochromines have bright light pink nymphs — Plinthisus, Antillocoris, Cryphula, Xestocoris as well as Peritrechus fraternus. It is noteworthy that all are loose litter dwellers, and none are long-legged runners or crevice seekers. Only four species are known to feign death : the two Stygnocoris, Eremocoris ferns , and Drymus unus. In Stygnocoris this is espe- cially noticeable for when disturbed while feeding above the ground level on composites, the insects will freeze and drop to the ground. Predators and Parasites Predators As mentioned in the preceding section, very little is known about the natural predators of the Rhyparochrominae. Enough, how- ever, is known to indicate a large number of predators. Thomas (1955) considered the important predators in England to be nabids, anthocorids and the centipede, Lithobius. Wilson (1938) has found that Pachybrachius bilobatus and P. vinctus are preyed upon by lizards ( Anolis spp.) in Puerto Rico. Knowlton and Nye (1946), and Knowlton (1944) working in Utah, found that a number of rhyparochromine species were fed upon by the sage sparrow, 4m- phispiza nevadensis (Ridgway). Knowlton, Maddock and Wood (1946) in Utah found rhyparochromines in the diet of the sage- brush swift lizard, Sceloporus graciosus graciosus (Baird and Gir- ard). Miller (1956) reports that certain African lizards consumed Dieuches sp.; Corby (1947) found Gryllodes sigillatus Walker feeding on adults and eggs of Elasmolomus sordidus (Fab.). I have found Ptochiomera nodosa being preyed on by Geocoris uliginosus (Say) ; Pseudocnemodns canadensis by Geocoris limbatus Stal ; Myodocha serripes, Cnemodus mavortius, Zeridoneus costalis by Melanolestes picipes (H.-S.) but not by Nabis roseipennis Reuter from the same environment. Trapezonotus is preyed on by all stages of Pagasa fusca (Stein). In the laboratory the eggs of various lygaeids were readily fed on by roaches. A few other examples are given under the species, but this in no wise is adequate, and merely shows that the rhyparochromines are preyed on by a large assort- 56 VOLUME XLIII ment of predators, and it appears unlikely, as Thomas (1955) thought, that the rhyparochromines suffer an unusually low natural mortality. Probable predators are wasps of the sphecid genera Diploplec- tron Fox and Dryndella Spinola, both members of the subfamily Astatinae, Williams (1946) found that Diploplectron provisioned its nests with nymphs of Emblethis vicarius, Megalonotus chira- grus, and Spliragisticus nebulosus. Parker (1962) notes that there is no apparent host specificity, but rather a size specificity for Heteroptera, the larger astatines capturing pentatomids, coreids, cydnids, or scutelleroids ; the smaller, lygaeids. It appears of some interest that these Heteroptera are all from the Trichophora section of the Pentatomomorpha. Evans (1958) describes the hunting pro- cedure of the wasps as follows, 4 4 The female hunts its prey in weedy areas and is often seen crawling up and down plant stems. When a bug is located the wasp grasps it and stings it to immobility. ’ 7 As emphasized by Wheeler (1910) and Remold (1962), ants form the most abundant and active arthropod predators, especially at the ground level. Remold found that although the different heteropterans, including rhyparochromines, were attacked by vari- ous ants, the ants were warded off by the temporary paralyzing ac- tion of the scent gland secretion. Moreover the secretion also acts as a general repellent. Further work is needed to evaluate the ac- tual effectiveness of the secretion in the field, and to what extent the insects are preyed upon by ants. At any rate this defense mecha- nism would make understandable the coexistence of large popula- tions of lygaeids with ants, and, in turn, also the evolution of ant mimicry among lygaeids and other Heteroptera. Parasites There are only a few records of parasitism among the rhyparo- chromines. Tachinid flies are known to parasitize Scolopostethus thomsoni and Eremocoris plebejus (Fall.) in Germany (Michalk 1935b, 1938a). The tachinid parasite of the Eremocoris was deter- mined as Cinochira atra Zett. (Michalk 1938a), and a species of Chalcididae as also recorded from this species (Michalk 1940). A scelionid parasite has been recorded from the eggs of Gastrodes gros- sipes (DeGeer) (Nageli 1933). Michalk (1938b) found Scolo- postethus affinis (Schill.) parasitized by the larvae of the mite, Allot hrombium. I have found Thrombidium mites on Myodocha serripes and Scolopostethus diffidens. Various rhyparochromines were found to be parasitized by tachinid flies of the genera Catharosia and Petia ( = Procatharosia) . A fairly large complex of Catharosia species appears to be involved. Dr. Paul H. Arnaud, Jr. (in litt.) is presently revising these genera. 57 ENTOMOLOGICA AMERICANA A more detailed presentation of the biology of the flies will be given in a later contribution as a considerable number of new species appears to be involved. A general outline may be sketched. Parasitism occurs at least by the late instars, but the parasite does not leave until the host becomes adult. The parasites leave by either pushing through the ovipositor in the female or through the conjunctiva between the genital capsule and segment eight in the male. The larva crawls a short distance away usually to a wetter site and pupates. Pupation usually takes approximately a week. Ashlock {in lift.) was able to rear Catharosia lust vans from a new species of Eremocoris. The fly developed very rapidly and was reared through its entire life history between April 9 and May 4. Neither of us have observed the type of oviposition by which tachinids are classified biologically. The catharosines were usually reared in the spring or fall, and very few were found in the summer generation of bivolt ine lygaeids. A parasitized female can be readily recognized because it is non- reproductive. The life cycle of the catharosines varies with whether the host species overwinters as an egg or an adult. If in a species which diapauses over the winter as an adult, the parasite overwinters in the hibernating host. If the host overwinters as an egg, then the parasite leaves the lygaeid host in the autumn. The following lygaeids were found parasitized by Catharosia spp.: Host adult { and parasite) overwintering. — Heraeus plebejus, Pachybrachius basalis, P. albocinctus, P. bilobatus, Cnemodus mavortius, Ptochiomera nodosa , Cryphula trimaculata, Peritrechus fraternus, and P. paludemaris. Host egg overwintering. — Pseudocnemodus canadensis, Sphae- robius insignia, Ligyrocoris diffusus, L. depictus, Z eriodoneus cos- talis, and Perigenes constrictus. Only one species, Catharosia nebulosa (Coq.) was determinable, (Arnaud in litt.) but the remarkable size range of the host species and corresponding variation in the size of the parasites, along with the quite different methods of overwintering of populations from adult or egg overwintering hosts, suggests that the species concept of C. nebulosa may be too broad, and warrants further studies. Hibernating Pachybrachius albocinctus were also parasitized by a different tachinid, Alophorella aeneoventris (Will.). Emblethis vicarius was found parasitized by another catharosine, Petia ( = Procatharosia) ? calva Coq., and Ashlock {in litt.) informs me that in California he too reared Petia from Emblethis. Moreover, the Procatharosia parasitized Emblethis were among and near myodo- 58 VOLUME XLIII chines which were parasitized by Catharosia, which further suggests a parasite-host specificity. Petia overwinters in its host, Emblethis vicarius. Both Sabrosky and Arnaud doubt (in lift.) that the tachinids could have such a narrow host range as appears to be indicated by these results. However, from the converse view, the parasitism of the rhyparochromines is very narrow as each species appears to have a specific catharosine parasite. This exploitation by a single taxon of parasites has a parallel in the Miridae which are almost exclusively parasitized by euphorine Braconidae (Leston 1961). The Catharosini are so far known only from the Lygaeidae, and these observations constitute the first host records (Arnaud in lift.). Of the total rhyparochromine fauna of 40 species, tachinid para- sites were found in 16 (40%). Of these hosts 12 (75%) were myodochines which is a significant proportion as the Myodochini (Table 1) comprise only 47% of the fauna. However, another important correlation is that no lygaeids of closed canopy habitats were so parasitized, and nearly all Myodo- chini are found in open habitats. It could be that the western species of Eremocoris investigated by Ashlock (in lift.) is an open area inhabitant like some European Eremocoris. Since Eremocoris plebejus is typically found in open habitats (Penth 1952), and the Catharosini are known from Europe (Arnaud in lift.), but without hosts, the determinations given by Michalk (1938a) should be checked. Since parasites form a major population control factor, it is interesting to relate the distribution of hosts to Gause’s (1934) statement, “When a new habitat is first colonized there is a strong probability that its normal predators, including parasites, will be absent and during the first period after colonization, the population (of the host), if the reproduction rate is sufficiently high, will build up quickly. ’ ’ The first aspect is that the inhabitants of permanent habitats with the significant exception of the salt marsh species Peritrechus paludemaris and the marsh edge species, P achyb rachitis albocinctus , were not parasitized. Parasitism was most pronounced among myodochines of temporary habitats especially the newest habitats. In populations of Ligyrocoris diffusus, Zeridoneus, P achyb rachitis, and Cnemodus, the parasites were found only in well-established colonies, never in new small populations. There is obviously a great need here for further study of preda- tion and parasitism especially its role in population control. In- formation on prey or host seeking and capture is needed to interpret protective coloration and competition among the rhyparochromines. 59 ENTOMOLOGICA AMERICANA Feeding Habits A general account of feeding habitats of the Lygaeidae was given earlier (Sweet 1960) in which it was shown that the Rhyparo- chrominae like most other Lygaeidae were seed feeders and not predacious. Therefore, only the recent literature is reviewed here. Thomas (1955) investigated the feeding habitats of a number of rhyparochromines. He found that the insects would feed on buds, moss, dead and dying insects, and concluded that materials of this sort formed the food sources of the Rhyparochrominae. Moreover, he noted that the Rhyparochrominae laid only few “batches” of eggs and concluded from this that the Rhyparochrominae with their biological success in their protected environment, needed only a low reproductive capacity. Putshkov and Putshkova (1956) and Putshkov (1956) found that most of the Lygaeidae were seed feeders, and their reproduc- tive potential was not low, but high. The work of Johansson (1858a, 1958b) effectively explains the disparity. He showed that in Oncopeltus fasciatus (Dallas), the corpora allata hormone pro- duction was controlled by seed feeding, for starving or placing the insect on a diet without seeds cuts off egg production. I have simi- larly found that on lettuce and other green material only a few eggs were laid, in contrast to the heavy egg production when fed on sunflower and other seeds. Beck, Edwards, and Medler (1958) found that Oncopeltus could not be reared readily upon a defined diet per se, for the insect needs to pierce through a milkweed seed coat to stimulate normal feeding. The seed coat thus appears to be very important, as some rhyparo- chromines react to seed coats, others do not. Most species, however, fed normally on a hulled sunflower seed. The general behavior of the insects during feeding is much as described by Feir and Beck (1963) for Oncopeltus and in Lepto- glossus (Koerber 1963). As in most Heteroptera the labium bends on its joints to allow feeding, and frequently the labium is entirely removed and feeding occurs by the stylets alone as in Leptoglossus (Koerber 1963). After a feeding period, as noted by Feir and Beck, a rhyparochromine always seeks out and imbibes free water. Salivary fluid Miles (1959) showed that Oncopeltus fasciatus like several pen- tatomids and coreids and all Homoptera secretes a salivary fluid which coagulates to form a sheath. In contrast, this fluid is absent in the Miridae, Tingidae, and predacious Hemiptera such as the Anthocoridae, Nabidae, and Reduviidae (Sweet, unpublished). He thought the function of this sheath was to seal the stylet entrance and line the bores to avoid wastage of the salivary-food solution. 60 VOLUME XLHI Saxena (1963), however, has shown that in the pyrrhocoricl Dysdercus, the sheath fluid is deposited only when piercing a tough barrier. lie found no loss of salivary fluid from the junction of the labium with the seed whether the sheath was present or absent. From the feeding behavior and conditions of sheath deposition, Saxena concluded that the sheath functions to affix and stabilize the beak during piercing, and unlike Oncopeltus (Miles 1959), it was not deposited along the path of the stylets in the seed. Saxena was also able to show that the salivary fluid in Dysdercus did not preorally digest the seed kernel. Instead, a suspension was formed along the path by the mechanical action of the stylets. Miles (1959) on the other hand thought preoral digestion occurred in the salivary fluid-filled feeding bores. Saxena suggested that the rapid saliva ejection-ingestion process of Dysdercus also occurs in Oncopeltus and thought Miles’ conclusions need further validation. Saxena moreover demonstrated that digestion occurred entirely within the gut in Dysdercus. This resolves the dilemma imposed by the slow action time of the enzymes in vitro as was observed by Nuorteva (1958) and Baptist (1941). I can only add that the salivary sheath cone was not formed when feeding on the freshly broken sunflower kernel but was formed upon the thin membrane (true seed coat) that surrounds the seed. Upon hard substrates the salivary cones may be deposited on bores already made, even upon a cone already present to the extent that as many as five cones have been observed on top of one another at a feeding site in P achybrackius basalis cultures. Cone formation may then be inhibited by a direct food or water stimulus rather than stimulated by surface hardness as Saxena suggested. At any rate, the salivary cone is more than a mere fleck of dried saliva as thought by Koerber (1963). I found the salivary sheath fluid in all the Khyparochrominae studied and throughout the Lygaeidae, including the predacious Geocorinae. Moreover, the predacious asopine Pentatomidae (. Podisus , Perillus ) secreted this fluid very copiously. Alydids and rhopalicls which can also be reared on sunflower seeds, similarly secrete a sheath fluid. It will be most interesting to trace further the presence of this fluid, which Miles (in litt.) considers to be strictly homologous in the Homoptera and the Pentatomomorpha. Food Plants In my 1960 paper I implied that the Lygaeidae were unusual in their seed feeding habits, but it appears that this tendency is wide spread, especially in the Trichophora families, the Pyrrhocori- dae, Rhopalidae, Alydidae, Stenocephalidae, (Southwood and Leston 1959), and common in the Pentatomidae and Coreidae (Les- ton in litt.). Nevertheless, the term “seed bugs” is still applicable, 61 ENTOMOLOGICA AMERICANA as this family is, in general, highly adapted to feeding on fallen loose seeds. As stated previously (1960), the rhyparochromines are rather oligophagons on seeds which makes the task of ascertaining their preferred seeds, if any, difficult. Some definite patterns emerge. P achybrachius basalis is very partial to Paspalum and Panicum seeds, P. albocinctus to Carex and Scriptus seeds; Peritrechus to Panicum , but also many others, also P . paludemaris for Spartina seeds ; Myodocha showed a preference for Hypericum and straw- berry seeds; Pseudocnemodus for V accinium and Festuca ; Cnemo- dus, Sphaerobius, and Sisamnes for Andropogon seeds ; Carpilis for Veronica seeds; Zeridoneus, Ligyrocoris diffusus, L. depictus, and L. sylvestris for composite seeds, especially Budbeckia, the latter also for Betula and Tsuga seeds ; L. caricis for Carex seeds ; Ptochio- mera for Rumex seeds, etc. ; Ozophora for Aster seeds ; Plinthisus for Tsuga, Picea, and Betula seeds; both species of Stygnocoris for composite seeds, especially Tanacetum ; Cryphula for Panicum and other grass seeds ; Xestocoris for Festuca , Danthonia, and Andropo- gon seeds ; Antillocoris for Betula populifolia seeds ; Trapezonotus for many seeds especially V accinium; Megalonotus for Centaurea; Sphragisticus for Rumex and Agropyron seeds ; Drymus spp. for Aster seeds; Eremocoris has a very wide seed range ; S colop ostethus atlanticus for V accinium and Viburnum seeds; Scolopostethus dif- fidens for Betula, Tsuga, and Picea seeds ; S. thomsoni A for Rumex seeds, etc. ; S. thomsoni B for Carex seeds ; Emblethis for Andropo- gon, Carex, and composite seeds. There is not any well-marked host specificity, bnt frequently as shown above there are preferences for certain seeds from the native habitat that may help to explain niche overlaps in the natural habitats. Frequently, in additions of these seeds may ameliorate the high mortality in the first instar. The later instars of most species as Putshkov (1956) found, are practically polyphagous on seeds although often retaining a preferential for a particular species of seed. This preferential was repeatedly observed in the feeding behavior. While I earlier (1960) could find no reference to insect predation among the rhyparochromines it has recently been found that the genus Mizaldus preys on beetles and on meal worms in grain respec- tively (Miyamoto 1955, Slater and Carayon 1963). As the latter authors point out, these insects may also feed on seeds much as does Geocoris (Sweet 1960). Poppius and Bergroth (1921) report that Xenydrium formiciforme Berg, is a predator on eggs and larvae of the ponerine ant Ectatomma ruida Boy. However it is not clear whether this was inferred from ant mimicry as at that time mimicry 62 VOLUME XLIII was considered closely associated with predation on ants (Wheeler 1910) and predation assumed. Evles (in litt.) has been able to show that Scolopostethus decor a- tus may, when starving, attempt to feed on animal food, but makes poor growth and oviposits very little upon such a diet. In my earlier paper (1960) I neglected to mention that when starving or in thirst, some of the species will attempt to scavenge upon dead or dying insects. However there was no oviposition, no or little nymphal growth, and great mortality even when an abun- dant supply of dead insects was kept available. The species which expressed such scavenging behavior were : Pachybrachius basalis, Pseudocnemodus canadensis, Cryphula trimaculata, Ozophora pic- turata, Stygnocoris pedestris, and Eremocoris ferns. It should again be emphasized that these insects made no attempt to attack live prey, nor did they feed on the eggs of other rhyparochromines. Symbionts It has been known since the early work of Glasgow (1914) that the phytophagous Heteroptera of the group Trichophora, including pentatomoid, lygaeoid, and coreoid families, all have bacterial caecae or mycetomes connected to or associated with the posterior or fourth part of the mid-intestine. Other heteropterous families may have different types of mycetomes. This subject including the transmis- sion of the symbionts has been reviewed by Carayon (1952). In some Pentatomidae the symbionts are smeared on the surface of eggs and the new hatched nymphs become contaminated by feed- ing on this after hatching. In others, as shown by Glasgow (1914), the infection occurs in the embryo. Among the Rhyparochrominae there does not appear to be any work upon the transmission but in other lygaeids as Ischnodemus sabuleti Fallen and Nysius spp. (Schneider 1940) the infection occurs within the egg before shell formation. In no instance when hatching was observed were the new nymphs ever observed to feed on the surface of the egg shell. Detailed work on this important biological subject was considered beyond the scope of the present study. Nevertheless all the rhyparochromines in the present study had the fingerlike groups of bacterial caeca as described by Glasgow (1914). As he indicated, but never carried through, the caeca configurations are species-specific and each species is inhabited by bacteria which are apparently host specific ( at least in form ) . Clearly, a careful study of the anatomy of the caecae, the types and relation of the bacteria to the host, and the methods of transmission should prove of great value in elucidating evolutionary patterns among the Lygaeidae, and of course, in other 63 ENTOMOLOGICA AMERICANA families. As indicated by Glasgow (1914), the presence of the caeca are somehow related (in the Pentatomomorpha) to a phytophagous feeding habit, as the predacious species lack them. Slater and Carayon (1963) have shown that the presence of bacteria is unstable in the predacious rhyparochromine Mizaldus. Economic Importance While it is true that very few (none in New England) of the rhyparochromines are economic pests, these insects may be of some economic importance as beneficial insects, especially those species adapted to field and garden habitats. It follows from their fre- quent high populations, ground level biotope, and seed feeding habit that these insects must be an important ecological factor in the destruction of fallen seeds and fill an important niche in the natural biota. This destruction would result not only from feed- ing but in piercing the seed coat, for probably fungal pathogens such as Fusarium and Pythium may more readily gain entrance. Moreover seeds deeply planted as in conventional agriculture would not be disturbed, while weed seeds on the surface of the ground would be susceptible to the ground living seed bugs. Thus for the most part such seed feeding would be beneficial. Behavior Seed Defense Many of the rhyparochromines exhibited remarkable behavior patterns relating to seeds. It was in the observation of this be- havior that the function of the fore-femora became evident. Only under what would be called dirty culture conditions could this repertoire be expressed, and sometimes only over certain seeds and when the bugs were allowed to become hungry. This behavior was displayed in essentially two ways. In the first, and this appears to be common to all Rhyparochrominae, the seeds are dragged or moved from exposed open sites into protected sites. This is very likely a thigmotactic response, the insects responding positively to contact with the substrate around them. Perhaps the function of the long abdominal and head trichobothria is to sense such contact, much after the analogous example of a mouse’s vibrissae. Tullgren (1918) thought these hairs may be auditory organs, but so far no evidence is at hand for such a func- tion. A thigmotactic function is consistent with modern concepts of orientation in arthropods (Fraenkel and Gunn 1940, Carthy 1958). This behavior was long ago noted in Elasmolomus sordidns by 64 VOLUME XLIII Maxwell-Lefroy (1909) who wrote, it has “been found to infect threshing floors and to carry off the wheat grains to the margins of the floor and hide them. What nourishment they can extract from a dry wheat grain seems doubtful, unless their salivary excre- tion has solvent powers, but they carry off the grain so abundantly that the cultivators require to collect them again every morning.” Ashlock (1958) observed one individual of Ligyrocoris latimargina- tus to pick up a seed between its fore legs and run with it. In clean cultures such a use of the fore legs was never observed. The insects would sense a seed, feed on it for a variable period, and then drag the seed by the labium, if large, or suspend it under the body affixed to the labium, if small, and carry it to shelter. But if the culture was full of litter and obstacles, the fore legs wTere clearly observed to aid in moving the larger seeds. The fore legs were placed over the seed and the seed was lifted over the obstacle. From the diverse and varied shapes and sizes of fore femora in the Lygaeidae, such a tool-like function may have been expected. But in no case was the seed observed actually being carried in this position. The second behavior response is one of direct defense. This frequently approaches a territorial sort of behavior, and is such at least in Pachybrachius bilobatus. Only in certain species was such an extreme intraspecific competition behavior seen, although it may perhaps be elicited in all species under the proper conditions. lit its most simple form as seen in most species, the seed is kept out of the aggressor’s reach by the defender keeping himself between the seed and the aggressor. The several successive stages of seed defense behavior are as follows : a vigorous wagging of the antennae as in Eremocoris and Drymus; then, a defense dash at the aggressor; and a clash with wildly flailing legs and antennae as in Ligyrocoris diffusus and Scolopostethus diffidens; finally the defense behavior reaches its highest expression in Pachyb rachius when the fore femora are spread stiffly and the two, the aggressor and the defender, stand upright with antennae and legs flailing. This final behavior pattern was only observed in Pachyb rachius basalis, P. bilobatus, and (?) Sisamnes contractus (Dist.) ( = Exptochiomera antennata V. D.). In P. bilobatus the seed defense or territoriality is involved in sexual behavior, and the males which are considerably larger than the females defend the seeds from other males. It is significant that the species which display this defense be- havior most vigorously are also those species which frequently exist in high local abundances in the natural habitat. These behavior patterns may play an important role in intraspecific and interspe- cific competition. 65 ENTQMOLOGICA AMERICANA A remarkable aspect is the ability of the rhyparochromines to relocate a particular seed after a defense display. The behavior patterns are described in detail under the individual species. Mating Behavior The individual species accounts should be consulted for details but, several basic mating behavior types are present: (1) Those which involve a shaking dance by the male. In these species there' is a stridulitrum (c.f. Ashlock and Lattin 1963) on the abdomen and a plectrum on the hind femora: Ligyrocoris spp., Sphaerobius insignis, Zeridoneus costalis. (2) Male employs a fore femora activity, ( stridulation ? ) and vibrating antennae: Pseudocnemodus canadensis. (3) The male recognizes the female before contact, and leaps suddenly on her : Stygnocoris species, Sphragisticus nebn- losus, Megalonotus chiragrus, and Plinthisus americanus. (4) The male vibrates the antennae rapidly near the female and climbs upon her deliberately: Pachybrachius basalis, P. albocinctus , Myodocha serripes , Heraeus plebejus, and most other species. The known species which employ the dance and stridulating behavior are all long legged myodochines of open habitats. Cleaning Behavior Another interesting behavior pattern involves cleaning behavior. This is readily elicited by shaking the bugs with debris, but also occurs frequently after feeding. First the labium is pulled through the fore tibial cleaning brushes, and then the antennae are similarly cleaned. Second, the fore tibia are rubbed together, and then one of the tibial brushes is brushed against the maxillary plate (lorum), and the first labial segment. With each fore tibial stroke the labium is partly opened. Thereafter the fore tibia are rubbed against the mesotibia and the meso- upon the metatibia. The metatibia and tarsi are next rubbed against the sides of the abdomen, and frequently with each other. Often at this point the genitalia are groomed. Frequently this sequence is repeated rapidly on each side always beginning with several strokes against this particular site on the base of the labium. This grooming be- havior has been observed in second to fifth instar lygaeids as well as in the adults. Parsons (1958) has summarized the literature on the presence of large cephalic glands in water bugs and described a similar gland in Gelastocoris. These glands have been found in the pyrrhocorids Dysdercus (Macgill 1947), Pyrrltocoris (Bugion and Popoff) ; and the lygaeids, Oncopeltus (Linder and Anderson 1955), and Mizal- dus, Ischnodemus and Dimorphopterus (Slater and Carayon 1963). 66 VOLUME XLIII Except in Gelastocoris (Parsons 1958) the glands in aquatic Heter- optera and Pyrrhocoris open on the margin of the maxillary plates at the base of the labium or at the labium. In the lygaeids, Oncopel- tus and Mizaldus, the orifices of the glands open in the preoral cavity. I have found similar glands in several other rhyparo- chromines. According to Parsons (1958) three functions have been sug- gested for these glands: (1) subduing prey; (2) defense against predators; or, (3) excretory organs. The last was shown to be untenable and in Gelastocoris, the posterior position of the glands made unlikely any poisonous action. Slater and Carayon suggested that the glands in Mizaldus secreted a toxic substance that paralyses the prey. Both Dysdercus and Oncopeltus exhibit the same sort of cleaning and grooming behavior seen in the rhyparochromines, and re- peatedly stroke their fore tibia against the maxillary plate-labial segment one region. It is here suggested that the secretion of the cephalic gland is spread over the body during the grooming process. It would be most interesting to ascertain the exact role of this secretion. Per- haps the secretion has a species-specific odor which has a role in sexual and aggregational activities. Nymphal Development and Reproduction In this section are considered the essentially developmental and reproductive aspects of the life history. All the New England species are oviparous. Carayon (1961), however, has found that the African rhyparochromine Stilb ocoris is ovoviviparous. Nymphal Instars As in other Heteroptera (Southwood and Leston 1959, Weber 1930) all the species have five distinct instars and none have the exceptional four instars associated with prothetely as in Dolichona- bis limbatus (Dahl.) (Southwood 1961b). The instars may readily be distinguished by the relative development of the wing pad (Guide 1919, Slater 1951, Putshkov 1958, Southwood and Scudder 1956). As Slater (1951) indicated, the first and second instars are frequently difficult to distinguish. The difference in length of the meso- and metathoracic segments used by Southwood and Leston (1959) is often difficult to ascertain and is not invariable (Putsh- kov 1956). In many rhyparochromines especially those of the tribes Myodochini and Ozophorini, the first and second instars undergo a striking change in color. Moreover the Y-suture is always absent in the first instar. In Eremocoris ferns, Scoloposte- 67 ENTOMOLOGICA AMERICANA thus atlanticus, and Malezonotus fuscosus the first instars have pale terminal antennal segments. The changes in color in Palearctic lygaeid species during the nymphal ontogeny are summarized by Putshkov (1956). The color patterns have already been alluded to under protective colora- tion, and only the ontogenetic changes are considered here. Accord- ing to Putshkov most of the species of the other subfamilies differ from the Rhyparochrominae (except EmMethis and Gonianotus) in that the body is generally pale colored and the basic color pattern appears in the second instar. In the Rhyparochrominae with the exception of EmMethis and Gonianotus the nymphs are either dark or contrastingly colored with the head and thorax dark, the abdo- men light. Putshkov distinguished four types of color changes each beginning with a red or pink first instar nymph. These four groups are largely adaptable to the New England fauna especially in genera also present in the Palearctic. A fifth type distinguished by Putshkov includes species with a pale band on abdominal seg- ment one to three in the first instar. This band gradually disap- pears in later instars. Only Megalonotus and Sphragisticus show this color pattern in New England. For most of the Myodochini and the known Ozophorini a sixth type must be noted. In this type the first instar is pale yellow with a narrow red band across segment four. In the second instar there is an abrupt color change. This includes all the New England Myodochini except Perigenes which has a dark red first instar and Kolonetrus which is unique with an entirely pale yellow abdomen. Into Putshkov ’s four types the other New England genera fit as follows: (1) Coloration remains nearly unchanged through nymphal development : Xestocoris, Plinthisus, Antillocoris, Drymus, Stygnocoris. (2) Coloration becomes more complex but not black: Cryphula , Peritrechus, Scolopostethus, Eremocoris, Perigenes. (3) Coloration becomes lighter — no Nearctic rhyparochromine fits here except perhaps some specimens of Drymus. (4) Coloration be- comes a deep black although the subcuticular layer remains red. These lack the Y-suture and include the megalonotines Megalonotus and SpJiragisticus as well as the gonianotines Malezonotus f Delochi- locoris, and Trapezonotus. As summarized by Putshkov there are very few structural changes during development in the Lygaeidae, each instar differing mostly in allometric changes in head size and appendage length, the head decreasing in relative size, and the appendages increasing in length, but to a variable extent, resulting in long legged and short legged genera. There is often an increase in pilosity and on the head there is frequently an increase in major setae. However, 68 VOLUME XLI1I in the first instar, much as in lepidopterous larvae (Fracker 1915) only the primary long setae (head trichobothria) are present like the abdominal trichobothria. Aside from the Y-suture appearing in the second instars and the appearance of a vestigial scent gland plate between segment three and four in Antillocoris, the various structures remain largely unchanged in nymphal ontogeny. The spiracles and trichobothria and sutures are much as they are in the adult. The known lygaeid nymphs are distinctive, compared to the Coreidae and Pentatomidae, in that their generalized smooth abdominal cuticle nearly always (except Malcinae) lacks spines and protuberances and rarely has punctures (Putshkov 1956), never in the Rhyparochrominae. Nymphal Development The details of the development periods are discussed under the individual species. There is no obvious relationship of develop- ment rate to size, the minute species of Antillocoris only 2 mm. long takes approximately five weeks to reach maturity, a period much longer than that of the large species of the Ligyrocoris group or the largest rhyparochromines in New England Perigenes and Cnemodus which are 8 to 9 mm. long. In general, as indicated under the seasonal cycle discussion, the development rates show adaptations to the habitat type and the type of seasonal cycles. The relation to temperature is only partial. Forest species de- velop fairly slowly in the egg state, but in the laboratory and dur- ing nymphal development in the field, growth proceeds rapidly. Xerothermic species like Carpilis and Xestocoris develop slowly while the cool adapted Eremocoris develops rapidly both in the field and laboratory. Species adapted to diversified forb habitats with a large seed production (these are all short lived subclimax habitats) have rapid life cycles which take only about three or four weeks and such species usually have bivoltine seasonal cycles. On the other hand species of habitats with low productivity, that is, the sparse open barren old sites, develop more slowly as do Carpilis and Xestocoris and are univoltine. These development patterns hold under labora- tory conditions of similar food and temperature, are not related to size, and so indicate a genetic difference. Reproduction A similar relationship is apparently present in the reproductive capacities as measured by oviposition capacity in the laboratory. However data on this aspect must always be treated with some 69 ENTOMOLOGICA AMERICANA skepticism because of the close control that seed feeding holds over egg production (See Feeding Habits). In the natural habitat, given this physiological mechanism, egg production would be con- trolled by the availability of seed food. However r, the intrinsic rate of increase or the reproductive rate (Andrewartha and Birch 1954) under optimum conditions is fairly high as shown by Putsh- kov and Putshkova (1956) rather than low, in the order of a few tens of eggs as thought by Thomas (1955). Thomas’ data, as indicated under feeding habits, is readily replicated by transferring these insects from seeds to foliage diet. Indeed the egg production figures given by Putshkov and Putshkova (1956) appear a little low (55-85), but as the authors state, this may result from the short term study in the rearing cages. Nevertheless laboratory fecundity cannot be assumed in the field, where not only food availability (Johannson 1958b), but in- cidence of feeding, weather, predation and the like would adversely affect fecundity. Density factors may be involved (Watt 1962) and this was apparent in P. bilobatus (see in Species Discussions). It was repeatedly observed that the total number of eggs laid per fe- male in mass culture was much less than the fecundity found in individual rearings. Nevertheless, if these uncertainties are assumed to cancel out, insofar as species differences are concerned, there were distinct differences in the reproductive rates among the species. In general the rapid developing species of temporary habitats had higher oviposition rates than slow developing species of permanent hab- itats. However the relation also seems to partly involve size (as expressed by length), the larger species laying a larger number of eggs. Among the larger myodochines (over 4 mm. long) the mean oviposition rate in the laboratory was 168 to 273 eggs, while the smaller species (under 4 mm.) averaged 70 to 130 eggs, and the rate per day was 6-7 eggs as compared with 3-4 eggs a day in the smaller species. However the smaller species are usually adapted to habitats with low productivity. Similarly the three gonianotines Trapezonotus, Malezonotus, and Delochilocoris averaged 3-4 eggs a day, and at least the first two are characteristic of sparse habitats. But Drymus unus and Pachyb rachitis basalis which are close to the above in size were found to lay a mean of 150 and 170 eggs each and each inhabit seed-rich habitats, the former in permanent woodland habitats, the other in temporary habitats. Several other factors influence egg productivity. When the virgin females are sexually isolated in most species their egg pro- ductivity is greatly reduced. The sole exception is Heraeus plebe- 70 VOLUME XLIII jus which would lay eggs freely whether mated or not. Among the species two conditions may be distinguished. In the first no eggs whatever were laid. These species include Pachybrachius basalis, P. albocinctus, Cnemodus mavortius, Sphaerobius insignis, Peri- genes constrictus, Plinthisus americanus, Xestocoris miens, Antil- locoris minutus, A. pilosulus, Drymus unus, Delochilocoris umbrosus , Malezonotus fuscosus, Embletkis and perhaps the two species of Stygnocoris. Among the remaining species from none to a few eggs were laid by sexually isolated females. These species include Myodocha serripes, Ligyrocoris diffusus, L. depictus, and L. caricis, Zeridoneus costalis, Sisamnes clavigera, Carpilis consimilis, Cryp- hula trimaculata, Ozophora picturata, Peritrechus f rat emus, Tra- pezonotus arenarius, Scolopostethus thomsoni, S. cliffidens, Eremo- coris ferus, and Megalonotus chiragrus. Logically, such oviposition control would be of considerable adaptive significance, for it would conserve resources and prevent the metabolic waste of unfertilized eggs. In correlation the virgin females lived a much longer time than the fertilized ovipositing females. For example in Myodocha serripes a virgin female lived for over a year while the normal adult life span is about 5 weeks. (See species discussions for further examples.) Another important factor is the availability of an appropriate substrate for oviposition. In many species such as in Ligyrocoris diffusus,. Pseudocnemodus canadensis, and Emblethis vicarius the egg production is sharply decreased if it is not provided with a methyl cellulose, cotton or soil substrate for oviposition. Eggs and Oviposition Southwood (1956) summarized the knowledge of the eggs of the terrestrial Heteroptera and discussed the phylogenetic significance of the different egg types. Like Pentatomomorpha eggs in general, the eggs of the Lygaeidae lack a true operculum and have an anter- ior ring of micropylar processes which serves for both sperm passage and air exchange. An egg-burster is always present. This is a small spine surrounded by a chitinized area on the embryonic vertex. The egg burster is inconspicuous in the Lygaeidae and best seen on the shed embryonic cuticle. The Lygaeoidea, with the exception of the Cleradini (sensu Stal) (Sweet, unpublished), never have a true pseudoperculum as in the Pentatomidae, although this condition is approached in Piesma (Southwood 1956, Putshkova 1956). Corby (1947) and Putshkova (1956) have described a thin polygonal area on the anterior end of the egg within the ring of micropylar processes. Each side of the polygon corresponds to a 71 ENTOMOLOGICA AMERICANA micropylar process and tlie cleavage of the eclosion rupture begins at the grooves which extend from each corner of the polygon. A micropylar process may be absent but the side of the polygon re- mains. Putshkova (1956) found that the cleavage pattern cor- responds to the various egg types. Eggs with rounded anterior ends usually split laterally along one or two sides and the cleavage curves ventrally. In eggs with flattened anterior ends, the cleav- ages may radiate from all the grooves to the central polygon. The exact type of cleavage or bursting is species specific (Putshkova 1956) and although there may be some variation, especially in lygaeoid type eggs, it is however not entirely correct to state that the chorion splits irregularly at ecdysis (Andre 1934, Southwood 1956). In most species, the micropylar processes show a considerable variation in number except in most of those species with four micro- pylar processes. The number of processes varies from 3 to 12 in the known Rhyparochrominae and are always located close together in a ring. It would appear from the variation of 5 to 9 in some species that the number of sides in the central polygon also varies. As in the eggs of Myrmus miriformis Fallen (Woodward 1952), egg development was in all cases apparently independent of free water absorption. However, in a dry atmosphere, the eggs, espe- cially those eggs which diapause, may desiccate, and at least a humid atmosphere was frequently necessary to prevent such des- sication. The eggs of different species varied in their resistance to drying. This difference was frequently expressed in the moistness of the preferred oviposition substrate. In the normal ground level biotope, the favorable microclimatic condition of humid still air and reduced evaporation rate prevents undue dessication (South- wood 1956). Similar conditions exist in a closed petri dish as the humidity is high but free moisture is lacking. No water uptake was observed in semidesiccated eggs placed on a moist substrate. The process of development and hatching is much as described by Southwood (1956). Lygaeid egg chorions are transparent and particularly convenient for observing their development. South- wood (1956) and Corby (1947) noted that the embryonic cuticle actually represents the first true instar. However, in this paper, the first instar refers to the first actively moving or normal instar. The embryonic cuticle in all cases was shed after complete emergence from the egg. The posterior apex of the embryonic cuticle sticks usually to the edge of the cleavage in the egg shell, or to the substrate a short distance away from the egg. Putshkova (1956) has studied in detail the eggs of Palearctic Lygaeidae and constructed a key based upon egg types and oviposi- *72 VOLUME XLIII tion. In a later contribution, I hope to present a similar and complementary study of the Nearctic lygaeid eggs. I wish to dis- cuss here only those aspects which may pertain to the ecology of the various species. Putshkova (1956) found lygaeid eggs to be quite diverse in form and she distinguished six types of eggs, four of which were also applicable to other heteropteran families. These types were named : lygaeoid, aphanoid, oxycarenoid, piesmoid, berytoicl, and macroparoid eggs. Kullenberg (1944) could find no systematic meaning in the egg types of the Miridae and considered the differ- ences present to represent specific adaptations to the site and man- ner of oviposition. While some similar adaptability is shown among the Lygaeidae, Putshkova ’s egg types appear to have some sys- tematic significance but not at the high level of significance sug- gested by Reuter (1910) for egg types. All the aphanoid type eggs were found only among the Rhyparochrominae, and in none of the other lygaeid subfamilies was this particular type of egg found. On the other hand, a few rhyparochromine species have eggs whose forms place them in certain other categories (Putshkova 1956). Among the New England Rhyparochrominae only two of Putshkova ’s egg types were distinguished, the aphanoid and pies- moid. The aphanoid eggs are elongate-cylindrical and one side, the ventral, is flattened or concave, the opposite side, convex. Most are roughly “cucumber-shaped”. The piesmoid type eggs are elongate with the anterior end flattened and the posterior end somewhat pointed. Putshkova, however, states that rhyparochro- mine eggs really only approach the piesmoid type. This type is illustrated by Southwood (1956). Actually these two types were not precisely exclusive, as some aphanoid eggs approach the piesmoid type. Moreover, the aphanoid eggs differ considerably and can be further subdivided. Both types are basically similar with the similar micropylar processes grouped relatively close together on top of the egg and have the venter concave. Putshkova classi- fied several rhyparochromines as having lygaeoid type eggs, but at least in the case of Sphragisticus, the eggs I examined had a defi- nite ventral curvature which is thicker in the middle and could not be clearly distinguished from the aphanoid type egg. The lygaeoid type, I think, should be limited to the oval almost football shaped egg with a wide circle of many micropylar processes. This dis- tinctive egg type is characteristic of the subfamily Lygaeinae sensu stricto. Putshkova places the eggs of Gastrodes among the beryti- noid type eggs. This type has longitudinal ribs along the side, but the anterior end is not flattened. 73 ENTOMOLOGICA AMERICANA We may distinguish the following types of eggs among the New England Rhyparochrominae. Piesmoid type eggs Group one. — eggs with the anterior end strongly flattened, so that a marginal rim appears to be present. These eggs have a smooth chorion which is fluted : Plinthisus americanus, Carpilis consimilis, Ptochiomera nodosa, Sisamnes clavigera. The eggs of Ischnocoris and Acompus are also of this type. Group two. — eggs with the anterior end flattened but a distinct rim is not evident : Peritrechus fraternus, Pacltybrachius basalis, Kolenetrus plenus, Ligyrocoris caricis. Putshkova found Peritre- chus nubilis and geniculatus to be of this type but not P. sylvestris. Aphanoid type eggs Group one. — eggs without setules. Eggs of this group vary from being quite short to rather elongate. The shorter ones are thickly cylindrical and show only a slight curvature. They are listed from relatively shorter to longer as follows : Xestocoris nitens, Crypkula trimaculata, Antillocoris minutus, Stygnocoris rusticus and 8. pedestris, Eremocoris ferus, Scolopostethus thomsoni, S. diffidens, S. atlanticus, Sphragisticus nebidosus, Heraeus plebejus, Pachybrachius albocinctus, Myodocha serripes, Cnemodus mavor- tius. Group two. — eggs covered with chorion setules or “nap”. This chorionic investiture varies from sparse to thick and dense : Ligyrocoris depictus, L. diffusus, L. sylvestris, Sphaerobius insignis Uhler, Zeridoneus costalis, Perigenes constrictus, Pseudocnemodus canadensis, Ozophora picturata, Megalonotus chiragrus (this genus has remarkable “tack-like” processes), Trapezonotus arenarius, Delochilocoris umbrosus, Malezonotus fuscosus, Emblethis vicarius, Dry mus units, and D. crassus. Systematic Relationships There is a diversity of egg types among the Myodochini, the largest tribe. The only similarity is the relatively elongated shape of their eggs. In contrast, the Lethaeini, Antillocorini, Stygno- corini, and in general, the Drymini, all have eggs proportionally shorter than the Myodochini eggs. All Gonianotini have setulose eggs. These similarities are in large part related to the oviposition habitats of the species and may reflect common adaptive similarities. This is especially true of the setulose investiture. While our two species of Drymus resemble the Palearctic D. brunneus in having 74 VOLUME XLIII setulose eggs, D. sylvestris has perfectly smooth eggs. Moreover, D. brunneus belongs to the same snbgenus as sylvestris ( Sylvadry - mas) while our species represent Drymus sensu strict o. While all our species of Scolopostethus have smooth chorions like the Palearc- tic 8. pilosulus and 8. lethierryi Jak., S. affinis Schill. and 8. decora- tus Hahn have setulose eggs. Eremocoris podagricus also has setu- lose eggs while those of our E. ferns are smooth. A similar situation is found in the eggs of Peritrechus and Rkyparochromus (= Raglius) (Putshkova 1956). The piesmoid type reappears independently in the Myodochini, Rhyparochromini, Stygnocorini, and Drymini. Oviposition Sites As described by Putshkov (1956) most of the Lygaeidae and all the Rhyparochrominae lay their eggs singly although a small pile of eggs is sometimes formed in some species. Only Heterogaster always lays a mass of eggs which are cemented together with a copious secretion (Putshkov 1956). When selecting a place for oviposition, the females investigate a site with the antennae, sometimes the labium, and subsequently carefully and repeatedly probe it with the ovipositor. These insects possess a remarkable and delicate control over the movements of the ovipositor and can work it into various kinds of substrates and crevices. In soft material the apex of the abdomen may be sub- merged during oviposition. It is noteworthy that the apex of the ovipositor has a series of erect stiff hairs and the repeated probing and pushing probably represent trials toward stimulating these sensory hairs equally. The actual process of oviposition lasts at least several minutes. The type of substrate preferred by the different species cor- responds in large part to the oviposition behavior and egg type. Southwood (1956) classified the oviposition types in the Heteroptera into four loose categories and criticized Michalk’s (1935a) over- rigid ten categories. All rhyparochromine egg sites fall into South- wood’s category of semi-exposed egg sites “e.g., in the soil, under rocks, amongst fallen leaves, in axils of stems. ...” The soft ly- gaeid ovipositor precludes oviposition into plant material (Putsh- kova 1956), but allows a definite placement of eggs into crevices, surface pilosities, soil, and even into rather tight substrates. Butler (1923) was then incorrect in referring to the lygaeid ovipositor as being “saw-like” with the single known exception of Ischnodemus sabuleti Fallen. According to Putshkova some species do not attach their eggs to the substratum but lay them free or at random. The chorion is usually smooth and the eggs fall readily into fissures between bits of ENTOMOLOGICA AMERICANA litter and the ground. On the other hand, eggs which are covered with different kinds of nap, or chorionic investiture are frequently laid on pubescent, hairy or similar substrates. However, as Putsh- kova notes, in some species the smooth eggs are laid in different fashions. The elongate piesmoid eggs with pointed apices and flattened anterior ends, were usually laid into firm substrates, either into wet or dry litter. These eggs were frequently forced into the tight cotton stoppers of the water vials (Pachybrachius basalis, Carpilis consimilis, Peritreckus f rat emus, Ligyrocoris caricis). In the other species the eggs were laid into the parenchyma of plant stems or narrow grass culms, or otherwise into cotton stoppers or loose litter. The elongate egg form would appear to facilitate oviposition in firm substrates. However, the thicker aphanoid eggs of Sphra- gisticus nebulosus are similarly laid. The slighter egg curvature of Sphragisticus may correspond to its habit of laying eggs into the parenchyma of fallen herb stems (Putshkova 1956). Plin- thisus oviposits into the tight crevices of hemlock and spruce litter, frequently placing the eggs into hollow needles. The eggs of Sisamnes also stick by a cement layer to the substrate in which they are laid. The other species which lay smooth, but aphanoid eggs fall into three groups. The first are those which oviposit directly into loose litter and lay eggs which are free and sift readily into crevices. These include the short, thick eggs of Cryphula, Xestocoris, Antil- locoris, Eremocoris, and Scolopostethus. The latter three are woodland species which preferably lay their eggs in moist litter, or, in the laboratory on loose cotton. The eggs of Scolopostethus, as shown by Putshkov have leathery egg shells which give readily on slight pressure, in contrast to brittle chorion typical of the other species. The second group includes those which lay eggs with a consider- able cement layer, and the eggs are stuck to the litter or soil of the oviposition site and to each other. These include the eggs of Stygnocoris which also are diapause eggs. Since all heteropteran eggs are apparently laid with a cement layer from the accessory glands (Southwood 1956), it appears that the cement coating must differ considerably among the different species. The eggs of these latter two groups, if no substrate is available, are laid “at random”. The species of the third group lay their eggs into the ground or into tight crevices and the eggs are relatively elongate. These are all myodochines and include Pacliybrachius albocinctus, Myo- docha serripes, Heraeus plebejus, and Cnemodus mavortius. The first three species prefer a fairly moist substrate for oviposition. 76 VOLUME XLIII All the eggs which have ‘ ‘ nap ’ ’ are laid and affixed upon definite substrata. These eggs are all aphanoid in contrast to the piesmoid egg per se (of Piesma) which also have a nap-like surface and are laid upon leaf epidermis. As mentioned, the rhyparochromine piesmoid eggs are laid into tight crevices or into soft parenchyma. All the setulose surfaced eggs stick tightly to the substrate, but the sticking appears to be accomplished more by the cement layer, and the setulose surface apparently prevents ready dislodging, especially on a fibrous substrate as on plant fuzz or pilosity. The setulose surface probably abets sticking to a rough surface as to sand grains. While there is a broad behavioral overlap there are different oviposition preferences among the species. These oviposi- tion sites may be classified into three general categories: (1) on plant fuzz or crevices, and (2) in loose fine litter on ground, and (3) under small surface objects. Accepting a considerable plas- ticity within a species, species may be classified by their preferred oviposition sites. Because of this plasticity the preferential of a few of the species might be misinterpreted. The species which prefer to lay their eggs in crevices or plant fuzz may also lay their eggs shallowly in the soil but rarely lay their eggs deeply into the ground. Frequently the eggs of this group are laid near or on fallen seeds or seed capsules as Putshkova notes. These include Malezonotus fuscosus, Delochilocoris umbrosus, Psen- docnemodus canadensis, and frequently the summer generation eggs of Ligyrocoris diffusus, Zeridoneus costalis, and Perigenes con- strictus. Ozophora picturata, Drymus unus, and P>. crassus lay their eggs in crevices in leaf mold but could be placed as readily in the second type. The eggs of Drymus species like Stygnocoris have a heavy cement coat and are frequently found stuck together in a group of two or three eggs. The second type of site, ground deposition, actually would in- clude all the species, since both of the other types frequently over- lap this type. However, certain species especially manifest this habit and lay their eggs quite deeply in loose soil. In the laboratory these species especially prefer to lay their eggs into deep mounds of dry cotton or methyl cellulose. In ovipositing, species of this group commonly oviposit in a vertical position with spread legs, and also often work the apex of the abdomen into loose soil as well. These species include Ligyrocoris diffusus, L. depict us, and L. sylvestris, Zeridoneus costalis, Sphaerobius insignis, Perigenes con- st-rictus, and Emblethis vicarius. The third type are those which lay their eggs preferably under small objects. These species show little inclination to lay their 77 ENTOMOLOGICA AMERICANA eggs into cotton or methyl cellulose, and never deeply, and more frequently will lay their eggs in narrow crevices or in plant fuzz. These species are readily recognized for they predominantly lay their eggs directly backward with the body horizontal, and show very little vertical displacement. These species include : Trapezo- notus arenarius, Megalonotus chiragrus and frequently also Malezo- notus and Delochilocoris. In no case was a hole observed dug into the sand for oviposition as Weber (1930) illustrates for Rhyparochromus pini (L.). Nor were the fore legs ever observed being used to help dig such a hole as he suggested. Southwood and Leston (1959) also note that this species lays its eggs deeply. While Drymus spp. and L. sylvestris lay their eggs in mesic or moist sites, all the other species with setulose eggs are typical in- habitants of drier locations where the exact egg placement and lodgement may be important in ensuring proper microclimatic con- ditions for development. This general relationship is also true of the Palearctic species as was shown by a comparison of Putshkova’s egg descriptions with the ecological data of Penth (1954) and Southwood and Leston (1959) . The eggs of this type, especially of Emblethis, Sphaerobius, and Malezonotus, when laid in the soil, become densely covered with sand grains and litter which makes the eggs difficult to locate in the cultures. It would be most interesting to compare the oviposition habits with egg parasitism which undoubtedly occurs. It would seem that the thick layer of attached sand grains and deep oviposi- tion of individually laid eggs should lower the parasitism rate. It would appear then, from the different types of oviposition sites and also from the large percentage which diapause (see Season- al Cycles), that such adaptations should result in different reactions of the species to weather vicissitudes and perhaps, to parasite and predation pressures. Competition Now that a general review of the biology of the New England Rhyparochrominae has been completed, the ecological relationships among the species may be considered. It was mentioned earlier that frequently there were characteristic assemblages of rhyparo- chromine species. Since each of these species is apparently oligo- phagous on seeds it would appear that these species assemblages are living together in the same functional niche in the same ground level biotope. The standard question which then presents itself is are these species really in the same niche and in direct competi- tion? It should be kept in mind that the following discussion is 78 VOLUME XLIII largely exploratory as a great deal of quantitative data and careful study is required to approach an accurate understanding of com- petition. The question of competition is of course an important one, one which forms a major foundation of evolutionary theory (Darwin 1859, Crombie 1947), of modern systematic thinking on speciation (Mayr 1942, Lack 1947, Brown and Wilson 1956), as well as a fundamental ecological concept (Gause 1934, Hutchison and Deevy 1949, Allee et al. 1949, Hardin 1960). Recently the concept has been attacked by Andrewartha and Birch (1954) and some phases are presently in a state of flux (Slobodkin 1961, Klopfer 1962) although the basic principles and implications of competition as laid out by Darwin (1859) are entirely valid (Kendeigh 1961). Much of the controversy has revolved around the basic unit of thought upon competition, the niche. Since these considerations are basic to any interpretation of the biology and ecology of a taxon (Lack 1944), it is apropos to the present discussion to review the concepts involved. The Niche When originally proposed independently by Grinnell (1917, 1924, 1928) and Elton (1927) the concept of the niche was given similar but distinctly different meanings (Clarke 1954). Grinnell conceived of it as a habitat niche, a discrete spatial and distribu- tional unit occupied by a species, and to which the species was held by structural and instinctive factors. Elton defined it more in terms of an animal’s position in a trophic hierarchy, or its functional place in a community. The common European use of the word biotope corresponds very closely to the spatial aspect of Grinnell ’s concept, and forms the unit equivalent of a species in a habitat analysis (Hesse, Allee and Schmidt 1951). Elton’s definition along with the instinctive and structural aspects forms the ecological niche or the present functional role concept of the niche. From this original structuring of space and function, the niche has been interpreted in terms of competition (Gause 1934) which led to the formulation of Gause ’s Law which stated that two species with the same ecology cannot persist in the same region, i.e., no two species can occupy the same niche. Although applicable to many species this concept has presented some difficulties when applied to several natural situations (Ross 1957, 1962) perhaps because of the then-prevailing concept of the niche. This has led to an operational definition in which an “ecological niche or space is that space which no two species can continue to occupy for an indefinitely long period of time” (Slobodkin 1961). Slobodkin fur- 79 ENTOMOLOGICA AMERICANA ther stated “if two species persist in a particular region it can be taken as axiomatic that some ecological distinction must exist be- tween them, and that their ecological niches, in the restricted sense, do not coincide.” Thus Hardin (1960) could erect this concept into an exclusion principle based on competition. This concept, and especially that of competition, has been sharply attacked (Andrewartha and Birch 1954, Birch 1957 and Andrewartha 1961) as not applicable to ordinary “rare” species or in large areas which regularly suffer great ecological changes. Moreover the size of the niche has become disconcertingly small in many situations. Hutchison (1959) reviewed the size ratio of trophic characters in ecologically overlapping species and obtained the ratio of 1.2 to 1.4 which he explained as an example of character displacement in feeding habits. However Klopfer and McArthur (1961) found that in some tropical mixed flocks of birds, the ratio was actually unity between some species. This they explained as resulting from extremely stereotyped behavior patterns. Park (1954) showed that only a small difference in environment adapta- tion to temperature and humidity separated two species of tene- brionids. Andrewartha (1961) called attention to the remarkably long time of 6 months to several years for the competition between the beetles to come to completion, and considered the situation im- possible in nature with insects of one generation a year in an extremely variable climate. Ross (1957) cited the example of six Erythroneura leaf hoppers on the same host plant and explained this situation (1962) on the basis of different microclimatic optima within the context of seasonally and locally variable weather con- ditions. These conditions produced an ecological oscillation among the species as the first one and then another is favored. In this context, then, the exclusive niche may be the small difference in climatic optima between these species (Slobodkin 1961). This appears far from the trophic role of Elton’s definition of the niche. Indeed the competitive niche concept is applicable with difficulty to preclimax species especially those which invade a habitat only briefly and are termed by Hutchinson (1951, 1953) as “fugitive” species. Such niches if so defined would have a large open or escape aspect and competition can scarcely be understood in such a species (Slobodkin 1961). Slobodkin admits these difficulties and states, “It is true that some field situations are difficult to explain in such a way as to conform to the Gause axiom and that occasionally the explanations are quite frankly speculative. The abandonment of the Gause axiom however, is equivalent to aban- doning the concept of competition, and competition is the only reasonable mechanism developed to explain the generally homeosta- 80 VOLUME XLIII tic properties of the natural world.” It is not in the scope of the present paper to evaluate this con- troversy except to cite Watt’s (1962) criticism of the tendency of ecologists to believe in a priori models without empirical testing. In such controversies, the reality usually lies between the semantic poles and is fairly complex (Allee et al., pg. 729). For this reason the interpretation of Ross (1962) may present the most balanced judgement of niches and competition. Competitive Relations In species of the same community Ross (1962) makes the stan- dard ecological distinctions among exploitation, mutualism and coexistence, and among different trophic levels. However he dis- tinguishes between direct and indirect competition for the same commodity. In direct competition the organisms feed side by side, simultaneously and in the same fashion on the same food. If some significant difference exists in the site, time or method of feeding, Ross considers this situation indirect competition, for these organ- isms, contrary to the common conception, do not avoid competition for potentially each species reduces the nutrition available to other species. This is a logical extension of the same reasoning which led Slobodkin (1961) to disagree with Hutchinson’s concept (1957) that a nocturnal organism occupies an entirely different niche than its diurnal counterpart. Slobodkin reasoned that these species were actually in competition, i.e., in the same niche since each utilizes the same fixed energy source of which any depletion results in a diminished supply available to other competitors. Competition then is indirect for the present purposes under the following conditions. (1) Host seed preferences differ even though there may be a complete overlap, (2) seasonal cycles distribute the feeding stages of different species in different periods of the year. For example, the orthopterans Arphia sulphur ea and A. xanthop- tera exist in similar habitats but the former overwinters as a nymph, and the latter as an egg. In different lygaeid species the time of oviposition and whether overwintering occurs as an egg or an adult is important. (3) Size differentials are usually cited, and may involve, among rhyparochromines, an ability to move and defend seeds or the ability to penetrate and feed on the seeds with their stylets. Direct competition occurs when either of two conditions or maxims are met (Ross 1962) : “ (1) If populations over the entire community range are at levels producing interspecific competition, coexistence will be possible only for species best adapted to some recurring variant of the ecological pattern of the community. (2) 81 ENTOMOLOGICA AMERICANA If populations are habitually or locally below levels producing interspecific competition, any number of potentially but not actually competing species may coexist.” Conceivably both of the above conditions can operate on a given population. The first situation has already been discussed in terms of unstable niches. The second situation is an important one upon which biological control is largely predicated (Glen 1954) in which a combination of inanimate environment resistance together with predation-parasite pressure reduces insect populations well below the carrying capacity of the nutrition available. This situation is characteristic of normal complex communities (Smith 1935, Glen 1954, Andrewartha and Birch 1954). It should be mentioned that along with standard predator-parasite-disease factors, host resist- ance in plants (Dethier 1947, 1954, Fraenkel 1959) is surely an important factor involved in the controlling of the insect popu- lations. In most situations the survival of a species requires adaptations to an inclement and variable climate which involves the seasonal cycle, diapause, cold hardiness, migration, and egg sites. There need also be adaptations to predators and parasites (Cott 1940, Klopfer 1962) which would include protective coloration and be- havior, concealment habits, density patterns. Obviously such adaptations also influence the competitive relationship among differ- ent species. In this context, among the Rhyparochrominae the procryptic, warning ( ? ) and ant mimicry coloration, repellent odors, and protective behavior takes on relevance, as do the seasonal cycle and brachyptery phenonema. Since all of the above attributes strongly affect the competitive survival of a species, they should (Klopfer 1962) be considered as a component of the niche of a species and utilizable in analyzing an apparent niche that contains several species. Ross (1962) emphasizes that survival adaptions per se may be difficult to dis- tinguish from the adaptations supposedly arising from niche segre- gation (Lack 1947). Brower L. (1958) suggests that many appar- ent host selections or ranges may instead represent behaviorial adaptations to predation pressure upon procryptic insects. Such adaptations would render predator learning more difficult in the mosaic distribution of the plant forming the background. Reason- ing along similar lines Slobodkin (1961) suggests that conceivably several species could inhabitat the same (?) apparent niche if mutual survival is enhanced. It follows then that such a symbiotic- like arrangement could involve several species with different pro- cryptic patterns or behavior, especially where the reproductive rate barely exceeds predator pressure so that food competition is 82 VOLUME XLIII low. This is analogous to the explanation given to polymorphic color patterns, especially in mimicry, in which predator learning is rendered more complex (Sheppard 1958). The problem posed by the many rare species as emphasized by Darwin (1859) and Andrewartha (1961) are hardly treated by population workers and indeed are almost jocularly dismissed by Slobodkin (1961) as due to “ exogenous” factors without explaining the limitations or acknowledging that many of these possible factors are not based on competition. Kendeigh (1961) suggested that the competitive niche concept can be extended to such organisms and others of densities lower than carrying capacity by postulating occasional high abundance levels when competition will then estab- lish the niche restriction of the species along the lines suggested by Lack (1947). Obviously care must be taken in assessing abund- ance in relation to the niche size. No attempt is here made to com- pare the abundance levels of the different seed feeding rhyparo- chromines in different communities with the species abundance distribution formulae developed by McArthur (1957). While the rhyparochromines qualify as a uniform trophic level, it is better to take the entire seed feeding trophic level into consideration, if competition of this sort is to be considered. Significance Thus one can readily see the complexity involved in assessing competition and niche definition, especially in sympatric competi- tors. As Klopfer (1962) said, it has become a routine exercise for ecologists to demonstrate feeding differences in sympatric species, or other differences that place the species in different niches. For the insects studied here the question which appears is, what differences? For all the rhyparochromine studied here are bio- logically distinct from each other, even those of the same genus. As Slobodkin admitted, often any biological difference ascertained are assumed a priori to account for the niche differences in species of different appearance. Such thinking is tautological since it has been repeatedly shown that morphological evolution itself is adap- tive (Lack 1944, 1947). Moreover as Ross (1962) and Andre- wartha (1961) emphasized, a species, when evolving, is bound to become biologically differentiated in some way. Thus most of the demonstrations of niche differences among species in the same habitat have been purely hypothetical since “what is not yet clear is the degree of ecological difference required to permit coexistence, and we are not even sure how this difference should be measured (Slobodkin 1961).” It then appears from this uncertainty and the various relation- 83 ENTOMOLOGICA AMERICANA ships possible (Ross 1962) that it would be difficult to accurately evaluate competitive relationships even in very well studied groups. For the present discussion then, it is best to point out the differences among the coexisting species of rhyparochromines. It must be emphasized that not enough is known about the causes of natural mortality, etc., to do more. It is assumed that the ecological segre- gation already observed evolved from former competitive situations (Lack 1947). Genera The best evidence for possible ecological displacement is in the distribution of related species in a genus. However generic size in New England rhyparochromines is relatively small and most genera are represented by only a single species. To further evaluate this aspect, the same sort of study needs to be done in a warm species- rich southern climatic area. Only six genera have more than one species present in New England. Ligyrocoris with four, Pachybrachius with two, Peri- trechus with two, Drymus with two, Scolopostethus with three (or four) and Stygnocoris with two. Only in the case of Stygnocoris were species of the same genus found together in the same habitat, and Stygnocoris species were much more frequently found separated than together. The species differ as follows : In Ligyrocoris, L. diffusus is typical of open ruder al habitats ; L. depict us of dry sparse bald habitats ; L. sylvestris of northern cool woodland margins ; L. caricis of a Carex stricta aquatic transition habitat. In Pachybrach- ius, while P. albocinctus inhabits aquatic transition, P. basalis is found under rank, often ruderal vegetation. The three species of Scolopostethus are found as follows : S. diffidens in climax or sub- climax forest of white birch-hemlock ; S. atlanticus is geographically separate and found in a mature shrub and red maple community, and S. thomsoni varies from aquatic transition to moist roadsides and meadows with scattered trees. In Drymus, D. unus is found in aster-rich subclimax deciduous woodlands, D. crassus in mature cli- max woodlands. In Peritrechus, P. f rat emus is usually found in beach wash ; P. paludemaris in salt marshes. Stygnocoris pedestris is often found on woodland margins and grass dominated habitats while S. rusticus is especially abundant under tall forb habitats. Species Assemblages Because of the importance of the steady state condition in Gausian competition, the species assemblages fall into two groups: inhabitants of short-lived habitats and of long-lived habitats. AVhile such short-lived (Tj) habitats should be of the type char- 84 VOLUME XLIII acteristic for fugitive species, they also yield the highest abundance of rhyparochromines, presumably because of the abundance of fallen seeds and perhaps a lack of population density controls. Competitive conditions presumably exist briefly under these con- ditions. The pattern then is a rapid succession of species whose habitats broadly overlap and should partly compete at times. These species generally show rather catholic feeding habitats and have higher reproductive potentials under similar conditions than most permanent habitat species. The species of this group can hardly be conceived of as species assemblages as the associations are sporadic and brief. Each of the species does select a fairly definite habitat and the species may be organized into a succession scheme (see Ecology). It is not clear whether these species succeed by competition or habitat choice. An important and interesting aspect is that the two introduced megalonotines Megalonotus and ( ?) Sphragisticus apparently com- pete rather poorly with the native rhyparochromine fauna, and are found in rather restricted man-influenced habitats in New England. Sphragisticus is nearly always found around gardens and in ruderal habitats, but even then much less abundantly than Ligyrocoris and Pachybrachius at the same sites. Megalonotus was nearly always restricted to patches of the European plant, Centaurea. Trape- zonotus arenarius, if introduced, does not show the wide ecological range of the Palearctic species (Southwood and Leston 1959, Penth 1952). In the seed-rich litter habitat, Peritrechus fraternus was domi- nant over the other rhyparochromines which invaded briefly- Pachybrachius basalis, Perigenes, Ileraeus. One species, Delocki- locoris umbrosus, has the most sporadic ecological distribution of all and occurred briefly in a large variety of open habitats and only once was found in any abundance in a peculiar thin wash habitat. Of all the rhyparochromines studied it appears' to best fit the fugi- tive species concept of Hutchinson (1953) as it simply did not appear to compete well with the other rhyparochromines for long. In a sense all of the species of temporary habitats may be considered fugitive species (Slobodkin 1961). The second broad group comprises the species of permanent (P) and semipermanent (T2) habitats (see Table 4). In this group the long persistence of the species allows discrimination of definite species assemblages which are present year after year and indicate a steady state relationship, especially since the majority of the individuals of such populations are flightless. While a particular assemblage will remain present, the exact species composition of the assemblage varies, especially on a north-south pattern, and 85 ENTOMOLOGICA AMERICANA component species may vary in relative abundance from site to site. Northern Woodland Assemblages Eremocoris ferus, Scolopostethus diffidens, and characteristic- ally, Plinthisus americanus compose this group. It is especially characteristic of mixed deciduous conifer woodlands. All species feed on Tsuga and birch seeds. Plinthisus was most abundant rela- tive to the other species in pure Tsuga or Picea woodlands. On the trophic level the species differ in that Plinthisus has one yearly generation and lays diapause eggs in autumn : Scolopostethus is also univoltine but diapauses as an adult and has a single early summer feeding period. Eremocoris feeds and oviposits through- out the summer. Moreover Eremocoris is about twice the size (5.2- 6.0 mm.) of Scoloposthethus (3-3.5 mm.). The life cycles differ as E. ferus hibernates, the others diapause over the winter. Plinthisus has procryptic coloration, the others are flash type ant mimics. All hide under litter but the drymines run rapidly when disturbed while Plinthisus moves slowly. Eremocoris is macropterous with a relatively wide distribution. The others are brachypterous and restricted in distribution. Drymus unus was found only in deciduous margins, D. crassus in rich climax deciduous forests. Both species of Drymus have one generation a year, overwinter as an egg, and the nymphs are bright red (warning coloration?) . Middle Woodland Assemblages This is a similar group except that Plinthisus drops out and Antillocoris minutus appears. This species is procryptic, slow moving, and minute (1.8-2. 2 mm.) but with a life cycle similar to Scolopostethus. Drymus unus is also a regular member and differs from the other species in that like Plinthisus it has a fall feeding and oviposition_ period with diapause eggs being produced. It moreover prefers aster seeds to birch seeds. Southern Woodland Assemblages This group is characteristic of a V accinium-V iburnum-Acer rubrum community. Eremocoris ferus and Antillocoris minutus are again members. Scolopostethus diffidens is replaced by S. atlan- ticus which has a similar life history. Both species feed on V ac- tinium and Viburnum seeds. Z>. unus is abundant, and Ozophora picturata is present. Ozophora has a late summer reproductive period which coincides with the second generation of Eremocoris ferus. At this time Scolopostethus and Drymus feed little, al- though Drymus has an autumn feeding period. Ozophora is pro- cryptic and rapid moving both as adults and nymphs in contrast 86 VOLUME XLIII to the other species which are self-concealing forms. Moreover Ozophora is larger (6. 0-6. 5 mm.) and is more characteristic of bnt less abundant in dry oak-hickory forests where the other species do not occur. Aquatic Transition Assemblage This group which includes Ligyrocoris caricis, Pachybrachius albocinetus and Scolopostethus thomsoni is characteristic of the Carex stricta community along the shore of Pink Ravine at Storrs, Connecticut. All three species feed on seeds of C. stricta, a clump (stool) forming sedge with cover at its base. On the trophic level, P. albocinetus is not represented by nymphs until after July 10 at which time L. caricis is already midway through its oviposition period and producing diapause eggs. P. albocinetus moreover de- velops here largely on the seeds of the tall bullrush Scirpus. Both myodochines climb the plants frequently to feed, but S. thomsoni which has a bivoltine life cycle which begins a month earlier than Pachybrachius, was not observed to do this. Comparing the cli- matic adaptations, L. caricis diapauses as an egg through late sum- mer and winter, and is not dependent upon equitable late summer conditions as is P. albocinetus. The late life cycle of P. albocinetus may reflect its southern distribution as is the case in aquatic Odonata of tropical derivation which become active only during the warmest part of the season (Kennedy 1927). On the protective level, P. albocinetus is macropterous and flies readily, the others are brachyp- terous and restricted to the location. Both Ligyrocoris and Pachy- brachius have ant mimicking nymphs but the ant resemblance is more pronounced in P. albocinetus. In Scolopostethus thomsoni the color pattern appears disruptive rather than ant-mimicking. In this same community feeding on the same seeds are several other non-rhyparochromine lygaeids, Cymus discors Horvath, Cymus luridus Stdl, and occasionally, Cymus robustus Barber, and Oedancala dorsalis (Say). These species are competitive in the sense that they reduce the available seed supply. However, they are rarely found on the ground where the rhyparochromines are concentrated, and may then be con- sidered as being in a separate ecological niche. Sparse Old Field or Bald Assemblage This assemblage includes Xestocoris nitens, Carpilis consimilis, Pseudocnemodus canadensis, Ligyrocoris depictus, and Trapezono- tus arenarius and is characteristic of dry sparse bald habitats. At any given site one of the species may be absent or infrequent. Xestocoris is the most abundant and characteristic species. On the trophic level Xestocoris is predominantly a grass seed 87 ENTOMOLOGICA AMERICANA feeder. Carpilis will also feed on Veronica seeds, and Trapezonotus, L. depict us and Pseudocnemodus feed on composite and Vaccinium seeds as well. The myodochine species all overwinter as eggs, but L. depictus emerges early and grows rapidly, becoming adnlt in early to late June. Pseudocnemodus is intermediate becoming adult around June 21. Carpilis however develops slowly and does not become adult until mid- J uly. These species begin ovipositing shortly after becoming adult. Some individuals of Pseudocnemodus, however, are bivoltine, rather than univoltine. The other two species overwinter as adults. Trapezonotus mat- urates early and produces diapausing adults by mid- July. At this time Xestocoris is ovipositing and nymphs are still only up to the third instar. The slower cycle of Xestocoris synchronizes its feed- ing period with the ripening of the seeds of Festuca spp. Thus the feeding periods are distributed differently among the species. On the climatic adaptation level, all species are univoltine or nearly so. The myodochines overwinter as eggs, the other species as adults. In the laboratory the eggs of Xestocoris are laid in loose litter while those of Trapezonotus are placed in glumes and crevices. Pseudocnemodus lays its eggs deeply in loose litter, while Carpilis and L. depictus push their eggs into the ground and tight crevices. L. depictus and Pseudocnemodus are found in an open microhabitat on the litter, while the other species keep under the litter. On the protective level, the adults and nymphs of Pseudo - cnemodus and the nymphs of L. depictus are ant mimics. The re- mainder have procry ptic coloration : Xestocoris is a glossy brown ; Trapezonotus, a mottled brown; and Carpilis is a contrasting pale gray and black. L. depictus and Pseudocnemodus run rapidly on the litter when disturbed while the other species seek cover. These species thus present a varied assemblage of forms, colors, and escape reactions to potential predators. Pseudocnemodus and L. depictus, were parasitized by tachinids while the others were not. At Warrensburg, New York, Konenetrus plenus was found with this assemblage. It also feeds on Festuca seeds and has a nymphal development like Pseudocnemodus. It is contrastingly colored to break up the body pattern. It has one yearly generation with diapause over the winter as an egg. In southern New England at Noank, Connecticut Xestocoris is largely replaced by Cryphula, and perhaps Trapezonotus is re- placed by the rather rare Malezonotus when the latter occurs. The latter relationship is speculative, but the Xestocoris-Cryphula ex- change is well marked. These species rarely occur together in the same habitats. The seasonal cycle of Crypula trimaculata is nearly 88 VOLUME XL III identical to that of Xestocoris nitens. Malezonotus is bivoltine rather than nnivoltine. Sphaerobius and Cnemodus are also never found together since the former is northern, the latter southern. These remarkable ant mimics are characteristic of dry Andropogon communities with considerable interspaces. They frequently form unispecific populations. They are each parasitized by different species of tachinids. Thus it is seen in the foregoing analysis of the assemblages that the various species are quite different from one another. The exact meaning of the differences must await studies which can establish what precisely controls the population sizes. In most cases despite the abundance of these insects I doubt that the controlling factor is food as there are simply too many seeds produced and available. Further study of these complexes should yield valuable informa- tion on their population structures. Speciation Lack (1944, 1947) has interpreted the habitat distinctions among sympatric species as steming from former isolation speciation and competitive interactions. The present species are derived from various sources and have migrated into once-glaciated New England. Since the New England fauna includes several zoogeographical elements, the habitat specificities may have derived from competi- tive interaction during the faunal mingling of diverse elements (Ross 1962). It may be illuminating to compare the habitat range of the related species from the center of the distribution of the genera with the New England representatives. The habitat ranges may be restricted in New England as a result of such a competitive mingling. If the species distribution and feeding patterns are interpreted as Kohn and Orians (1962) suggest, the speciation pattern in the Rhyparochrominae is horizontal and habitat-specific as with small mammals and not related to host plants and vertical stratification which results in high speciation. This interpretation would also help explain the lower speciation level of the Lygaeidae as com- pared with the Miridae. Conclusions 1. It has been shown that the species of Rhyparochrominae ex- hibit a well marked ecological distribution. 2. There is a close relation between habitat permanency and the phenomena of brachyptery or flightlessness, and between ma- croptery, temporary habitats, and dispersal records. 89 ENTOMOLOGICA AMERICANA 3. Certain correlations were evident among habitat selection, zoogeographical origin, and systematics. 4. The life cycles are strongly structured, and adapted to the habitat types. Either one or two generations are present and diapause occurs in the egg or the adult. 5. A variety of protective coloration and behavior patterns are noted, and related to the habitat background. Many of the species are ant mimics. 6. Many species especially of temporary habitats are parasitized by the tachinids Catharosia and Petia (= Procatharosia) . The parasites’ overwintering patterns are adapted to the hosts’. 7. While oligophagous on seeds, many species show distinct pref- erence for the seeds of certain species. 8. Many of the species possess a remarkable seed defense behavior and also actively move seeds about to more protected sites, often with aid of the fore femora. 9. The mating behavior of the different species vary from simple to complex with stridulatory devices occurring in some species. 10. Development rates vary among the species and the variation is largely independent of food and temperature and correlates with life cycle whether univoltine or bivoltine and the produc- tivity of the habitat. 11. Reproductive rates can be varied directly with seed supply, and reproduction ceases on foliage diets. Total productivity in the laboratory appears to vary with habitat productivity and size. 12. Ontogenetic changes during nymphal development in color and appendage allometry occurs. 13. Several types of eggs are discerned and the different types in large part correlate with oviposition habits although some systematic relationships seem present. 14. The biology and ecology of the species are considered in rela- tion to competition theory. Species assemblage are shown to be very different among themselves and each species may be considered as being in different niches in the narrow Gausian sense. 15. Species of all habitat assemblages are generically distinct. 90 VOLUME XLIII THE SPECIES ACCOUNTS TRIBE MYODOCHINI Myodocha serripes (Olivier) This species is easily recognized by its strikingly elongated head and is better known than most other rhvparochromines. Many workers have been impressed by its apparent “predacious” appear- ance and in the early literature Myodocha was even placed in the Reduviidae (Spinola 1837, Kirby 1837). Barber (1932) thought the species was at least in part predacious. Blatchley (1926) compared it to the carabid beetle Casnonia Pennsylvania (L.) and surmised that the long slender necks of both species had developed through reaching into crevices. Later workers, as summarized by Sweet (1960), have found it to be phytophagous. I have little positive evidence about the adaptive significance of the elongated head, except that I have frequently observed the insect probing inside the fallen seed capsules of St. Johnswort ( Hypericum spp.) and the whole head may enter into the capsule. However, many rhyparochromines do this, and this answer can only be suggestive. Actually, the significance of the long “neck” may probably be better sought among tropical species of Myodocha which have much longer necks — which inspired Stal (1862) to name one species giraffa. Myodocha serripes represents the most northern extension of Myodocha , a predominantly Central American genus of seven species. M. serripes has the widest known distribution of any of these species but this may be largely a result of more thor- ough collecting and the distribution patterns of the Neotropical species will undoubtedly be extended. In fact, the genus is already known from Ecuador (Campos 1925) and southern Brazil (Slater, in lift.). In southern Florida, M. serripes is replaced by an entirely differ- ent species, the endemic M. annulicornis Blatchley. Unfortunately no records of Myodocha are at hand from northern Florida and Georgia. M. serripes is recorded (Slater, Catalogue) north to Quebec and Minnesota, west to Colorado and Texas, and south to Louisiana and South Carolina. Brimley (1944) found the species only to the west of Raleigh, North Carolina, and not on the low coastal plain. While Moore (1950) collected it in Quebec and Parshley (1917b) recorded it from Maine to New Hampshire, I found it extremely uncommon from the northwestern highlands of Connecticut northward. 91 ENTOMOLOGICA AMERICANA Environment As Uhler (1876, 1878, 1884) early noted, M. serripes over- winters in woodlands and moves into field habitats during the spring. The autumn migration is quite marked and begins in Connecticut in late August (first date, August 25) well before the advent of cold weather. This migration continues throughout September and into October as the nymphs mature in the fields. The movement into woodlands extends very little beyond the forest side of the ecotone between field and forest. Within woodlands of appreciable extent relatively few specimens are found. A definite preference is shown for a light woodland of mixed black birch, red maple, and red oak. The litter beneath such a combination is of medium looseness (4-6 inches deep), and moderately moist in con- trast with the dry loose litter of oak alone or the very tight litter as of conifers and Betula populifolia Marsh. Despite the great abundance in the area studied of pure oak litters, no colonies are found in them, while five separate overwintering colonies were found in the mixed litter type, to one of which Myodocha returned for four successive years. Dowdy (1955) also found Myodocha hibernating in a climax oak hickory forest in Missouri under rocks and logs, often in large colonies. I never collected the insect under rocks and logs but rather in litter along such objects. In the hibernating areas the bugs are found aggregating in groups of from 2-3 to as many as 25 in a square of four inches. They are not found clustered tightly together, but merely near each other. I could distinguish no conditions in the hibernation loci different from surrounding microhabitats. In May when the adults disperse into field habitats they are found in many types of field habitats but in June and later, espe- cially as the ground becomes warmer and drier, the habitat distribu- tion becomes much more limited. This species is found most abundantly in relatively new habitats such as fallow fields, gardens, or embankments which are colonized by a dense, species-rich associa- tion of forbs 1-2.5 feet in height. These forbs include such species as Achillea millefolium, Chrysanthemum leucanthemum, Arthemis sp., Ambrosia art emisii folia, Solidago spp., Bumex spp., Galium spp., and especially Hypericum spp. and Fragaria. These habitats persist at most only 2-3 years, and accordingly, only once have I collected Myodocha two years in succession in the same habitat. The very sparse first stages of plant succession are avoided, as are the more xeric hillsides characterized by morainic gravelly soil. Occasionally a few nymphs are found along an ecotone of southern exposure and it is noteworthy that when this occurs, Hypericum is usually present. The preferred habitats are also characterized by 92 VOLUME XLIII a brown, friable, and moderately (5-7) moist soil which is semi- shaded by the plants, and is covered with a thin litter of broken stems. Litter temperature during the hottest days never exceeds 32°C. Under apparently optimum conditions the popula- tion may reach 25 per square meter. More often, the population is about 5-8 per square meter. It is difficult to follow a population increase as the populations are quite unstable. While the species may certainly be considered common, its abundance is more ac- curately a reflection of the abundance of relatively new forb covered habitats. At no time was Myodocha found off the ground on the plants. No specimens of Myodocha could be collected by sweep- ing at night. There are several literature records of the summer generation. Myodocha sometimes becomes a pest of strawberries in a habitat which is probably similar to natural rank forb habitats (Osborn 1900, Bryson 1939, Neiswander 1944). Walkden and Wilbur (1944) collected the species in alfalfa fields in Kansas. Torre- Bueno (1929a) sifted the species from grass piles in Massachusetts. Wray and Brimley (1943) found specimens of Myodocha in the pitcher plants, Sarracenia flava L. and 8. purpurea L. At Naranja, Florida, I found second and third instars of the closely related Myodocha annulicornis Blatchley in a rank forb habitat at the margin of an avocado grove, a habitat very similar in aspect to those favored by M. serripes. General Biology The dispersion of the species may occur at night as it has been frequently collected at lights (Summers 1891, Tucker 1907, Torre- Bueno 1908, Barber 1923, Froeschner 1944). 1 collected the species in early June at lights which may mean the species disperses as teneral adults at the end of the first generation (see General Dis- cussion), and also in late August which may represent the autumnal hibernation migration of the second generation. As spring nights are quite cool during the vernal movement back into the fields, it may be relevant that I observed a male Myodocha to fly across a field flitting from the top of one herb to another on May 8. Fe- males can be induced to fly throughout the summer, even during oviposition periods. It is noteworthy that all species of the genus Myodocha are entirely macropterous. When presented in the laboratory with a light or dark back- ground Myodocha persistently comes to rest upon the dark one. In the field, the insect blends very well into the dark, stem-strewn surface of its habitat. Its movements further suggest a reliance on protective coloration as it normally moves rather deliberately 93 ENTOMOLOGICA AMERICANA and slowly. Its color pattern is a good example of disruptive coloration as the three white spots, one at the apex of the membrane, the others at the apex of the coriiun, the elongated shining head, and the yellow legs with the dark apices of the long fore femora, together serve to break up the form of the insect and blend it into the background. The second to fifth instars show protective colora- tion of a very different sort. These nymphs are unusual among the Rhyparochrominae in possessing three longitudinal dark bands that extend the length of the abdomen which is otherwise a pale yellow. The bands blend the nymph into the background litter of stems in the summer habitat. When disturbed, Myodocha moves very rapidly, its long legs abetting its movement over the rough litter. No insect parasites emerged from 123 specimens collected. Nabis ferns (L.) and Melanolestes picipes (H.-S.) fed readily on Myodocha adults. In the laboratory, the bug readily feeds on various seeds from the field litter of its habitat. A strong preference is shown for Hypericum sp. and Frag aria seeds, but seeds of Taraxacum, Achil- lea, Erigeon, Aster spp., Potentilla canadensis L., Bumex crispus L., Aquilegia canadensis L., and Paspalum muhlenbergii Nash, are readily fed on. Other grass seeds than Paspalum are, in general, avoided. I never observed this species ever to make any overt move toward aphids, beetle larvae, sow bugs, nor to scavenge on a dying or dead bug as does Pachybrachius basalis. Neiswander (1944), Osborn (1900) found this species to seriously injure everbearing varieties of strawberries by causing the berries to become soft and covered with a mold. In the laboratory it is observed that the feeding is largely restricted to the achenes on the surface of the fruit, and the fruit is pierced only occasionally, and infrequently when water is present. The older nymphs when placed with lettuce grow slowly but in the absence of seeds the adults do not oviposit. It seems very likely that when the strawberry fruit is pierced in search of water, the wound allows the entrance of infection hypha which cause the damage recorded by Neiswander (1944). Without water Myodocha dehydrates and dies in a day. Myodocha does not exhibit a complex “seed territoriality” behavior with a threatening stance. Seeds, if large, are dragged to secluded sites. If the seeds are small, as with Hypericum seeds, the seeds are carried at the end of the labium and beneath the body. Once when a seed was lodged in some methyl cellulose a female used a three quarters open fore leg draped over the seed to dislodge it. This suggests that the fore legs as in other species may have this function in the natural habitat. This species mates quite infrequently and the courtship behavior 94 VOLUME XLIII was only briefly observed. The male apparently does not recognize the female until after contact, and even then only after repeated contacts. When excited, the male vibrates his antennae very rapidly, protrudes his genital capsule, and attempts to climb upon the female who usually evades him. It is noteworthy that the male spends a good deal of time “grooming” himself. The males have a long spur on the fore femora which may function in the mating behavior. Life History Myodocha serripes has a bivoltine life cycle as Neiswander (1944) predicted. The adults overwinter (Blatchley 1895, Wirtner 1905, Torre Bueno 1908, 1925, Dowdy 1955). Blatchley (1926) states that the fourth and fifth instar nymphs may also overwinter. In New England, however, I have found only the adults overwinter- ing. The adults appear in the fields in May at Storrs, Connecticut with the earliest date May 10. The observed phenology at Storrs is as in Table 5. The percentage of adults in the field in September is of little significance as the adults migrate into woodlands from August 25 on. The two generations overlap, as the early instars of the second generation and ovipositing females of the first generation occur together with late instar nymphs of the first generation. Adults collected in late August and September were in reproductive dia- pause. TABLE 5 Phenology of Myodoclia serripes Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adults June 1 June 13 June 19 6.2% 30% 62.5% 21.3% 100% 70% 10% June 28 12.5% 45% 25% 12.5% 5% July 6 July 15 10% 12% 38% 23% i% 10% 9% 80% 17% Aug. 3 10% 20% 20% 20% 30% Aug. 20 Aug. 31 Sept, 20 Oct. 10 5% 1% 20% 29% 10% 45% 45% 34% 50% 30% 25% 56% 50% 95 ENTOMOLOGICA AMERICANA The late summer cessation of reproductive activity and mi- gration into woodlands well before the advent of cooler weather sug- gests that the diapause state is initiated by a short photoperiod. Members of a population sample collected on March 21 were in reproductive diapause. As fertilization occurs during the spring, the function of photoperiod could be assessed. When kept under equinoctal daylight (12 hours) the females remained in reproductive diapause. Under long day (16 hours) an aliquot of overwintered adults came out of diapause after 16 days and reproduced several months before the field and greenhouse populations. It is note- worthy that copulation occurs quite independently of reproduction and that the males do not exhibit reproductive control by photo- period under these conditions. In the fall a somewhat different situation exists. When popu- lations from late July and early August are brought into the labora- tory and kept under long daylight they do not enter into the facultative diapause. However, when exposed to long photoperiod (16 hours) or exposed independently to cold the insects did not come out of diapause. The diapause was a strong one, and if left in warmth during the autumn, the insects died without reproducing in the laboratory. Only after a combined cold-long photoperiod exposure could diapause be broken. The exact day length which initiates diapause was not ascertained, but the field evidence ap- pears to indicate that it is less than the summer solstice (15 hours 15 minutes at 42° latitude), and is near 14 hours for this day length coincides with the earliest (May 15) and latest (early Au- gust) occurrence of gravid females. In the laboratory, the mean stadii and extremes were as in Table 6. Neiswander (1944) stated that under experimental conditions the newly hatched nymphs reached the adult stage in from 19 to 25 days, and that under field conditions probably twice this period was required for development. The phenology data indicates that in field conditions at Storrs, Connecticut, one month or a little more is required for this development period. The laboratory life cycle happens to be in essential agreement with this interval, rather than more rapid as is usual in other species. My longer development periods than Neiswander may result from the more moderate (75°F.) laboratory temperatures against higher field temperatures. I was unable to determine the precopulatory or preoviposition period of a non-diapause female. Release from diapause under long photoperiod (16 hours) took 13 days as measured by oviposi- tion time. As remarked under diapause, oviposition began by mid- May and continued normally until death in mid-June. If the fe- 96 VOLUME XLIII TABLE 6 Stadia of Myodocha serripes Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10.5 (10-12) 7 (6-9) 7 7 7 10 37 (6-10) (6-10) (6-10) (8-12) (27-41) male is not fertilized she may live for a very long time without laying eggs. One virgin female collected April 14 remained alive until April 29 of the next year without laying any eggs. Under 16 hour illumination, other virgin females sometimes laid a few (mean 9.4 (0-26) ) eggs. The mean normal fecundity in the labora- tory was 107 eggs, the range 55 to 161. The rate varied from 4.5 to 10.1 eggs a day (mean 6.8). Contrary to Neiswander (1944) the eggs are not laid at random. The female carefully examines a site with her antennae and then probes it repeatedly with her ovipositor. This may be repeated several times until a suitable location is found. In the laboratory, the smooth, unadorned, elongate eggs were laid singly into the soil, crevices in litter, under small stones, and into methyl cellulose and cotton. Heraeus plebejus Stal The genus Heraeus differs from Myodocha primarily in that the head is only constricted behind the eyes rather than elongated into a slender “neck.” Its appearance similarly suggested to Barber (1923) that the species was at least partly predacious. In view of the apparent close taxonomic relationship of these genera, it is perhaps significant that H. plebejus shows distributional, ecological, and bionomic similarities to Myodocha serripes. H. plebejus appears to be the only species of the 10 known which is adapted to a cool temperate climax. Most of the other species, insofar as is known, are found in Central America and the West Indies. Only one, H. cincticornis Stal is recorded from South America (Bolivia and Argentina) and another, H. pacificus Bar- ber is endemic to the Galapagos. H. plebejus has a wide range, extending north to Quebec, south to Florida, Bahamas, and Haiti, and west to Kansas and Arizona (Slater, Catalogue). In New England, I found the species to have a scattered and 97 ENTOMOLOGICA AMERICANA spotty local distribution, and did not collect the insect north of Connecticut and Rhode Island. Parshley (1926) had no records of the species north of Massachusetts. Blatchley (1926) considered it scarce in Indiana, especially northward. Moore (1944, 1950) however has recorded the species from Quebec. Nevertheless, H. plebejus is at the northern fringe of its range in southern New England. Consequently, the biology of this species should even- tually be amplified by work nearer to the center of its distribution. Environment Several authors found Heraeus to overwinter as an adult : con- cealed in moss in January near Buffalo, New York (Van Duzee 1894) ; sheltering under boards and other loose objects (Barber 1923); and under stones in December (Torre-Bueno 1910). In Florida, Blatchley (1926) captured numerous specimens in winter by beating Spanish moss and the dead leaves of the cabbage palmetto. In Connecticut there is an annual movement to woodlands in early autumn to hibernate and a redispersal in spring into open habitats, as in Myodocha. While these two species are rarely found together in summer habitats, they frequently hibernate together. In fact every hibernation capture of Heraeus in late fall has been with Myodocha, at edges of light mesic woodlands. The earliest fall record in woodlands is September 9, and only nymphs and very few adults remain in the summer habitats during September. In the spring, the species reappears in open mesic habitats, but very in- frequently re-establishes itself in the same locale occupied the previous year. Of course, this temporary presence in a habitat in part results from the scarcity and the dispersal pattern of the species, but also and perhaps more important is its preference for a particular open, mesic, rank forb type of habitat. It is also found along streams and shaded ecotones between woodlands and fields or marshes. Torre-Bueno ’s (1910, 1924, 1925) records of Heraeus from near swampy areas, and perhaps Hussey’s (1922) collection of it in a sassafrass strip between white pine and black oak communities cor- respond to these observations. In the summer habitats, only four large colonies of Heraeus with nymphs were found. More frequently, only a few specimens are found at a site. These favorable habitats are all open rela- tively moist ecotones between fields and marshes or mesic wood- lands. They agree in having moist soil (8-9), dark with humus, which is shaded by the herbs and so remains relatively cool (no more than 75° F.) on hot days. As I was fortunate enough to visit 98 VOLUME XLIII each of these areas earlier, the colonies definitely persisted for only one year. Two of the habitats were similar, as ecotone of rank forbs between a drier field slope and a marsh stream with wood- lands 30-50 feet away. The plant association consisted of various tall grasses, the rank forbs Impatiens biflora Walt, Eupatorium purpureum L., E. perfoliatum L., Galium sp., Bidens laevis (L.), Solidago patula Muhl., Solidago sp., and in one area, a few young Sambucus canadensis L. The third habitat was on the edge of a but- ternut-maple-oak wood with a drier field above it, and the fourth favorable habitat was at Rocky Neck State Park, Connecticut where the insects were found among rank tall grasses leading down a slope toward a sheltered salt marsh, but well above and away from the influence of the salt level. Among the various locations where Heraeus occurred in low abundance, several may be mentioned. One was a thick strand of flood litter on a partly shaded bank at Mansfield Center, Connecti- cut. Here, in the seed-rich litter, it occurred with three other rhyparochromines, and it was significant that in the drying litter Heraeus occupied the wettest level while the other species occupied drier levels. At Noank, Connecticut a surprisingly dry habitat (soil 4-5), was a sandy roadside with scattered clumps of Andro- pogon scoparius. As mentioned earlier, a study at the center of its range should be made, as I have found it in June upon the cool grassy balds at 5,000 feet on Mt. Pisgah and Mt. Mitchell, North Carolina in the Great Smoky Mountains, as well as in roadsides in the valleys at 1,000 feet. Not only does Heraeus have a scattered scarce overall distribu- tion, but it is found in populations of low densities, usually 1-3 per square meter, at most, 8 per square meter, and the total colonies are of small size, covering no more than 10-20 square meters. General Biology The seasonal movements and preference for a relatively short lived serai stage, suggests that the bug readily disperses, which cor- relates with its entirely macropterous condition. H. plebejus may migrate at night at it has been taken at lights in New York by Torre-Bueno (1908) on July 17, 1907, and subsequently (1930). Blatchley (1926) collected it on March 15 and October 27 at Dune- din, Florida. I collected the species at lights July 10, 1957 at Noank, Connecticut and July 25, 28, 1960 at Storrs, Connecticut. Another species of this genus, H. guttatus was captured in Puerto Rico at lights (Ramos 1946). The species may be capable of larger dispersal movements at Glick (1939) captured this species up to an altitude of 1,000 feet over Louisiana. 99 ENTOMOLOGICA AMERICANA Like Myodoclia, the adult Heraeus has a dark color pattern with white pattern-breaking markings on the lateral margin of the corium and apex of the membrane. The bug ordinarily moves slowly and deliberately and blends well into its natural habitat. The nymphs are conspicuously colored with a white band along the Y-snture which gives a good impression of ant mimicry when the insect runs. When the nymph is motionless, however, the color pattern breaks up the form and renders it difficult to see. The first instar has a longi- tudinal mesal red band as well as the usual transverse red band across the yellow abdomen which is typical of the Myodochini. Unlike Myodocha, Heraeus is parasitized by the tachinid Cath- arosia nebulosa (Coq.). All individuals were reared from diapaus- ing adults of Heraeus. The parasites emerged from the host in the laboratory at the same time that Heraeus ordinarily breaks dia- pause. Thus the parasite overwinters in the host bug. Feeding was not observed in the field. In the laboratory aside from sunflower seeds, it feeds upon various small seeds : Galium sp., and Monarda sp. from its habitat, and also upon small composite seeds and grass ( Festuca and Andropogon) seeds. It would seem to be a general seed feeder but more critical work needs to be done. I have reared it through the life cycle only on sunflower seeds on which the bug feeds very readily. However the mortality on this seed is very high (90% ), most (ca. 70%) dying in the first instar. H. plebejus dehydrates and dies very quickly away from water. While I frequently observed mating end to end, I have not been able to record courtship or initiation of mating. Nor did this species show any manifestation of seed possession or aggressive dis- play as in many other rhyparochromines. These seeds are moved to more secluded places for feeding. Life History Heraeus plebejus has two generations a year. The earliest over- wintered adults were collected in late spring (June 7 and 9) and were already gravid and laying eggs. Blatchley (1926) collected the bug on March 15 in Indiana. Only one population was followed as it developed. Its phenology is given in Table 7. On July 25 some new females of the first generation were gravid. However, all adults of the second generation were in repro- ductive diapause. When brought into the laboratory, they remained in diapause until October 10 or later. Evidently the dia- pause is not a strong one and the adults become reproductive after only two or three months in warmth. This short cessation, how- ever, serves to suspend reproductive activity until after the advent of cold weather. It is interesting that the females of the first gen- 100 VOLUME XLIII eration, when sexually isolated, would survive until October but did not go into reproductive diapause like the second generation. One was mated with a second generation male which broke diapause in early October and a few fertile eggs were produced. This re- productive cessation is probably initiated by a short photoperiod as in Myodocha. TABLE 7 Phenology of Heraeus plebejus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 7 — 100% June 21 11% 11% 34% 22% n% 11%* J uly 5 10% 20% 40% 30% July 25 23% 77% Aug. 2 30% 20% 15% 35% Aug. 20 10% 15% 40% 30% 5% Sept. 11 5% 15% 25% 65% * new adults The life cycle in the laboratory correlates with the field notes. The stadia were measured accurately only up to the third instar. Beyond that, too few developed and too erratically to give a mean- ingful average. TABLE 8 Walmer 12-2 Stadia of Heraeus plebejus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10 9 10 11 5 8 53 (9.5-11) (7-13) (8-12) (8-13) (5-14) (42-68) The adult longevity varies with generation, of course, and for the summer generation it is two months, and for the winter genera- tion (overwintered), nine months or, in laboratory warmth, four months. One female after breaking diapause on October 18 laid 455 eggs 101 ENTOMOLOGICA AMERICANA at a rate of 7.6 eggs a day over two months. The number of eggs per female varied from 30 to the above figure but the sample is too small to give an accurate average fecundity. Egg production occurs regardless of fertilization. The mean preovipositional period of four virgin females was 18 days. A female mates repeatedly any time during her reproductive period, but one mating suffices to fertilize all eggs to be laid. The smooth elongate eggs are laid singly in the laboratory into loose sub- strates as sand, loose soil, wet or dry cotton or methyl cellulose and are not laid at random or loose in the culture dish. The female searches assiduously, prior each egg laying, for a proper site. LIGYROCORIS Stal In New England four species of the genus Ligyrocoris occur, one of them described only recently (Sweet, 1963). There is some controversy concerning the limits of this genus as some authors (Van Duzee 1914, Ashlock 1957, Ashlock and Lattin 1963) have questioned whether the possession of the lunate or stigose abdominal vitta (stridulitrum of Ashlock and Lattin 1963) indicates a natural genus. Nevertheless the four closely related species treated here should remain nomenclaturally unaffected by any further generic changes because among them is the type species for the genus, Ligrocoris sylvestris (Linn. ) . As sylvestris and diffusus Uhl. were confused by synonymy, and depict us Barb, not yet recognized, American literature prior to Barber’s 1921 revision of the genus must be used with great caution. Distribution At present the genus includes two subgenera, Pseudopamera Dist. and Ligyrocoris Stal. Pseudopamera, originally established as a separate genus, includes nine species and is restricted to south- western North America and Central America. Ligyrocoris ( sensu strict o) includes fourteen species and while also apparently centered in Central America, has a much wider distribution. Although several species of Ligyrocoris extend to South America and the West Indies, the present complex of species is apparently adapted to a temperate climate. While sympatric in New England, the ranges of the four species to be considered differ greatly; L. sylvestris is evidently a boreal form (Krogerus 1960) and is widely distributed over the north Holarctic region. The very closely related L. depictus Barb, occurs from Quebec south along the Appalachians to North Carolina. L. diffusus Uhl. extends across North America, roughly between the 102 VOLUME XLHI 38th and 50th parallels and broadly overlaps on the southern limits of L. sylvestris. The fourth species, L. caricis Sweet is so far limited to one locality each from Connecticut and Maine. The origin of this pattern of distribution and overlap poses an intriguing problem, but an analysis must await a better knowledge of the range of the species, especially in southeastern and southwestern United States. Ligyrocoris diffusus Uhler This is one of the most abundant and widespread of the rhyparo- chromines studied. Its range extends as far north as Newfound- land (Lindberg 1959), and the north shore of the Gulf of St. Lawrence (Walley 1932) ; west to British Columbia and California (Barber 1921) ; and south to Missouri (Froeschner 1944) and North Carolina (Barber 1921). Blatchley (1926) noted that it was fre- quent in the northern counties of Indiana, but had not been taken in southern Indiana. Slater (1952) said that diffusus is one of the most common members of the subfamily in the midwest, but scarce in southern Illinois. Over this wide range the species exhibits some variation in size and darkness and may eventually be separated into subspecies. Specimens from northern New England and the Adirondacks are definitely smaller and duskier, but the differences overlap greatly those of specimens from Connecticut. As the specimens from Connecticut closely resemble those from the Midwest it may be that these represent a large western population that has moved eastward with the creation of extensive open habitats in eastern United States. While it is true that Barber (1921) could say that he had never seen a brachypterous form of this species, wing lengths are not uniformly the same. Smaller female specimens from the northern and western limits of its range frequently exhibit a slightly shorter hemelytra which, while exceeding tergum seven, does not exceed tergum eight, the apex of the abdomen. The hind wing in each is slightly shorter than the hemelytra at rest. In a very few males the membrane may not quite attain the apex of tergum seven, the apical segment. These slightly brachypterous forms also have a narrower pronotum with a proportionally larger anterior lobe. Environment This species is abundant throughout New England, but it is ecologically restricted and its abundance is in large part an expres- sion of the abundance of its preferred habitats. While in reality it is primarily a ground-living insect, it frequently ascends plants 103 ENTOMOLOGICA AMERICANA to feed on seeds, and so has often been recorded as collected by sweeping which greatly contributes to its 4 ‘ abundance. ’ * Sweeping and ground collecting each produces a very different impression of the species’ habitat preferences. Hussey (1922) in Michigan said it was abundant in fields, especially on goldenrod and ragweed. Blatchley (1926), in Indiana, swept it from ‘Hall grasses and other herbage along margins of wet meadows and marshes. ’ ’ Froeschner (1944) in Missouri swept it from weedy fields and open woods between July 1 and October 15. Hendrickson (1930) in his study of insect populations in relatively undisturbed prairie communities in Iowa made more detailed observations. He swept diffusus from all communities, and found it common to numerous in both climax and subclimax grasslands, but absent from wetter consocies. Sig- nificantly, he noted diffusus as especially numerous on Andropogon furcatus associes where herbacious plant flowers attracted many of this species. Lindberg (1958) however stated that the species was found in Newfoundland in humid localities, along margins of lakes and pools, on wet meadows, and Sphagnum bogs. In New England, this species is typical of open disturbed habi- tats, roadsides, as well as old fields with many forbs present, especially composites. In the latter habitat diffusus is fre- quently collected by sweeping, for it prefers the seeds of certain composites such as Chrysanthemum leucanthemum L., Solidago spp., Tanacetum vulgar e L., and especially Rudbeckia serotina Nutt. In late summer when the seeds are ripe the insects will congregate on the ripe seed heads. In such a site where it is readily collected and appears abundant, ground sampling will frequently indicate a lower overall abundance at this site than at a nearby waste habitat. Indeed, diffusus will be apparently absent from the tall f orb habitats earlier in the year, when abundant in other habitats. Barber (1928b) found it under ground cover and Procter (1946) said it was to be looked for under boards, etc. These notes do little justice to the great predominance of this species on the ground. Nearly all collections of diffusus have been in relatively open un- shaded habitats, especially on level exposed areas. In general the population density of diffusus decreases as1 the herbacious level becomes more dense and shades the ground surface. The greatest densities occur in areas with pioneer stages of herbacious vegetation frequently in ruderal communities on poor sandy soil which are aptly described as wastelands. Frequently the margins of culti- vated fields and recently fallow fields support high densities of this insect. Plants found in such preferred sites usually include Ambrosia artemisifolia L., Silene spp., Daucus carota L., Agropyron repens (L.), Potentilla recta L. and P. canadensis L., Rumex ace- 104: VOLUME XLIII tosella L ., Panicum spp., Polyganum sp., Stellaria sp., Chrysanthe- mum leucanthemum L., Tanacetum vulgare L., Oenothera spp., Verbascum thapsus L., Erigeron sp. ; in short many of the rapidly invading plants which are commonly called weeds. In sparser areas abundances may reach 30-50 per square meter. By far the most favorable site was a corn field 40 by 150 meters, left fallow for a year. The ground was heavily littered with corn husks, stalks, and cobs, fallen stems of grasses and composites, and in mid- June had a heavy population of 50-70 adults and fifth instars of diffusus per square meter. Areas nearly or completely devoid of plants are not colonized unless a good deal of litter is present. Also, as the proportion of grass to forbs increases, the numbers of L. diffusus decreases. I never swept this species from grass, and it is completely absent from the dry Festuca-Andropogon-Cladonia habitats. The great majority of collections of diffusus are from habitats with but a thin sparse layer of litter, which, in the open conditions prevailing, is very dry. Surface soil on the moisture scale varies from 1 to 4 with an average dryness of 2.5. As is to be expected such habitats are very hot in early afternoon, with surface soil temperatures up to 140 °F. This strong insolation very likely accounts at least in part for the rapid spring development of this species as compared wth many other rhyparochromines. In autumn when diapause eggs are laid, a definite change in habitat preference occurs as the weather becomes cooler. The adults are then frequently found in more shaded marginal areas than during the summer, and the soil moisture level is usually 5-6. It is important to note, however, that such areas in the following spring are unshaded, quite warm (80°F.) and drier (3-5) at mid- day as the leaves are small and herbs have fallen during the winter leaving a seed rich litter. For over five years several new fields in Storrs, Noank, and Canaan, Connecticut were left fallow and in the invasion, expansion and subsequent decline of populations of diffusus could be followed. This sequence strongly shows a peak abundance at an early sere succession stage and subsequent tapering off as herbs, especially grasses, more thickly cover the ground. General Biology As the herbs grew and filled in the habitats open in early spring, the later instars and new adults often dispersed into sur- rounding, more sparsely vegetated, xeric areas. (Frequently adults and older nymphs dispersed into nearby flood wash litter along river banks). In late June and early July populations appear in 105 ENTOMOLOGICA AMERICANA habitats — frequently very new — which earlier did not have diffusus present. This, combined with its ubiquitous presence in fallow fields, indicates that it is a rapidly dispersing species. It has been collected at lights (Torre-Bueno 1930) but apparently much less frequently than might be anticipated from its abundance. L. diffusus is one of the most abundant species found in beach wash (Torre-Bueno 1915, 1927, Parshley 1917a). Torre-Bueno sug- gested that such insects were on dispersal flights which happened to lead over water. The nearly total macroptery attests to the importance of dispersal in this species. It would be interesting to see if the slightly shorter wing forms have reduced flight or dispersal ability. Two female specimens were collected above the tree line at 5,400 and 5,500 feet respectively on the roadside of Mt. Washington, New Hampshire on August 9, 1961 and had probably dispersed there from the valley floor where the species was abundant along road sides. As in many “weedy” fields which apparently had no diffusus present earlier in the year it is probable that these were dispersing adults. The mottled brown coloration of L. diffusus allows it to blend well with the thin litter and sandy soil of its preferred habitats. As it is not a sub-litter inhabitant and often lives in areas normally with little litter, its main defense is to run and freeze near some object or to crawl into loose litter. The nymphs of instars two to five are fuscous with the abdomen marked with small white spots. The second to fourth instars have a lateral white band along the suture between the third and fourth segments, which, when the nymph runs gives a definite impression of ant mimicry. Balduf (1939) found L. diffusus preyed on by Phymata penn- sylvanica Melin which gains significance when it is recalled that Phymata waits in ambush on flowers. I have fed diffusus to Nabis femes (L.), Pagasa fusca (Stein), and Melanolestes picipes (H.-S.). I have found diffusus parasitized by a small undescribed tachinid of the genus Catharosia. The degree of parasitism was low (less than 5% ), and only a few diffusus populations yielded any taehinids. As mentioned earlier (Sweet 1960) L. diffusus is fairly unusual among the rhyparochromines for it does on occasion climb into plants to feed on ripening composite seed heads. Rudbeckia sero- tina Nutt, was by far the most preferred host plant. When ob- served feeding in the field on a seed head, the insect had pierced the achene through near the pappus, and the usual pumping mo- tions of the head indicated its feeding. Such feeding has been observed on Tanacetum vulgare L., Chrysanthemum Leucanthemum 106 VOLUME XLIII L., and Solidago spp. This feeding on seed heads explains the relatively late date of recorded collections: July 23 (Hendrickson 1930), July 1 (Froeschner 1944), July 15 (Procter 1946), and July 11 (Blatchley 1926), which corresponds to ripening of composite seed heads. In the field the nymphs feed on fallen seeds of the previous year as indicated by several rearings of the insects on litter from the collection locality. By laboratory observations it was found that the nymphs and adults feed on the seeds of Rumex acetosella, Rumex sp., Polyganum sp., Potentilla recta, P. canadensis, Fragaria vir- giniana Duchesne, Acer rubrum L., Medicago sp., Trifolium spp., Monarda sp., and some other unidentifiable seeds, but grass seeds and Daucus carota seeds did not elicit a feeding reaction, which correlates with the absence of diffusus on these plants. The most vigorous feeding response was toward Rudbeckia and sunflower seeds. In general, L. diffusus should be considered as oligophagous on seeds, especially of the Compositae. In this connection, it may be relevant to note that Rudbeckia serotina is noted by Fernald (1950) as a most aggressive Great Plains species which has rapidly spread eastward with the clearing of the forests. The midwestern population of L. diffusus may have moved east in company with Rudbeckia . There are several other references in the literature which men- tion the presence of diffusus on a plant but these do not indicate definite feeding. Phillips (1951) found it on sour cherry in orchards; Adam (1915) collected it on Lepachys (= Ratibida sp.), the prairie coneflower ; Forbes (1905) found it on corn silk. Vestal (1913) described diffusus as a non-selective plant feeder. In the field diffusus was never observed to feed on any part of the plant other than the seeds. It fed readily on lettuce for water but no eggs were produced. With the addition of a vial of water such “feeding” was greatly reduced. The addition of seeds brought about egg production ; the removal, a cessation in egg production. L. diffusus was reared through several generations on Rudbeckia seeds and numerous times on sunflower seeds. The culture results were quite variable, very likely because of disease interference. Yet nymphs of any instar could be reared quite easily to adults which oviposited vigorously in culture. However nymphs reared from the eggs, suffered very high mortality on the same food. Most of the mortality occurred during the earlier instars. Not only was this species in no way observed as actively preda- cious, but it did not probe at a dead or dying insect as shown by an absence of any cones of salivary sheath fluid on the dead insects. It is therefore interesting that Slater (in lift.) found one feeding on 107 ENTOMOLOGICA AMERICANA a large black aphid. Like other rhyparochromines diffusus fares badly away from water, thus it is interesting to speculate on the source of water available in its dry habitat. Vegetation is an obvious choice, but dew, which condenses heavily in such open habitats is very likely quite important. In various dry habitats diffusus was found active until early afternoon on very hot days, when, the insect retreated into clumps of litter in late afternoon. I frequently observed that the nymphs had slightly shrunken abdomens by late afternoon and specimens from the field at this time did not feed unless water was provided, in contrast with those collected in the forenoon which fed readily on sunflower seeds. This would indicate, then, that the insects were dehydrated. It also strongly suggests that dew is the im- portant water source, especially as free water was much preferred to water from plant tissues in the laboratory. When in thirst, however, L. diffusus will pierce nearly any green plant in search of water. As in other members of this genus and the related Zeridoneus a fertile male reacts to a receptive female with a sort of courtship dance which probably involves stridulation. The peculiar jerky movements of an excited male are produced by a rocking of the body back and forth while at the same time the male jerkily moves forward and moves his antennae in an uneven tapping or shaking fashion. The male does not react unless he comes in contact with the female, in contrast with some other rhyparochromines which react to the female before actual contact. The female usually resists the excited male by turning on him with rapidly vibrating antennae which is an annoyance signal, and moving away a short distance. This may be repeated several times as the male endeavors to mount her from the side. Finally the female may become quiet and the male, still continuously jerking, mounts her, usually on the right side. The extruded pygophore is turned over 180° and placed upon the apex of the female’s abdomen. At this point the female may release the ovipositor and the claspers, which work like little claws, then grip the apex of the ovipositor, and copulation ensues. The male swings away into an end to end position. With considerable pulling and pulsating the male slowly forces the aedeagus into the genital chamber and the spermatheca. The slow course of intromis- sion can be observed readily as the spines of the conjunctiva are clearly visible through the semi-transparent valvulae of the oviposi- tor. Length of copulation varies from 1 to 4 hours. Repeated attempts to cross L. diffusus wth the other species of 108 VOLUME XLIII Ligyrocoris, showed that the males, except caricis, reacted with the various females in very similar fashion, but that the female after a few usually preliminary circlings became extremely excited if the male was of another species and she ran very rapidly and 1 1 frantic- ally” about the dish as the alien male jerkily “danced” about. In only one case, a diffusus female of the second generation mated with a sylvestris male, when the male was dropped suddenly beside the female. The male made copulation before the female could react to it. Repeated attempts to duplicate this have failed, and it seems highly unlikely that this would occur in the unconfined natural environment where ecological factors also keep the species apart. Neither L. caricis male nor Zeridoneus male seemed to recognize the diffusus female, but L. diffusus male was very inter- ested in the tiny caricis female, but on the other hand, was not stimu- lated by a Zeridoneus female. The annoyance waggling of the antennae mentioned earlier plays an important part in the “seed possession” behavior which is strongly developed in diffusus. By starving the insects for a few days this behavior can be readily elicited. When first disturbed by an intruder, the insect (usually female) simply places itself between the intruder and the seed. Further intrusion results in a rapid alternate waggling of the antennae and vigorous attempts to keep the intruder away by kicking at it with the hind legs. If this fails the seed possesser will leave the seed and rapidly charge on the intruder flailing at it with antennae and fore legs. The fore legs are not spread as in Pachybrachius , but two may flail at and “fight” one another actively until one ceases to intrude. The insect may return to the seed, carefully search it over with its labium until the original feeding hole is found, and resume feeding. Life History L. diffusus j in New England, overwinters as an egg and has two generations a year. The statement by Forbes (1905) that it over- winters as an adult and Torre-Bueno’s collection (1924) of the species in New York by sifting in December are very likely the result of the cold hardiness of diffusus. It lays eggs until very late in fall, and the latest ovipositing females were collected on November 10, 1959 and a male as late as November 29, 1960. The eggs hatched in spring at a staggered rate, for while some appeared as early as April 25, 1960 first instars continued to be present on May 14, 1960 and May 27-29, 1959 when some nymphs were already just becoming adults. Thus all instars would be found in the field in May although the mode instar changes, as 109 ENTOMOLOGICA AMERICANA shown by the instar distributions in Table 9. In the first generation adults, mating was observed in the field on June 24. The adults from duly 10 on represent the second generation. The adults of this second generation remain sexually immature until the last week of August. It is interesting that in a cool valley at the base of Mt. Washington, New Hampshire the second generation appeared much later and in late July as only first to third instar nymphs were present, and first instars were still present on August 3. Last instar nymphs of the second generation were found as late as August 26 in Storrs, Connecticut and Septem- ber 8, in Canaan, Connecticut and evidently occur much later in New Hampshire. TABLE 9 Phenology of Ligyrocoris diffusus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult Apr. 29 93% 7% May 7 50% 35% 10% 5% — May 17 13% 50% 25% 12% June 9 10% 15% 35% 40% June 16 — 20% 80% June 22 100% June 28 50% 30% 20% July 5 55% 30% 15% July 10 24% 35% 17% 15% 8% 1% July 17 5% 5% 10% 30% 35% 15% July 27 5% 20% 35% 40% Aug. 7 15% 25% 60% Aug. 26 — 8% 92% Sept. 10 100% As the second generation adults do not become reproductively active in the field until late August or the beginning of September, this indicates that the early maturing adults are in a reproductive diapause for a period of a month or more, in sharp contrast with the rapid attainment of reproductive activity (6-8 days) in the first generation. This is then not an immature state in the sense of a normal pre-reproductive period as the late fifth instar nymphs at Storrs, Connecticut and the late developing population from New Hampshire all attained reproductive activity at the same time as 110 VOLUME XLIII the earlier developing- individuals in the southern New England populations. These events, including the assumption of reproductive activity between August 29 to September 6, occur well before the advent of cold weather and also in field populations exposed to normal day length in the laboratory. Moreover the second generation colonies when reared under long photoperiod in the laboratory, laid non- diapause eggs in early and mid-August. These results indicate that a photoperiodic response is involved in this seasonal cycle. How- ever the reaction of the first generation to short photoperiods has not been tested. These results may be most readily explained as follows. There is no reproductive diapause under summer solstice long photoperiods (15.5 hours) when the first generation matures. In shorter photoperiods from late July (14 hours) to late August, the adults remain in diapause. After late August (13.5 hours) diapause eggs are laid. The picture which emerges is a species very finely adjusted to its environment. Its bivoltine life cycle coupled with actively dispersing macropterous forms allows it to readily fill the transient sere it prefers, and egg production in the fall allows the females to make use of the ripening composite seeds, and place the diapause eggs around the site of such plants. The emerging nymphs can make use of the fallen seeds before the seeds sprout or succumb to molds. The diapause eggs are darker and slightly larger (ave. 1.07 mm.') than non-diapause eggs (ave. 0.98 mm.). The diapause eggs de- velop until the eye spots are visible through the chorion. At this time, coincidentally at the blastokinetic stage Wheeler (1893) termed diapause, physiological diapause intervenes. The diapause is a fairly strong one, but its strength differs in various females when the eggs are left in warmth ; in some females, a few eggs would resume development in two weeks; in other females, none would hatch until 60 to 150 days later. Eggs from adults collected in Laconia, New Hampshire did not develop at all in warmth, but required a cold exposure. Diapause development occurs in cold (35°F.). Nymphs hatched from the diapause eggs of April 14 after restoration to warmth March 21. The development, how- ever, was irregular, and some eggs did not hatch until early May. This variability correlates with the field observations of apparent staggered egg hatching. It would seem that this would be an excellent physiological mechanism to scatter the risk of hatching during unfavorable weather conditions. Since a few eggs of a few females did hatch early, it is interesting that several third instars were collected in early October and certainly represent a 111 ENTOMOLOGICA AMERICANA field example of an early diapause release. It may then be that three generations will be present further south. Two females which were collected at the late dates of November 8, 1959 and November 10, 1960 each laid 50-60 eggs all of which did not diapause. This indicates that the exposure to cold had broken the diapause initiating phenomenon in the female before the eggs were formed. TABLE 10 Stadia of Ligyrocoris diffusus Egg* Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10.0 7.6 6.6 4.5 6.7 8.2 30.6 (7-12) (6-8) (3-11) (3-6) (3-10) (5-9) (22-40) ^Generation 1 In the laboratory the stadia (in days) are quite variable (Table 10). The more rapid development always occurs in nymphs taken from field and slower development occurs in nymphs which are reared from eggs in the laboratory. The life span of the adults varies with the generation and mating. Fertilized females live 19 to 31 days, males and unfertilized females, 45 to 75 days. In the second generation with its brief diapause period fertilized females live longer, 44 to 78 days, unfertilized adults 60 to 85 days. The precopulatory and preoviposition period for the female is eight days, and the precopulatory period for the male is four to seven days. There is no marked difference in egg productivity in females of each generation. The productivity varies in 26 mated females from 51 to 325 eggs (mean 166.5). In 11 unmated females the number varies from none to 29 eggs. In some sexually isolated females of both generations, a few eggs are laid at the beginning of the normal oviposition period. Laying soon ceases and the abdomen of the virgin females becomes very distended. Dissection showed well developed eggs. The egg-lay- ing stimulus is probably copulatory as the following two experi- ments illustrate : ( 1 ) a twelve day old virgin female when mated with a three day old immature male immediately laid a normal, but infertile complement of 29 eggs in three days, after which the female ceased laying eggs and once again became swollen; (2) a virgin female mated with a very old male (45 days) produced a normal 170 but infertile complement of eggs. This suggests that the male 112 VOLUME XLIII may transmit some substance to the female which induces oviposi- tion, and which was in low quantity in the new male, but present in the old male which apparently was infertile. The females lay their eggs singly into litter, on grass culms, hollow stems, and in soil, especially loose sandy soil. The eggs are cylindrical, slightly curved (cucumber-shaped) and beset with tiny clubbed hairs. Small sand grains and other debris cling to the egg, concealing it. Wet substrates are shunned unless no others are present. The females prefer to lay their eggs deeply into sand or such loose dry substrates as cotton and methyl cellulose, and will even work their abdomens into the substrate to insure deep place- ment of the eggs. On sand the females spread their legs wide, tilt their abdomen to 80° from the horizontal, and work their entire abdomen into the sand. After each egg is laid the female makes several jabbing movements which tumble sand into the oviposition site. Egg production becomes much retarded and reduced when no suitable substrate is provided. Such females become greatly swollen and egg production may entirely cease. Ligyrocoris depictus Barber This species, which was not recognized until 1921, very strongly resembles sylvestris and differs from it chiefly in coloration and slightly different labial segment ratios. From the type localities of New Jersey, Massachusetts, and New York it has been found north to Quebec (Moore 1950) and Maine (Procter 1946). I have further found the species in Connecticut, New Hampshire, and also in the Great Smoky Mountains of North Carolina. It has an Appalachian distribution. It is to be noted that the specimens from lowland habitats — New Jersey and Storrs, Connecticut — are much the same in size and coloration, while the specimens from the mountainous areas are larger and darker and more nearly resemble sylvestris. The habitat selection of this species is entirely different from that of L. sylvestris, and the two were never found together. Environment Despite the apparent rareness of L. clepictus, it is actually a fairly common species on mountain bald habitats with Xestocoris nitens V.D., Carpilis consimilis Barb., and Trapezonotus arenarius L. It is also found on similar lower slopes where ericaceous shrubs, principally Vaccinium angustifolium Ait. along with Danthonia spicata (L.), Aristida dichotoma, and open grasses form a relatively permanent, long persisting open habitat. The habitats of the balds varies from low dense ericaceous shrubs 113 ENTOMOLOGICA AMERICANA mostly V accinium on Mt. Everett and Mt. Greylock in Massachu- setts, to grass, blueberry, and lichen margins around large open rock outcrops among scrub oak ( Quercus licifolia Wang.) on Canaan Mountain, Connecticut and to a nearly pure grass and sedge herb layer around Rhododendron catawbiense Michx. clumps on An- drew’s Bald, Clingman’s Dome, North Carolina. In Storrs it was found in an old open habitat dominated by fine leaved Festuca capillata community. At Canaan it was also found in a Festuca- Gnaphalium-V accinium community surrounded by white pine, as well as some essentially disclimax old ecotones between pastures and woodlands on xeric morainic hillsides. These habitats for the most part have in common a xeric, open aspect, a relatively open vegetation community with a growth of low ericaceous shrubs. In each the soil is sandy, gray, and very poor with a considerable lichen ( Cladonia ) flora present. These exposed habitats become quite warm (105°F.) on the ground at mid-day in summer but not as hot as in diffusus habitats. In the late summer or aestival period these sites become extremely dry. The species is always present in relatively low densities of 3-10 per square meter and reached the latter density on Canaan Mt. balds. Barber (1928b) collected the species in apparently similar open dry habitats in the Adirondacks. Torre-Bueno (1925), however, swept L. depictus from bunch grass in a marshy place. This record might possibly refer to L. caricis (see L. caricis). Procter (1946) found it on Atriplex patula on Mt. Desert Island, Maine. General Biology As all known specimens are brachypterous or submacropterous, the maeropterous forms must be produced at rare intervals, if at all, and dispersion would be rather limited which correlates with the relative permanency of the habitat. This is a very rapid moving and agile insect which is difficult to collect. When disturbed its escape response is to run rapidly for a considerable distance. It does not actively seek refuge in litter crevices, and would come to rest as often in the open as in some coarse loose litter. When found in the field it was upon the ground litter and not concealed in it. In the resting position the brown and black coloration of the adults appears clearly procryptic. The rather light coloration of this species, in contrast to diffusus, and the more linear effect of the mesal zone of dark coloring presumably better blends this species into its usual dry V accinium — grass habitats, similar to other grass community insects. The nymphs are similar to diffusus except that 114 VOLUME XLIII the white band between segment three and four is pronounced in the last instar as well as in the earlier instars. When running with their characteristic jerky gait, a definite ant mimicry effect is apparent. Only one population found near Mt. Greylock, Massachusetts yielded Catharosia tachinid parasites. The nymphs at least were definitely fed on by Pagasa fusca (Stein) nymphs. Although Rudbeckia was not found in the habitat of depictus, the seeds elicit a strong feeding response in the insect, as do sun- flower seeds. It also feeds on the seeds of V actinium, Betida populi- folia L., and Gnaphium, but is negative to Centaurea seeds. Ap- parently Rudbeckia seeds have a definite odor for when a few are introduced into the culture dish, the Ligyrocoris nymphs present become quite excited and search vigorously until the seeds are found. There was only some cursory probing at fescue and Pani- cum seeds which were the major seed component in the litter of several habitats. Seed possession behavior is similar to that of diffusus , but much more vigorous. Here the third to fifth instar nymphs were also observed to express this behavior. A fifth instar nymph usually effectively dislodges third and fourth instars from a seed by swarm- ing over it with flailing legs and wagging antennae. The possessor often clings flat over the seed when under such an attack. The mating behavior is similar to that of L. diffusus. L. de- pictus is a much more active and vigorous species. The shaking dance was timed at 3.2 motions per second. While a strong avoid- ance reaction is present which would preclude field matings, pro- longed enclosure is also negative. Virgin females of depictus were reared with diffusus males from third instar on, and kept together in a four inch petri dish without any cross mating occurring. Life History This species has but one generation a year and lays eggs which diapause through the late summer and winter to hatch the next spring. It appears evident that this life cycle corresponds to a moist spring and early summer, with dry conditions in late summer, often a severe drought. This climate pattern is strongly accentu- ated in the open, overdrained habitat of this species. The available phenological record is rather heterogeneous. While first and second instars were collected May 17 at Storrs, Connecticut, those collected May 20 and May 21 at Mt. Everett and Canaan Mt. were third and fourth instars. Second instars were collected May 28 at a cool location on Canaan Mt. Fifth in- stars were found with third and fourth instars May 28 on a warmer 115 ENTOMOLOGICA AMERICANA slope. In the laboratory and field adults become mature between June 8 and June 25. Instar ratios obtained are too variable and represent too many different small populations to form a meaning- ful phenological sequence. For example, third instars were present until June 10 in some localities. This variability suggests that staggered hatching occurs here as in diffusus. Oviposition begins between June 6 and June 21 and continues until mid- July and early August. The eggs are laid in a state of strong diapause which commences in early anatrepsis when the egg is still white. The eggs of one female began to show some develop- ment from early August on — these eggs malfunctioned and died before completing anatrepsis. Another female’s eggs in contrast showed no development whatever until October 1 when they all quickly began to develop, but only to early anatrepsis, if at all. I was unable to get adequate hatching under 35 °F. conditions, and in the spring, of 311 eggs, three-quarters of them developed, one- half to katatrepsis, but only one hatched. The stadia in days of field collected instars reared on sunflower seeds were as in Table 11. TABLE 11 The Stadia of L. depictus Instar 2 Instar 3 Instar 4 Instar 5 4 4 5.5 7.0 (4-7) (5-9) This development rate is much more rapid than in the field populations. The longevity of the adults in the laboratory varies from 36-75 days, the average being 60.1 days. Most died by early August, but a few lingered longer in the laboratory, one to August 21. Preoviposition period ranges from 10-13 days (ave. 11.5). I could not induce four old virgin females (32-45 days) to mate which suggests a post copulatory period. 116 VOLUME XLIII Ligyrocoris sylvestris (Linnaeus) This boreal species is one of the few rhyparochromines with Holarctic distribution. Its distribution largely coincides with the taiga biome, and it is absent from the areas of western Europe with an Atlantic climate. It was found from Lappland south into the boreal coniferous zone in the Alps (Hedicke 1942), the Pyrenees at 1,600 meters (5,248 ft.) (Wagner 1958), the Caucasus (Kiritshenko 1918), and Bulgaria (Strawinski 1961). It extends east through Siberia and Alaska and across North America to Newfoundland, and south to New Jersey, northeastern Illinois, South Dakota, and in the mountains to Colorado, Idaho, and British Columbia (Slater, Catalogue). It is, however, conspicuously absent from the highlands of Scotland where suitable habitats appear to be present. Since dis- junct populations occur in the Caucasus and the Pyrenees it is probable that land connection was broken between Britain and the continent before L. sylvestris reached this area. Since sylvestris is a boreal species, it is possible that it represents a post- Wisconsin glaciation introgression of Ligyrocoris into the Palearctic region. As may be anticipated in a species so widespread, considerable variation exists. Horvath (1901) noted that specimens from Siberia had much darker femora than the European representatives. Specimens examined from Anchorage and Fairbanks, Alaska were considerably smaller and lighter colored than the New England population. In L. diffusus the northern New England specimens are usually smaller and dingier with weaker hemelytral fascia. It therefore may seem that by parallel both L. diffusus and L. sylvestris are smaller and lighter at the northern extreme of its range. Since the dingy diffusus populations occur where the ranges of diffusus and sylvestris overlap, another interpretation is conceivable. This may be an example of overlap of two related species (Blair 1955) which is described as character displacement (Brown and Wilson 1956). L. sylvestris in New England is fairly common in the appropriate habitats in the North but becomes extremely scarce in Connecticut and only a few specimens were collected in the northwestern high- lands at Canaan and Norfolk, Connecticut. According to Parsh- ley (1917) it reaches the New England transition zone from the north. Barber (1921) noted the species as very uncommon in New Jersey, and Slater found records of sylvestris only from the north- eastern corner of Illinois (1952). It is therefore most interesting that Froeschner (1944) records it from Missouri and refers to it as a u. . . so-called northern species (which) has been taken as far 117 ENTOMOLOGICA AMERICANA south as the southern border, where it occurs along with many of the more truly (sic) southern species.” Could this possibly be a disjunct Ozarkian population which has adapted to a warmer habitat? Or, may it represent another species? Environment In New England sylvestris is found limited to cool, mesic, and semishaded habitats along north exposed ecotones between meadows or bogs and beech-maple-birch or spruce-fir woodlands. It is always found in the field aspect side of the ecotone, never in the closed canopy forest. This field formation is one to two feet in height, of medium density and is characteristically dominated by forest type mesophytes as Festuca obtusa Bicher, Carex spp., Aster acuminatus Michx., A. lateriflorus (L.), Solidago caesia L., and Dryopteris spinulosa (Muell.). The ground biotope usually has only a thin litter layer on the mesic soil (moisture 5-7). A few specimens were swept from composites as in L. cliff usus. There is a clear ecological separation between diffusus and sylvestris. On the exposed drier portion of an open slope, only L. diffusus is collected. But as the more mesic ecotone is approached, diffusus drops out and the sparse population of sylvestris appears. The same phenomenon was noted by Slater (in litt.) at the Moosehorn National Wildlife Refuge, Maine and at Lake George, New York. The close contiguity of these populations adds more credence to the species difference accentuation hypothesis mentioned previously. L. sylvestris was found in several sphagnum-black spruce-larch bogs in Maine and in Myrica gale L., at the margin of black spruce, at Norfolk, Connecticut. The other Connecticut collection was in an alder swamp at Canaan, Connecticut. These bog-type ground habitats were, of course, nearly hygric with moisture levels at 1-2. The abundance here was 1-2 per square meter, and this was the only rhyparochromine found in these habitats. Uhler (1875) described it from wild grassy spots adjacent to Sphagnum swamps and in high mountains in North Carolina. These North Carolina records, however, may refer to L. depict us. The extensive European literature indicates a very similar habitat type. Stys (1960) collected it in birch litter and sphagnum in a coniferous forest in Czechoslovakia. Sahlberg (1920) in Sweden and Stichel (1925) in north Germany collected it on the ground in coniferous woods. Pfaler (1936) found it regularly on the margin of spruce woods in Finland. Hedicke (1942) records it in coniferous woods in high alpine areas in the Alps. Wagner (1958) swept it from alpine fields in the Pyrenees. Krogerus (1932, 1960) however describes it in Finland as an ubiquitous spe- 118 VOLUME XLIII cies found in Calluna healthlands, Festuca-Elymus open associa- tions, spruce and pine woodland margins, and birch-alder associa- tions. It appears from Krogerus’ data, that L. sylvestris has a wider ecological amplitude in high latitudes in Europe than at lower latitudes there and in New England. This is perhaps to be expected, and the ecological range of the species in northern parts of North America may be similarly broader. However, it remains possible that in Europe, in the absence of competition with other Ligyrocoris species, L. sylvestris may assume a wider ecological range, much as have many introduced species when released from competition with congeners (Elton 1958, Ross 1962). General Biology As all the specimens available from New England are brach- ypterous, the macropters and submacropters must be rare which would place an important limitation on dispersal in the discontinu- ous mosaic of favorable habitats in the southern portion of its range. It is therefore significant that in Europe, according to Reuter (1875), long winged forms are found southward, short winged forms northward, which may correlate with a discontinuum of favorable habitats in the southern part of its European range. At any rate, many of the habitats, especially bog margins, would be long lasting habitats. The behavior and movements of this species are similar to cle- pictus. The darker and more sharply contrasting hemelytral col- oration of sylvestris forms a disruptive type coloration which con- forms to the mosaic of dark substrate and light leaves in its mesic ecotone habitat. In the laboratory immediately on introduction sylvestris feeds on the seeds of Aster spp., Solidago caesia L., Betula populifolia Marsh, B. alba L., and Tsuga canadensis L., all from its habitat, but not on Festuca obtusa Bichler. It oviposits when feeding on these seeds. It appears especially attracted to sunflower seeds. It has been recorded from a number of plants, especially in Europe, but it is not at all certain that these records represent feeding: goldenrod ( Solidago sp.) in Indiana (Blatchley 1926), Ledum palustre and Betula nana (Sahlberg 1920, Stichel 1925), and Myrica gale (Stichel 1926). Mating behavior and seed defense behavior is similar to L. de- pictus, but less vigorous in execution. While the males will mate with already fertilized young females, the older actively ovipositing females would not permit copulation. Life History Jensen-Haarup (1912) cited L. sylvestris as over-wintering as 119 ENTOMOLOGICA AMERICANA an adult. Pfaler (1936) refuted this and found that sylvestris overwintered as an egg and had a single generation in Finland. The nymphs are present from June 15 to July 15. Immature adults appear in early July, and oviposit from late July to the end of August. Prohaska (1923) found both adults and nymphs in equal numbers on July 18 and 25 in moist meadows in Austria. In New England a nearly identical pattern to Pfaler ’s was obtained at Laconia, New Hampshire. Very few last instar nymphs (4%) were found with the adults in late July, and oviposition in the laboratory continued through August. Slater (in litt.) found fourth and fifth instars as late as August 18 in Maine. In light of this cycle it is most interesting that Strawinski (1960) reported sylvestris in Bulgaria as overwintering as an adult. Moreover in the population of L. sylvestris in Missouri, Froeschner (1944) reported adults from as early as May 24 to as late as October 19. Either this population is not L. sylvestris as mentioned earlier, or it may represent an adaptation of a southern disjunct population to a longer, warmer season. The onset of diapause occurs in early anatrepsis when the egg is still white. When left in warmth, development occurs slowly and some eggs during late September and October become pink colored at late anatrepsis, and a few later slowly reach katatrepsis. These eggs uniformly died without hatching. Some viable eggs never developed. Cold exposure (20° and 34°F.) for several months did not permit hatching, although the eggs developed to katatreptic position. Only a long cold exposure (September 12 to April 30) yielded any nymphs (28% hatched). Only data on oviposition is available. The fecundity in the laboratory is high: the mean, 272 eggs; the range, 161-475 eggs. Both moist and dry substrates are readily oviposited into, and no preference is discerned except for a loose substrate. The eggs are beset with hairs and cling to the substrate. Ligyrocoris caricis Sweet This is the only new species discovered during the present study on the New England Rhyparochrominae (Sweet 1963). This slender small species was found only at Pink’s Ravine Pond near Storrs, Connecticut, and was collected on Mt. Desert Island, Maine (Great Health) by F. B. Shaw. It is very unlikely that its distribution is this limited and it should be found in suitable habi- tats at other locations at least in the northeastern United States. 120 VOLUME XLIII Environment As mentioned earlier (Sweet 1963), the habitat of this species is entirely different from its close relatives. It is found among the outcropping stones and stool-tussocks of Carex stricta Lam. at the margin of Pink’s Ravine Pond, an artificial but dystrophic pond with dark humus-colored water. This is a rather limited area only about 50 yards long. The plant community is the typical transition from a wet red maple forest through shrubs to the emergent pond vegetation. Shrubs of Cletha alnifolia L., Rosa carolinianum L., and Cephalanthus occidentalis L ., form the shrub layer. In the open area, other shore herbs are Scirpus sp., Hyperi- cum sp., Drosera rotundifolia L., and emergent further out, Alisma sp., Sparganium sp., and Sagittaria sp. Around the bases of the Carex stools and rocks are a few scattered patches of the bog moss Sphagnum sp. This association approaches the Caricetum strictae of Conrad (1935). L. car ids is collected chiefly in the open area on the substratum , on the Carex stools, at the edges of rock out- crops, and on fallen Carex litter at the water’s edge. During early June at all times of the day adults and last instar nymphs are frequently found feeding on the Carex heads on the seeds. By late June and early July the seeds have largely fallen and the insects are then restricted to the ground level and rarely could be swept. At this same time, the adults are occasionally found in the mixture of Sphagnum and leaf litter beneath the shrubs. The biotope of caricis then varies from open exposures to semi- shade in Carex clumps or in the shrub margin. The moisture level is, of course, saturated to very wet (9-10) and the temperatures moderate 62° to 76°F. in late June. The population level of L. caricis remained approximately the same from 1960 to 1962, despite the draining of the entire artificial pond in August of 1961 which left the area dry for several weeks. The population as sampled reaches a peak in mid- June and declines steadily until late July when none could be found. The abundance at its peak was 5-6 per square meter of Carex tussock, or 1-2 per square meter of straight area and declined to less than 1 per square meter of tussock by July 10. It was found with Scolopostethus thomsoni and Pachybrachius albocinctus Barb, (see Competition Discussion). As mentioned under L. depictus, Torre-Bueno’s (1925) collec- tion of L. depictus from sedges in New York may very well represent this species. Moreover, the extent of the hemelytral fascia in L. caricis is variable and some individuals may be keyed out as L. depictus in Barber’s 1921 key (Sweet 1963). The association of Scolopostethus “ atlanticus” (=“ thomsoni”) with Torre-Bueno’s 121 ENTOMOLOGICA AMERICANA depictus is also similar to the present association. General Biology It is clear that this species must have considerable difficulty in colonizing* new habitats as all of the specimens collected or seen were brachypterous or submacropterous, and incapable of flight. When collected or disturbed on Carex heads the insects either dropped or ran down the stalks to the ground layer, and at no time attempted flight. Collecting at night by lights at this location drew Pachy- brachius albocinctus, but not L. caricis. Yet Pink’s Ravine pond is definitely an artificial one and is of no great age. This is an extremely rapid moving insect and, in its speed, very similar to L. depictus, a much larger species but its closest relative (Sweet 1963). In the field it appears that the narrow form of this species (much narrower than other Ligyrocoris) , along with the pale mar- gin of the hemelytra which increases the apparent slimness, may be a camouflaging adaptation to the narrow leaves of the Carex as in many other grass and sedge feeding insects. Otherwise the colora- tion is a procryptic fuscous and black. The nymphs are clearly ant mimics with a broad white band across their fuscous and white spotted abdomens. This mimicry is especially apparent in their rapid jerky movements when dis- turbed. While these insects pass their entire nymphal life history on Carex seeds, they feed quite readily on sunflower seeds. Indeed such is the low fertility of the Carex seeds brought into the labora- tory that it is safer to rear the species largely on sunflower seeds, although they can be reared readily on the sedge seeds. Mating behavior is, in general, similar to that of the other species. The male courts the female with a jerky dance which is probably a stridulation dance as discussed under diffusus. In the mating patterns, there are some significant differences. First the male is much less excitable than in the other species and requires a number of contacts with a female before any sexual response is elicited. Second, the female takes a much more active role in the courtship pattern. When contacting a male, and before the male shows any recognition, the female in each case advances shortly on the male, her antennae wagging rapidly. This prior advance was not seen in the other species. After three or four such advances the male, which has moved but little, suddenly begins the courtship dance whereon the receptive female quiets a little and usually after two or three attempts which the female repels in the fashion of diffusus the male will accomplish copulation. As in diffusus the 122 VOLUME XLIII male jerks continuously and vibrates the antennae rapidly when initiating copulation. One copulation lasted 2 hours 15 minutes. As in L. diffusus, copulation may occur repeatedly in the young female, but an old female (in this case over 20 days old) would not mate. Males caged together would occasionally advance on one another. A female which lost its antennae was unable to recognize the males and never copulated as in L. diffusus. It is rather interesting that although the males of the other species of Ligyrocoris reacted with the different female species, they showed no response to L. caricis females and nor did L. caricis males react to females of other species. However, the introduction of the male of another species with L. caricis stimulated a very obvious avoidance reaction, in which the female raced to and fro across the dish despite the quietude of the alien male. As in diffusus the wagging antennae indicates “annoyance” during feeding when intruded on by another individual. Often the insects would run about with the small Carex seeds suspended on the end of the labium when it was folded under the body. Life Cycle As in L. sylvestris and L. depict us, there is one generation a year with a long diapause as an egg. Partly due to the high water table in May, and perhaps because of their small size, I could not find the early instars in the field. The observed phenology is as in Table 12. TABLE 12 Phenology of Ligyrocoris caricis Date Instar 3 Instar 4 Instar May 29 25% 75% June 8 67% 33% June 12 — 100% June 25 84% June 28 44% July 11 Adult 16% 56% 100% Diapause intervenes in early anatrepsis when the egg is white. No further development occurs in the laboratory indicating a strong diapause. I was unable to break diapause with cold expo- sures of two to four months at 4°C. As the species oviposits shortly after becoming adult, egg diapause in the field must last as in L. depict us through the summer as well as through the winter. Such 123 ENTOMOLOGICA AMERICANA a seasonal cycle may not only be adapted to the host sedge plant, but may also serve to allow a late summer drought period. Such a rapid life cycle may also adapt this species to a much shorter summer season. It may be conjectured that this species will be found much farther north and indeed some of the records of L. diffusus as Lindberghs (1958) from Newfoundland may represent this species. Since only the late instar nymphs were found the stadia are not available except for the last instar. It averaged 6.8 days, and ranged from 6-7 days. The male and fertilized female longevities are similar: mean, 33.0 days, range 31-36 days. A virgin female lived longer, 48 days. The precopulatory and preoviposition periods are nearly the same, 5.0 and 5.5 days respectively, as oviposition occurs shortly after fertilization. The average fecundity of four females is 133 eggs (range, 125-151 eggs). The oviposition period is from mid or late June to late July. The eggs are laid singly at the rate of 4.8 a day. The rate could be varied by not providing an appropri- ate substrate to oviposit into. The female would retain its eggs for a few days and become quite swollen, and then would lay a large complement of 10-11 eggs. The eggs are slender and smooth, not beset with knobbed hairs as in L. diffusus and do not stick to objects. They are laid prefer- ably on loose wet methyl cellulose, wet soft Carex debris, and not on dry substrates as in L. diffusus and L. depictus. To Be Continued 124 PRINTED BY BUSINESS PRESS, INCORPORATED Americana A Journal of Entomology. VOLUME XLIV (NEW SERIES) for 1964 PUBLICATION COMMITTEE JAMES A. SLATER, EDITOR GEORGE S. TULLOCH JOHN HANSON PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY 1964 ENTOMOLOGICA AMERICANA Yol. XLIV (N.S.) 1964 CONTENTS The Biology and Ecology of the Rhyparochrominae of New England (Heteroptera : Lygaeidae) Part II . . . Merrill H. Sweet Pages 1-201 VOL. XLIV (NEW SERIES) AmeriqAna A Journal of Entomology. PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY PUBLICATION COMMITTEE JAMES A. SLATER, EDITOR GEORGE S. TULLOCH JOHN HANSON Published for the Society by the Business Press Inc. N. Queen St. and McG-ovem Ave., Lancaster, Pa. Subscription $6.00 Per Year Date of Issue December 29, 1964 AMERICANA VOLUME XLIV THE BIOLOGY AND ECOLOGY OF THE RHYPAROCHROMINAE OF NEW ENGLAND (HETEROPTERA: LYGAEIDAE). PART II1 By Merrill Henry Sweet2 Table of Contents Myodochini Plinthisini Antillocorini Lethaeini Ozophorini Stygnocorini Drymini Rhyparochromini Megalonotini Gonianotini Bibliography Index Page ' 59 62 67 75 83 95 124 133 144 165 194 In Part I the ecology of the Rhyparochrominae was discussed in general and six of the species were taken up. In the present and final part the remaining rhyparochromine species of New Eng- land are considered. Attention should be called to the recent work of Eyles (1963a, b, c, d) upon the biology of nine British rhyparochromine species which came too late to be included in Part I. 1 A dissertation submitted in partial fulfillment of the require- ments for the degree of Doctor of Philosophy at the University of Connecticut. 2 Department of Biology, Texas A & M University, College Sta- tion ; formerly at University of Connecticut, Storrs. 1 iNSTlTUHON A 0 ENTOMOLOGICA AMERICANA Zeridoneus costalis (V. D.) Zeridoneus differs from Ligyrocoris in lacking an abdominal stridnlatory area. At present the genus contains only two species. However as shown by Ashlock and Lattin (1963), Z. costalis, along with such Ligyrocoris species as L. latimarginatus Barb., L. litigi- osus Barb., L. obscurus Barber (and perhaps a few others) forms a rather compact group of species that are similar in general struc- ture and type of aedeagus. Under high magnification the epi- dermal cells of the normal stridnlatory area in Z. costalis are clearly aligned in raised parallel rows essentially similar to the much more distinct condition in Ligyrocoris. Even with the potential inclusion of these other species, Zeri- doneus has a distinctly Nearctic distribution. The other known species', Z . knulli Barb., was described from western Texas (Barber 1948a). Of the species of Ligyrocoris related to Zeridoneus, L. obscurus is known from the eastern United States in a narrow band from Maryland to Kansas ; latimarginatus Barber from the northwest Pacific coast of United States ; and litigiosus from Florida to Arizona and Mexico (Barber 1921). Zeridoneus costalis itself is a north temperate species found in the eastern half of northern North America from Alberta and Manitoba to Quebec and south to Iowa, northern Illinois, southern New England (Slater, Catalogue), and also in the highlands of North Carolina (Brimley 1944). It may be significant that the ranges of the above species, for the most part, are allopatric. Froeschner (1944) recorded a specimen of Zeridoneus from Mis- souri, but stated that Barber was uncertain whether it was Zeri- doneus costalis as it differed considerably in color. This specimen may represent a relict Ozarkian population or a separate species. While Z. costalis is generally considered rare (Blatchley 1926, Slater 1952), in Connecticut I found it to be of average abundance compared to other rhyparochromines. Environment Zeridoneus is found in open field habitats in relatively early succession stages, but the general aspect of these habitats varies considerably. It is found at such ruderal sites as vacant lots and road sides, on flood plain pastures, along the disturbed field side of mesic woodland ecotones, in tall herb fields, and even in some small drying marshes. These sites vary in exposure from completely open to semi-shaded. It is clear, however, that Zeridoneus has a mesic (5-6) ground layer biotope preference, and even in the drier open habitats it is found in relatively moister sites than L. diffusus, 2 VOLUME XLIV such as in the margin of litter of a thick grass pile, in the shade of a Panicum clump, or near a riverside. It is never found on dry, well drained slopes such as those dominated by Festuca, Stipa, or Andropogon. The highest populations (20 per square meter) of Zeridoneus were found on several newly abandoned gardens in the semi-shade of a woodland margin, among the pioneering plants Panicum spp., Agropyron repens (L.), Rumex obtusifolium L., Chenopodium album L. and other forbs. At more ruderal sites, Zeridoneus is frequently found in moder- ate abundances (6-8 per square meter) at the base of Panicum sp., Potentilla recta L., Centaurea, Cichorium intybus L., Chrysanthe- mum, leucanthemum L., Daucus carota L., Plantago sp., and other rank plants. While the vegetation cover varies from complete to areas with considerable bare soil, the greater abundances are found in the more sparsely covered disturbed sites, or where the general cover is quite short, under 6 inches, interspersed with larger grass clumps. The soil is sandy or loamy, frequently dark, reflecting the relatively mesic sites. The litter layer is usually sparse, except where once it was artificially piled and provided a habitat for Zeridoneus in a site otherwise dominated by Ligyrocoris diffusus. Although Zeridoneus was not swept from plants in the favorable habitats, it was swept in small numbers at sites where it was' not found on the ground layer. Slater {in litt.) swept it from tall rank herbs near Lake George, New York but also did not find it on the ground. (Only adults were collected by sweeping.) Accord- ing to Blatchley (1926) Barber collected a specimen from high weeds along the bank of a stream in the Adirondacks. Barber (1928b) collected it on a semi-xerie hillside in the Adirondacks and, significantly, described the habitat as adjoining a woodland. Hend- rickson (1930) collected it in a Stipa-spartea- Andropogon scopar- ius association and a Bouteloua curtipendula association in Iowa. While the former Iowa community is a climax association (Hendrickson 1930), in the northern and eastern parts of the range of Zeridoneus the climax is forest. Thus in the eastern part of its range, Zeridoneus is a subclimax species inhabiting open mesic disturbed habitats which are relatively short lived. General Biology Zeridoneus is a completely macropterous species. This corre- lates with its temporary habitat preferences. While there are no recorded light captures, it was frequently observed in the labora- tory that new adults are especially active in the evening after sun- down and fly readily to nearby lights. 3 ENTOMOLOGICA AMERICANA This is another rapid moving long legged myodochine which when disturbed responds by actively running instead of taking refuge in litter. It is usually found on the ground or litter and infrequently in other than loose thin litter. Its dark coloration blends into the more mesic habitats but renders it conspicuous in some of the sandier habitats. The nymphs are darker than are those of Ligyrocoris which they strongly resemble. As in Ligyro- coris, the running nymphs mimic ants. A few tachinid parasites of the genus Catharosia were reared from the second generation of this species. They emerge shortly after the host becomes adult. The late developing instars that were collected as late as September 21 were all parasitized by Catha- rosia. This insect utilizes a wide range of ripe food seeds. In the field adults were occasionally observed on the plants feeding on ripe seeds of Carex lupulina Muhl., Potentilla recta L., Cichorium intybus L., and Chrysanthemum lecucanthemum L. In the labora- tory they feed on these seeds, especially those of Potentilla; and also on the hedge-nettle Stwchys sp., Festuca sp., Budbeckia sp., and sunflower seeds. It displays the strongest response to Budbeckia and Potentilla. It does not appear to recognize as food the seeds of Paspalum sp., Panicum muhlenbergii Nash, Solidago sp., Chen- opodium alba L., or Hypericum sp. It is readily reared on sun- flower seeds from egg to adult. The mortality is lower and the rate of development more rapid when Potentilla seeds are added. Nymphs as young as the second instar can pierce the thick sunflower seed coat and feed on the kernel. The seeds are frequently dragged to more protected sites under methyl cellulose in the laboratory, but no seed defense behavior was observed. Like Perigenes, the male smeared its defecations in a thin line, but no associated odor was perceived. The mating behavior is rather similar to that of Ligyrocoris and may constitute additional evidence of the close relationship be- tween these two genera. After contacting a sexually receptive female, the male may become sexually excited. It responds by moving in short bursts on high stiff legs ; the abdomen shakes back and forth at the rate of 2 or 3 times a second ; and the distal three antennal segments are vibrated rapidly in a horizontal plane and at a right angle to the body and the antennal scape. With this courtship “dance” the male advances on the female frequently touching her with his vibrating antennae. The male often follows a circular path around the female in his attempt to mate with her. Initially, the female, in several trials, gave a quick brief jerk 4 VOLUME XLIV of her legs similar to the male’s shaking and actually advanced toward the male. Only in a very few trials was the male observed to complete mating, in the same fashion as described in Ligyrocoris cliff usus. In 90% of the trials the female decamped. An interest- ing aspect is that in a culture with eleven pairs of these insects, one male was responsive to only one of the females and would dance only when contacting her, and one other male was responsive to one other female. The excitement of a male may spread by contact to other males, causing them to begin “dancing.” When caged alone an unmated male will spontaneously begin to dance despite the absence of a stimulus from the female. This qualifies as a “spontaneous release” of the behavior pattern. When several virgin males are isolated together one male may dance briefly on contact with another male. This release in isolation was not ob- served in Ligyrocoris. Finally, despite the faintness of the stridu- litrum, it is possible that these insects may stridulate by the shaking movements of the abdomen as was postulated also in Ligyrocoris. When placed with other large myodochine species such as Pachybrachius, Cnemodus, and Pseudocnemodus, no reaction was elicited from sexually active males or from the alien females. It is therefore significant that while males of Zeridoneus did not react to females of Perigenes, Ligyrocoris depictus, and L. diffusus, the females of these genera gave a strong avoidance reaction to the presence of the Zeridoneus males. This may further substantiate the apparent close relationship of these genera. Life History While the bivoltine seasonal cycle with an egg diapause is similar to that of Ligyrocoris , the precise phenology is different. The phenology at Storrs, Connecticut is shown on Table 13. Adults were found en copulo in the field September 9. Oviposit- ing females of the first generation were found after July 11 and the second generation after September 9. The reproductive pause from mid- June to early July cannot be considered merely as an immature period, for in the second generation oviposition occurs shortly after this generation becomes adult. The result is that the second generation occurs largely in August, which is about a month later than in Ligyrocoris. This data suggests that reproductive activity occurs during shortening photoperiods and during long photoperiods non-diapause eggs are laid while in short photoperiods diapause eggs are laid. While no experiments were performed, this hypothesis explains the occurrence of 26% non-diapause eggs in two second generation females which were forced to early maturity under the prevailing ENTOMOLOGICA AMERICANA photoperiod in the laboratory by August 18. This aspect is dis- cussed later. Autumn eggs entered diapause in early anatrepsis. The dia- pausing eggs never broke diapause in warmth and died in mid- winter. Short cold exposures of 1 to 6 weeks and three months did not terminate diapause. Diapause was broken by cold ex- posures (35°F.) of 192 and 212 days (six to seven months). Unlike other species, the majority of the diapause-released eggs developed nearly in unison and the hatching was only weakly staggered, with a few of the eggs hatching later. TABLE 13 Phenology of Zeridoneus Cost alls Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 27 73% 20% 7% June 10 20% 20% 45% 15% June 18 18% 32% 50% July 5 9% 91% July 12 100% July 27 72% 28% Aug. 7 25% 40% 15% 20% Aug. 18 2% 8% 22% 35% 23% 10% Sept. 3 15% 85% Sept. 20 10%* 90% parasitized It would seem that the later eclosion of Zeridoneus eggs in the field than those of Ligyrocoris of almost contiguous habitats stems from the earlier onset of diapause and consequent longer spring development. Moreover Zeridoneus eggs are laid in cooler more shaded biotopes than Ligyrocoris which would become warmer later in spring. The rapid field development of the stadia of laboratory reared specimens is reflected in Table 14. After release from diapause the overwintered eggs develop in 14 to 17 days. Adults of the spring generation persist until mid- August and the average longevity is 47 days (range 35-57 days) in the laboratory. In the autumnal or second generation the adults, presumably since they become reproductive earlier, live for a shorter period in the laboratory (mean, 34.5; range, 28-39 days). The 6 VOLUME XLIV insects are relatively cold hardy, and persist in the field until as late as November 6. This field longevity is distinctly longer than that found in laboratory specimens, undoubtedly because of the cooler field temperatures. Cold exposure of the adult females for a week does not affect the egg diapause condition. TABLE 14 Stadia of Zeridoneus costalis Egg* Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 11.0 4.2 4.9 5.0 5.6 6.7 36.5 (11-12) (4-5) (4-6) (4-6) (5-7) (5-10) (32-45) * Generation 1 Virgin females have a much longer adult life span in probable correlation with their low reproduction activity (mean 57 days, range 43-71 days) . As noted under the diapause discussion, the preoviposition pe- riods of the spring and summer generation are quite different. The first generation adults do not become sexually active until the first week in July, and the precopulatory and preoviposition periods vary accordingly and average 20 days. In the second generation the precopulatory period is as short as 3 days, the preovipositional period 4 days in the field adults which have become mature from August 29 on. However, adults reared to maturity under prevail- ing daylight earlier than mid-August likewise did not become mature until the first week of September. This oviposition pattern may be readily explained by a photoperiodicity mechanism with the critical points after the summer solstice (15 hours) and in late August (13.5 hours). The fall and spring generations have similar laboratory fecundi- ties which vary from 99 to 231 eggs (mean 164). The eggs are laid at the rate of 5. 5-8. 7 eggs a day (mean 6.4). Sexually isolated females laid none or very few eggs (0-20, mean 7.3). One female, however, laid 50 eggs at the rate of 2.2 eggs a day. Like Ligyrocoris diffusus, the rounded cylindrical eggs are covered with a layer of short fine hairs and the eggs similarly cling to debris. They are laid singly into litter crevices, into fallen ripe seed heads and sepals of Potentilla flowers (from which they could not be shaken), in hollow stems, and into dry loose soil or litter. The female spends considerable time probing at a site before ENTOMOLOGICA AMERICANA ovipositing. This is apparently related to a definite spatial con- tact requirement for oviposition. A gradient of sand grain sizes was formed by shaking sand in a petri dish. On such a gradient, the female selectively laid her eggs in a zone of sand particles which were similar in size (and in interspace) to Zeridoneus eggs. Perigenes constrictus (Say) Only the more recent literature can be accurately associated with Perigenes constrictus, as this large dark species was thoroughly confused in the literature with Zeridoneus costalis (Say) and Ligyrocoris abdominalis Guerin (Heidemann 1903, Van Duzee 1909) which it superficially resembles. As presently defined Perigenes Dist. contains two other species ; P. dispositus, the type, from Guatemala, and P. similis from the southeastern United States. P. constrictus is a northern species and the southern records are undoubtedly referable to L. abdominalis Guer. (Barber 1914b, Van Duzee 1914). It is recorded from Quebec and South Dakota south to North Carolina and Texas, and also from Colorado, California, and Alaska (Slater, Catalogue). The great majority of the records are from the central and eastern United States. In the Midwest its range apparently broadly over- laps that of the more southern P. similis. While Slater (1952) re- ported it as relatively common in Illinois and Iowa, Proeschner (1944) found it uncommon in Missouri and similis to be the much more abundant species. Blatchley (1895) considered it rare in Indiana. In New England, I collected it only in Connecticut, and only a few temporary populations were found. It appears to be more common along the coast. It cannot then be considered a par- ticularly common species, but its large size and ruderal habitat preference would cause it to be more frequently collected than in- dicated by its abundance after precise collecting. Environment P. constrictus is typically collected in exposed, level ruderal habitats in vacant lots, roadsides, and newly fallow fields. In general it is found in a slightly later serai stage than L. diffusus, and in a community of rank forbs and grasses such as Angropyron repens L., Chenopodium album L., Erigeron canadensis L., Ambrosia artemsiifolia L. The herb layer is usually two to three feet in depth. This is a distinctly more mesic habitat type than that of L. dif- fusus. The ground biotope is shaded from direct sunlight by the 8 VOLUME XLIV field layer, and little litter is present on the ground. The soil is usually a loam or sandy loam, moderately dry (3-5), and the soil temperature moderate. Only once was the species swept from low ruderal forbs. An exceptional occurrence was the presence of nymphs in a wash habitat at Mansfield, Connecticut along with a number of other rhyparochromines. However in this favorable seed-filled habitat, Perigenes was found in a zone of moisture and tempera- ture which approximated the more normal habitats. The abundance of this species varied from one or two to as many as 20 per square meter at one favorable habitat in a vacant lot at Noank, Connecticut which was dominated by Agropyron repens L. At none of the sites in Connecticut was the species collected more than two years in succession, and this only with fall ovipositing adults and the resultant spring generation. The available habitat notes largely confirm this habitat choice : Hussey (1922) collected it on Indiana sand dunes area on ragweed, and found another on sand by a road. Torre-Bueno (1910) col- lected it from short grasses in New York, and Blatchley (1926) records sweeping it from herbage along streams and on mulleins in June and July, in sphagnum moss in August, and under logs in November. Froeschner (1944) collected it among weeds in a low marshy field. Dowdy (1947) collected it from the herb layer in a oak-hickory forest margin. General Biology In correlation with its preference for temporary habitats, Peri- genes is entirely macropterous. While it has not been previously recorded from lights, I have collected three specimens during early July at lights at Noank, Connecticut. Perigenes similis has been collected at lights in Missouri by Froeschner (1944), and in Florida by Hussey (1952). Glick and Noble (1961) collected P. constrictus (?) by airplane at 200 feet in Louisiana. This large species is procryptically colored, its dark fuscous coloration blending into the dark loam substrate of many of its habitats. Occasionally, however, it is found on lighter sandy loam on which it is very conspicuous, especially when moving. The later instar nymphs are similarly colored a dark fuscous, the ab- domen patterned with minute pale spots. This is a relatively slow moving heavy-bodied myodochine, and appears to rely on its colora- tion and concealment in its rank plant habitat. It shows little tendency to take cover under litter or debris. A Catkarosia tachinid fly parasitizes Perigenes. All parasites were reared from the second generation Perigenes, and emerged 9 ENTOMOLOGICA AMERICANA shortly after the host became adult. This early emergence cor- relates with the egg overwintering diapause condition of this species. The rate of parasitism is low, under 10%. No predators were as- certained. While this species is readily reared on sunflower seeds from any field-collected nymphs, only a few first instar nymphs, despite active feeding, develop past the first or second instar. Those that did were reared through the life cycle. This species feeds on the seeds of Chrysanthemum leucanthemum L., Ambrosia sp., a grass ( Agropyron repens ), Budbeckia seeds, and probably feeds on many others. No expression of seed territorially was elicited from Perigenes. Unfortunately the mating behavior was not observed. The males, however, possess an unique peculiar odor which is associated with their defecations. Moreover, when defecating, the male, as the drop is released, moves forward a few steps and smears a long streak of the odorous material. Presumably the odor and the smearing behavior play a part in this species’s mating behavior. This odor is very persistent, and several dishes still retained the odor after four years. Life History While Blatchley (1895, 1926) considered this species to over- winter as an adult in Indiana, it actually overwinters as an egg, but the adults are cold resistant and persist until late in autumn. Blatchley recorded specimens “under logs” in November and on December 10. I have taken actively ovipositing females as late as November 16. There are two generations a year but the non-dia- pause spring generation was indeed scarce. Only in the second generation was Perigenes found in any numbers. The available phenology at Storrs, Connecticut is as given in Table 15. While this life cycle parallels that of Ligyrocoris diffusus, the second generation occurs later in the summer. Probably a similar photoperiodicity is involved. The onset of diapause occurs in early anatrepsis. Unlike dif- fusus cold exposure does not affect the female ’s capacity to lay dia- pause eggs. The strength of the diapause condition is quite vari- able. A few eggs develop within a month in warmth after being laid and a few others continue to come out of diapause all through the winter. About two-thirds of the eggs, however, never develop in warmth. When placed in cold (35°F.) for five months, the eggs hatch readily but again at staggered intervals, which suggests a protective spreading out of the spring eclosion. Once diapause 10 VOLUME XLIV terminates egg development occurs rapidly. In thirteen days after removal from cold, first instars hatched. TABLE 15 Phenology of Perigenes constrictus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 18 33% 67% May 26 June 9 50% 50% 50% 50% June 20 100% July 12 100% July 15 5% 15% 35% 20% 25% Aug. 20 10% 16% 30% 44% Aug. 28 12% 44% 44% Sept. 15 100% The stadia in Table 16 are based on rather small samples and no averages are given. Spring adults lived in the laboratory until late July and early August and averaged 45 days (34-56 days). The fall generation lives longer, averaging 76 days (60-82 days). TABLE 16 Stadia of Perigenes constrictus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 12 ( ?) 6 6-7 6 6-7 7-8 45 The reproductive capacity is very great, averaging 268 eggs (184—330) which are laid at a rate of 7 to 8.5 eggs a day. The eggs are laid singly when an appropriate substrate is available or in batches of 8-10 when a substrate is withheld. The eggs are prefer- ably laid in dry substrates, and like Ligyrocoris the eggs are ovi- posited deeply into the substrate of methyl cellulose, cotton, or sand. These eggs do not cling to the litter or sand. Even into firm sand the female was able to oviposite. Two virgin females laid no eggs. 11 ENTOMOLOGICA AMERICANA Sphaerobius insignis Uhler Among the rhyparochromines of New England, Sphaerobius is the most remarkable ant mimic. Three species are presently placed under Sphaerobius, but Slater (in lift.) informs me that the type, S. gracilis Uhler from St. Vincent Island does not appear to be con- generic with S. insignis. It is unfortunate then that Van Duzee (1917) fixed gracilis as the type, not the well known insignis. The third species, quadristriatus Barber, was placed close to insignis. Bearing this situation in mind, we may consider “Sphaerobius” in the sense of insignis, to be a purely Nearctic genus. The two species are quite different in distribution. S. quadri- striatus is known at present only from New Jersey. Other records refer to insignis. S. insignis has a northern or boreal distribution across North America from Newfoundland to British Columbia, and south to northern New York, Connecticut, Iowa, and in the western mountains to Colorado (Slater, Catalogue). In New Eng- land, insignis is found only as far south as northwestern Connecti- cut, and moreover, is scarce south of the northern tier of New England states. Environment Like Cnemodus this species is especially common on open dry barren sites among sparse clumps of the grass Andropogon scoparius Michx. It is frequently found on roadsides and railroad right-of- ways which are dominated by the above association. It is not found at all in ruderal sites dominated by new field forbs. Truly remark- able abundance levels (up to 60 per square meter) were found in an andropogonetum association along a railroad right of way in the southern Adirondacks near Warrensburg, New York. The soil on such open xeric sites is overdrained and very dry (1-2), and during the day becomes very hot (up to 50°C.). It was observed at several sites that in the early morning and late afternoon, the insects were found in the open bare areas, and as the day progressed became restricted to the litter of the Andropogon clumps. When the ground cover becomes completely filled in by other grasses, the abundance of the insect is much lower (1-5 per square meter). At a site in Canaan, Connecticut a definite gradient in abundance was traced from a filled-in margin to a bare soil area. It is clear that these habitats are essentially temporary. The poor overdrained gravelly soil, however allows only a very slow succes- sion to proceed. Hendrickson (1930) found Sphaerobius to be very abundant in a climax prairie association of Andropogon and Stipa. It may be 12 VOLUME XLIV that the eastern populations represent an influx of a western popula- tion, but Lindberg (1958) reports that it was found in Newfound- land on open swampy ground with Car ex and Sphagnum, near the seashore. It is evident that much more field work needs to he done over the entire range. General Biology The populations studied consisted of a majority of brachypterous (flightless) adults with a minority (ca. 20%) of macropters. No relation appears to exist, however, between population density and macroptery. It is evident from the natural situation that the pro- duction of macropters is sufficient to maintain Sphaerobius in the scattered mosaic of suitable sites. It would be interesting to see whether north midwestern populations in presumably widespread permanent habitats exhibit a lower frequency of macroptery. Sphaerobius exhibits a most interesting color polymorphism which is unique among the rhyparochromines studied. In large populations about 50% consisted of dark individuals, most of the remainder of light tan individuals, and a few intermediates. This is not a generation or developmental change in color as in many mirids (Kullenberg 1944), for the final color pattern develops shortly after molting. There is no apparent relation of color to sex or brachyptery. The ant mimicry effect is gained by transverse white bands on the corium, a narrow swollen thorax and a somewhat myrmeform head. The nymphs are yellow-brown in color with white markings on the abdomen, and the abdomen is narrowed near the junction with the thorax. The mimicry effect is greatly emphasized by an irregular ant-like gait when running. The effect of this combina- tion in the field is of a population of very large black, red, yellow- red, and smaller yellow ants (the earlier instars). The only ant found anywhere near the adult range was the much smaller Formica rufa, and the lygaeids were frequently as abundant or much more abundant than the ants at a given site. The ant mimicry effect was noted by Barber (1922) and Parshley (1921) who also noted an association with ants. Such an association, however, is inevitable in the dry habitats preferred by Sphaerobius. Sphaerobius is parasitized by Catharosia tachinid flies. Para- sitism appears to occur during the late instars and emergence occurs from the adults. There is no diapause, the fly emerging a short time after the insect becomes adult, which correlates with the egg diapause in Sphaerobius. Sphaerobius feeds on the seeds of the grasses Andropogon scop- arius, Panicum sp. and Paspalum sp. It feeds readily in the labora- 13 ENTOMOLOGICA AMERICANA tory on sunflower seeds on which it can be reared very easily. Seeds are defended by quick rushes with flailing antennae, and annoyance is shown by rapid antennal wagging. The courtship pattern in Sphaerobius involves a rapid shaking jerking motion by the male when approaching the female much as in Ligyrocoris and Zeridoneus. It is therefore interesting that Sphaerobius, like Ligyrocoris also has a faint striated area (or stridulitrum ) on the sides of abdominal segments three to four (Ashlock and Lattin 1963). The remaining mating process is much as in Ligyrocoris. Life Cycle Sphaerobius insignis has a bivoltine seasonal cycle with a facultative egg diapause. Only a general description can be given here as much of the quantitative data on this species was lost. The phenology at Canaan, Connecticut is as follows : Early instar nymphs were found as early as mid-May, but third and fourth in- stars were present as late as mid- June, apparently from a staggered eclosion. The first generation adults appeared in mid- June and oviposition began in late June. The late maturing nymphs became adult by early July. In late July at Warrensberg, New York, second generation nymphs of all instars were present. As in Ligyrocoris depictus there is no mid-summer reproductive pause, and the oviposition of diapause eggs begins in August after a brief precopulatory period. Fourth and fifth instars were found as late as mid-August. Diapause occurs during early anatrepsis, and is a strong one. The diapause is probably under photoperiodic control since several second generation cultures laid non-diapause eggs under long photoperiods. A few eggs broke diapause after several months but nearly all required cold exposure of several months to terminate the diapause state. This bivoltine seasonal cycle is unusual for a rhyparochromine with a northern distribution. Very probably this is made possible by the generally hot exposed habitats selected by Sphaerobius. Sphaerobius has a very high reproductive capacity, and in the laboratory the fecundity varies from 180 to 350 eggs per female. Sexually isolated females laid no eggs. The eggs are curved-cylin- drical with rounded ends and heavily and densely beset with hairs. The eggs stick to objects and become densely covered with sand grains and debris when laid in the ground. An egg is laid after a careful examination of the site. The eggs are laid singly in loose fine litter into soft ground, and into methyl cellulose and cotton. The eggs are not laid on wet substrates or bare substrates. 14 VOLUME XLIV Sphaerobius quadristriatus Barber I have not been able to find this most interesting and apparently very rare species which was described from Lakehurst, New Jersey by Barber in 1911. From the type locality and a history similar to Malezonotus fuscosus it is very likely that S. quadristriatus is one of the endemic pine barren forms (see Malezonotus) . The three known males were all brachypterons, and collected on July 4 and September 7, 1909. Pachybrachius Hahn This large and cosmopolitan genus is known from all zoogeo- graphical regions, especially from tropical areas. Presently 71 species are placed in this genus, but many do not belong here, for the genus was a convenient one for placing new myodochine species, especially under the name Pamera. Scudder (1962), for example, has moved many species into several other genera. In New England three species are recorded, basalis, albocinctus , and bilobatus, each of which represents a different northern ex- tension in New England. Only the former two were found in the course of this study. Eventually two others might be found, the European P. fracticollis and P. luridus which may have been suc- cessfully introduced into Canada as both species are adapted to boreal swamps and sphagnum bogs (Krogerus 1960, Southwood and Leston 1959, et aL), a niche which is apparently unfilled in Canada. One other species, occult us Barber, occurs in the western United States, in Idaho, Montana, and Colorado. Ashlock (in litt.) says the genus Pachybrachius does not occur in California. Pachybrachius basalis (Dallas) This species is one of the most abundant rhyparochromines in New England. It is distributed in eastern North America north to Quebec and Minnesota, west to Iowa and New Mexico, and south to Texas and south Florida (Slater, Catalogue). Uhler’s (1894) record of it from Grenada in the West Indies is probably incorrect in the light of the otherwise known distribution. It is found more abundantly in the southern parts of New England than farther north. 15 ENTOMOLOGICA AMERICANA Environment P. basalis also appears to have perhaps the largest ecological range among the New England rhyparochromines, which consider- ably magnifies its apparent abundance. While it is occasionally found hibernating in late autumn with Myodocha and Heraeus in light woodland sites, it is definitely an insect of field habitats. It is found from relatively moist mesic streamside meadows with rank plants to fairly dry upland old fields. Its greatest abundances are reached on rank hillsides and roadsides covered with the large coarse panic grass, Panicum spp. It is commonlv collected at ruderal sites and along garden margins. Uhler (1876) recorded it from wheat and grass fields in spring and summer, and Blatchley (1895) found it rather common on the borders of cultivated fields. This wide ecological amplitude is further shown by the collection of it in a small marsh in Michigan (Hussey 1922). Indeed such is the variety of habitat types in New England that it frequently appears that two species are involved since the wet site populations are distinctly larger and darker than the dry slope populations. However the two forms interbreed readily in the laboratory, in- tergrade completely with each other when insects from all habitats are compared, and have the same sort of life cycle, so these popu- lation types are perhaps comparable to ecophenotypes. It appears that these forms may be what Blatchley (1926) was considering when he separated a smaller curvipes Stal from a larger basalis. Barber (1953b) synonymized the two. It is easier to state where P. basalis was not found. It was not found on dry very old fields especially those with north exposures ; on hot exposed open ground ; nor in ericaceous scrub, and uncom- monly in woodland glades and margins. It was never swept from vegetation. A few were sometimes found in flood wash with Peritrechus fraternus. In Florida it was found in grass clumps in the slash pine- palmetto association, but was not found in ruderal sites as these appeared to be preempted by other rhyparochromines especially P. bilobatus. As Blatchley noted, only the smaller form appears to be present in Florida perhaps for the above reason. The ground biotope was then always at least semishaded by the herbaceous vegetation. Soil moisture estimates ranged from 3 to 8, and litter temperatures were rarely over 30° C. Among the tall Panicum grass clumps the abundance frequently attains 60-70 per square meter, while the lowest abundances (1-3 per square meter) are in the more xeric sites with a shorter grass. It is significant that the most favored habitats are relatively short 16 VOLUME XLIV lived succession stages. Overwintering largely occurs in situ , although some appear along woodland margins in autumn, a few move even into wood- lands. General Biology In agreement with its preference for early succession stages, P. basalis is entirely macropterous. In probable dispersal, it has been collected at lights (Tucker 1907, Torre-Bueno 1914, Froesehner 1944). Torre-Bueno (1927) found it washed up in beach drift on Long Island. Glick (1939) collected it by airplane at 1,000 feet over Louisiana. In general form, Pachybrachius basalis is fairly generalized, and is not an especially fast runner, nor does it conceal itself among the tangled litter of stems and grass. The coloration is distinctly procryptic, a dull brown with a few light flecks on the hemelytra to break up the pattern. The nymphs display a disruptive color pattern in the later instars, and are pale yellow and red in the earlier instars. P. basalis is extensively parasitized by the tachinid Catharosia. The parasite was reared only from the second or summer genera- tion. The parasite overwinters in the host Pachybrachius. P. basalis is especially partial to feeding on grass seeds, es- pecially Panicum spp. and Paspalum spp. ; it also feeds on Festu'ca spp., Bromus sp., Andropogon scoparius Michx, Poa spp., but ignores these and various forb seeds when Panicum or Paspalum seeds are present. It would feed sparingly on Oenothera and Rumex seeds, and could be reared from field first instars on sun- flower seeds. However, the mortality of the laboratory reared first instars is very high when reared on sunflower seeds alone. P. basalis does not show any feeding reaction to pepper grass ( Lepi - dium sp.), Hypericum sp., Aquilegia canadensis L., Chenopodium albus L. or various small composite seeds. As Panicum is an enor- mous genus with many large and small species, this grass was present in most of the habitats. Torre-Bueno (1946) mentions P. basalis as being found on strawberries, but I was unable to rear even late instars upon this plant or its seeds. An excellent and effective rearing technique is a mixture of Paspalum and sunflower seeds. The presence of a few Paspalum (or Panicum) seeds cir- cumvents much of the first instar mortality. It is noteworthy that the labium or beak of this species is unusually short, and probably correlates with the small size of the preferred grass seeds. Only with Panicum and Paspalum seeds could the remarkable seed defense behavior of this species be elicited. The seeds are 17 ENTOMOLOGICA AMERICANA frequently picked up with the labium and moved to a more pro- tected site for feeding. When intruded on by another, a feeding bug endeavors to keep the seed away from the intruder. If the aggressor persists, the defender flutters or rapidly flicks its antennae back and forth through a 35° arc directly above its head. If this fails to dissuade the aggressor, the defender then ceases feed- ing, and with the antennae continuing to flutter, it quickly advances on the intruder. On contacting the intruder, the defender rapidly flails his legs. Usually the intruder decamps at this point. If not, the defender spreads its fore femora wide, holding them stiffly at right angles to the body, and rears up on its hind and middle legs. The intruding Pachybrachius immediately responds with a similar posture, and the two advance on each other, and rise into a vertical biped position on their hind legs. Through this en- counter the insects vigorously flail their fore tibia, and middle legs, and flutter the antennae very rapidly until one insect topples over. When this happens they immediately desist, and the loser, which in the observed cases was always the intruder, decamps. If the insects are hungry, this activity may be repeated again and again by the same insects. In no apparent way was any harm done to either insect. Of considerable phylogenetic interest is the sim- ilarity of the total defense pattern to that of P. bilobatus. In P. basalis both sexes are involved and there is no association with sexual behavior, although in P. basalis the male may, and does, take advantage of the female’s passive feeding posture. The mating behavior appears to be simple. A male responds to an appropriate female on contact by rapidly vibrating his an- tennae. If the female remains still the male usually attempts to mount, whereupon the female gives a rapid ‘ ‘ annoyance ’ ’ fluttering of her antennae. If the male at this point succeeds in mounting, he rapidly vibrates his antennae upon her head, and if the female is receptive, copulation is quickly accomplished. The male then drops off into an end to end position. More often the female dislodges the male, and decamps or may continue to stand him off with her antennae fluttering, especially if she is feeding on a seed, and the male may again attempt to mount the female. Cer- tainly, at least under the laboratory conditions, the male’s chances of mating with the female are greatly enhanced when she is feeding on a seed. Life History P. basalis in New England has a bivoltine seasonal cycle with a faculative adult diapause. Fertilization occurs in early spring as most females collected in April were fertilized, while those collected 18 VOLUME XLIV before March 20 were not. A pair was found en copulo in the field on March 29. However, as in Myodocha, the earliest nymphs are not found until early June and only the fourth instar was attained by June 23. Some first generation adults, however, appear be- fore July 1, and a few first generation fifth instars were present as late as July 17. With this maturation variability and es- pecially given the long oviposition period of one to two months, the second generation nymphs were of considerably different age ranges in different local populations. Thus, second generation first instars were found from July 14 to September 9, and second gen- eration fifth instars as late as November 1. However, many of the late autumn nymphs were parasitized. The observed overall phe- nology at Storrs is given in Table 17. TABLE 17 Phenology of Pachybrachius basalis Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 15 100% June 1 10% 90% June 15 15% 15% 30% — 40%- July 3 15% 20% 15% 10% 10% 30% July 17 6% 9% 12% 33% 40% Aug. 5 12% 10% 3% 15% 15% 45% Aug. 19 5% 35% 30% 2% 28% Sept. 5 8% 8% 12% 12% 47% 13% Sept, 15 5% 5% 40% 50% Oct. 1 1% 3% 11% 85% Oct. 15 2% 10% 88% Nov. 1 5% 95% The second generation adults, which appear from mid- August on, enter diapause under field conditions and overwinter. This is a facultative diapause because it did not appear when the second generation was reared under long day (15 hour) conditions. The converse, rearing the first generation under deliberate short day conditions, was not attempted. However, such a photoperiod re- lationship renders comprehensible early spring cultures in which the first generation was forced to maturing in the laboratory (nor- mal spring daylight) before early June. These cultures went di- rectly into diapause. The diapause state is only moderately strong 19 ENTOMOLOGICA AMERICANA and is spontaneously broken under laboratory conditions in late October or early November in adults reared in August. Adults which became mature in September did not break diapause in the laboratory until January. The first generation which went directly into diapause in June became reproductive by mid-August. The period of reproductive cessation averaged two months and varied from 45 days in summer to three months in autumn. P. basalis population collected at Cherokee and Roam Mt., North Carolina displayed a similar seasonal periodicity and a P. basalts population from south Florida collected November 19 at Naranja, was in a state of reproductive diapause similar to the northern populations and similarly required a period of warm laboratory con- ditions, but here only a five week interval was required to break reproductive cessation. Photoperiods in the late fall were long because of the lighting conditions prevalent in the laboratory. In the laboratory, on a sunflower seed diet alone, the stadia of the surviving nymphs are variable and unusually prolonged growth rates similar to nymphs captured in the field are obtained on mixed sunflower — Panicum grass seeds. The stadia information on this diet under room temperatures are given in Table 18. TABLE 18 Stadia of Packybrachius basalis Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 9.6 (7-12) 7.1 6.3 5.4 6.6 (6-9) (5-8) (4-7) (4-8) 8.0 (6-10) 43.5 (38-51) The longevity of laboratory summer generations varied from 34-52 days (mean 44). April collected adults survived 26-58 days (mean 39). Diapausing adults in warmth lived 2.5 to 7 months (mean 5). The longevity is then clearly related to reproductive activitv. P. basalis has a fairly long preoviposition period (first genera- tion) of about a week. In the laboratory 62 to 210 eggs (mean 108) per female were laid. In adults which are allowed to complete diapause under warm conditions the fecundities are much lower, varying from 7 to 36 eggs. The eggs are laid singly at the rate of 6-7 per day. Ten sexually isolated females laid no eggs, but one spring female laid 9 eggs. Oviposition occurs freely with sunflower ;seeds alone, although complete rearing was rare on this diet. 20 VOLUME XLIV The female investigates each oviposition site first with her antennae, and then with her ovipositor. The eggs are oviposited into crevices, soil, and rather tight wet or dry cotton, even into parenchyma of herb stems and the inner coat of sunflower seeds. The eggs are long, slender, with the apical end somewhat pointed, the anterior flattened, and the chorion apparently has no sticking mechanism. This shape appears adapted for oviposition into firm substrates. Pachybrachius albocinctus Barber This species was only recently described (Barber 1953a) as it has been much confused with P. bilobatus or following Stal (1862), was erroneously called P. servillei Guerin. It may be easily dis- tinguished from P. bilobatus by the basal white band on the ter- minal antennal segment. Like P. bilobatus, P. albocinctus has a remarkably broad distri- bution throughout the American tropics but extends somewhat farther north to Illinois, Michigan, Ohio, and New York, and is here recorded from Connecticut. Its presence in Connecticut, then, is at the northern edge of its range. A population of this species has been present for at least eight years in Storrs, Con- necticut (Slater, in lift.) so a permanent population appears to be established. Environment This species is closely adapted to wet habitats. Froeschner (1944) collected the species from shrubs and weeds near water and considered it not uncommon in Missouri. Blatchley (1926) mentions sweeping it from low moist ground, and beating it from Spanish moss in Florida. Not surprisingly, then, Wray and Brim- ley (1943) recovered this species from pitcher plants (Sarracenia) in North Carolina. In Florida the relation of the species to water margins was striking and effectively isolated albocinctus from bilobatus which was found in much drier sites. In New England P. albocinctus was found only at two stations near Storrs, Connecticut. At Pink Kavine (in Storrs), it was found abundantly with Ligyrocoris car ids in the Car ex strict a- Scirpus sp. community on the margin of the pond, frequently above standing water (see L. caricis for a description of this habitat). In this community adults of albocinctus are frequently swept with Ligyrocoris from the early maturing clump-forming Car ex where thev are feeding on the seeds. Much more frequently both adults 21 ENTOMOLOGICA AMERICANA and nymphs are found feeding on the seeds of the non-clump form- ing Scirpus sp. which matures considerably later. However, only a minority occur on the plants ; the majority are found on the ground at the base and in the clumps of Car ex, frequently even on mud and wet plant debris. The substrate moisture level is very saturated (10) and the temperature moderate. The abun- dance was low (1-3 per square meter of Car ex clumps) in spring and rose to about 20-30 per square meter in late summer. These abundances are only approximate because of the counting difficul- ties encountered in this habitat. General Biology P. albocinctus is completely macropterous and when provoked readily takes flight from a Scirpus seed head. It was taken by airplane, presumably dispersing, at 200 feet in Louisiana (Glick 1939). It was taken at lights (Barber 1954) and I have frequently collected it at a lighted sheet in July and August. The ability of this species to disperse is perhaps best attested to by its broad dis- tribution not only in continental areas but also throughout the West Indies. The adults are procryptically colored brown and tan, as is fre- quent in species living in rank sites. The terminal antennal seg- ment has a basal white band which is especially conspicuous when the resting insects move the antennae to and fro against the dark background of the marsh habitat. Such a band (Cott 1960) is usually interpreted as a device to distract a predator’s attention. In contrast, the nymphs, except the first instar, are conspicuous ant mimics with the “petiole” formed by white patches upon the anterior abdominal area. P. albocinctus was found parasitized by a tachinid fly Alopho- rella aeneoventris (Will.) a parasitism which is unusual in that it does not involve Catharosia. Sabrosky (in lift.) tells me that Dr. Medler has reared a species of this tachinid genus from a mirid bug, Lygus spp. The parasites were recovered from specimens of P. albocinctus which were found hibernating in late November. The parasites emerged only after a long interval in the laboratory, and so apparently were overwintering in the host bug. Catharosia was also reared from P. albocinctus collected in Florida. The adults and all instars feed upon the seeds of Jussiacea. The insects could be readily reared in the laboratory on sunflower seeds. For some unknown reason, feeding turned the seeds to a bright green color. No seed defense behavior was observed in P. albocinctus. The mating behavior is as follows : When introduced to a female the 22 VOLUME XLTV male does not overtly respond until he comes in contact with her. However, the female seeks out the male in each case and responds first by rapidly vibrating her antennae. After about 12 seconds the male responds by similarly vibrating his antennae. It is noteworthy that the white antennal band is very conspicuous when so vibrated, which suggests another, perhaps true, function to the white band. After a remarkably long time of such face to face antennae vibrating (45 seconds) the male slowly climbs upon the passive female’s right side and vibrates his antennae close to the female’s head. The pygophore is extruded, and with the male’s claspers (parameres) working he moves the pygophore up and down the side of the lateral side of the abdomen. If the female does not release her ovipositor, the male decamps. Again, the female may seek out the male, and the procedure be repeated. If effective, the female raises her abdomen high, and releases her ovipositor which is then clasped by the male ’s parameres ; in this position each sex is at a considerable angle with the left sides tilted down, a re- lation which persists when the male drops off into an end to end position. During the initiation of copulation the male suddenly flickes his antennae several times through a 45° arc. This flick is the essential component of the rapid up and down antennal fluttering, the “annoyance” signal of P. basalis. Copulations last from 1 hr. 45 min. to 2 hr. 30 min. P. albocinctus is completely isolated sexually from P. basalts and other large myodochine genera, apparently by a complete nonrecognition. Life History In Connecticut, P. albocinctus has a bivoltine seasonal cycle with no diapause intervention. The earliest nymphs, however, are not found until late June, and, largely third and fourth instars are found in the field as late as July 25. First generation fifth instars are found as late as August 28. Adults of the new genera- tion appear in early August, rarely in the last week of July. The second generation appears in mid- August, and new adults appear in late September. Many nymphs are still present on the latter date but these are presumably killed by the advent of cold weather as only the adults are found overwintering. It is clear that this seasonal cycle is barely adequate to allow the completion of two generations. There is also no reproductive diapause in the adults whether reared in the laboratory or taken from the field, and re- productive cessation is established by cold dormancy. The over- wintered females collected in early spring were unfertilized. The life cycle in the field took approximately 7 weeks, in the laboratory about 6 weeks. A considerable increment of warmth 23 ENTOMOLOGICA AMERICANA seems required as the early spring adults would readily oviposit in the laboratory. The tardiness of the spring generation is probably best explained by the cool substrate conditions, since laboratory behavior indicates that eggs are laid in the moist substrate and egg development is relatively slow (11-13 days) even under the warmer laboratory conditions. The laboratory stadia based on the progeny of three females are given in Table 19. Nymphal de- velopment occurs much more rapidly and with a much smaller mortality rate on a mixture of Scripus-Carex and sunflower seeds than on sunflower seeds alone. The longevity in the laboratory of two mated females was 57 and 74 days of two virgin females 71 and 76 days and of two males, 43 and 45 days. TABLE 19 Stadia of PachybracJiius albocinctus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 12 10.4 9.5 8.5 8.3 8.4 52.6 (11-13) (7-18) (9-10) (6-12) (5-11) (6-10) (45-58) The precopulatory period is 6-8 days and oviposition occurs 1-2 days later in mated females. Unmated females of the summer generation lay no eggs. The fecundity of 5 females varies from 82 to 231 eggs (mean 140). From 2 to 10 eggs (mean 5.2) are laid daily. The insects oviposit on wet substrates, but dry substrates are acceptable if no wet ones are present. The eggs are frequently forced into very tight wet cotton stoppers. In correlation, the eggs are smooth, slender, and somewhat pointed at the posterior end. Pachybrachius bilobatus (Say) This dominant and widespread species is known from throughout the American tropics north to Missouri, Pennsylvania, New Jersey, and has been recorded once from New Haven, Connecticut (Barber 1953b). It was not found in New England in the present study, but from its habitat preferences in Florida, it is to be searched for in warm ruderal sites, dry lawns and gardens, a fact which, along with its propensity to occasionally climb into vegetation and thus be swept, has led to its ready collection. 24 VOLUME XLIV While a Florida population has been reared readily on sunflower seeds, I will not discuss this species in much detail as it has been recorded as a pest of strawberries along with P. vinctus (Sweet 1960, for references) and undoubtedly economic entomology work- ers will work out its biology in closer detail. For the present discussion, it is most interesting to note that this species exhibits an elaborate seed defense behavior in part sim- ilar to P. basalis, but also a distinct sexual element is involved. When a few seeds are available some of the males station themselves by the seeds, only occasionally feeding, and drive away other males, nymphs, and some females. But if an appropriate female ap- proaches, she is allowed to feed on the seed and the male attempts to copulate, mounting her and rapidly vibrating his antennae against her head. Usually the female would rapidly wag her an- tennae in “annoyance” as in basalis and decamp. Occasionally copulation would ensue, instead, if the female was receptive. When nymphs or some females intrude on the seed they are repulsed by the male who advances on the intruder and touches it with his antennae which are held stiffly and vibrated rapidly in a very slight plane. Once when a dominant male returned to a seed, after he had copulated with a female, he dislodged four in- sects, a weak male, a female, a third and fifth instar which were feeding on the seed, by climbing upon the back of each and 'em- ploying this slight but rapid antennae vibration. The territorial limits of this seed possession extends to about 1 cm. from the seed and frequently the defending male is at the out- skirts of this zone, not on the seed. When a male intrudes, he is repulsed with great vigor and sometimes chased across the rearing dish. In several such chases the dominant male became, so to speak, lost. Thereupon a formerly recessive male took over the defense of the seed, and attempted to copulate with a female. Later the dominant male returned to this particular seed, despite the pres- ence of other similar seeds, and touched the smaller male with his vibrating antennae in the usual manner. This time a different re- sult ensued. The small male clung close to the seed and would not dislodge, whereupon the dominant male suddenly began to flutter his antennae rapidly and stiffly spread his fore femora wide much as in P. basalis. The small male almost simultaneously likewise spread his fore femora and fluttered his antennae. The two males reared upon their hind legs and clashed for 5-8 seconds with flailing mid legs and fore tibia and very rapidly fluttering antennae. Suddenly both desisted, and began to clean their antennae. This appears to be displacement behavior. The large male then re- sumed possession of the seed. 25 ENTOMOLOGICA AMERICANA This behavior is described for it illuminates an unusual sexual dimorphism, for the males of P. bilobatus while variable in size among themselves are usually larger than the females, an unusual situation among insects (Darwin 1871, p. 628). Among mammals the greater size of the male is explained as a result of intraspecific selection among males in competition for females (Darwin 1871, p. 831, Simpson 1951, p. 86), and a similar explanation seems apparent here as among stag beetles (Darwin 1871, p. 628). Also of considerable interest is that cultures with many com- peting males and resultant continuous agitation, underwent a great population decrease which was prevented in one culture by re- moving the adults. Clearly this species would be an excellent sub- ject for a study of a possible stress syndrome. Wilson (1938) found P. bilobatus parasitized by a fungus Beauveris bassiana (Bals.) and preyed on by three species of lizards in Puerto Rico ; Anolis klugii Barbour, Anolis stratulus Cope, and Anolis cristabellus Dumeril and Bibron. Pseudocnemodus canadensis (Provancher) This monotypic genus bears a pronounced resemblance to Cnemodus. Pseudocnemodus canadensis has a boreal Nearctic distribution from Quebec to British Columbia, south to Iowa, northern Indiana, southern New England and New York, and in the Appalachian Mountains to North Carolina (Slater, Catalogue). It appears to be scarce in Iowa and Illinois (Slater 1952). Torre- Bueno (1912) considered it to be a “pretty common and wide- spread species” in New York. Environment In New England Pseudocnemodus is typically found on dry overdrained slopes which have a sparse but complete ground cover especially on dry edge habitats between forests and an old fields. Such areas are dominated by low bunch-forming grasses such as Festuca rubra L., Andropogon scoparius Michx., and Aristida dicho- toma Michx. Open patches of the low blueberry V accinium au- gustifolium Ait. also form a common habitat. These sites have rather uniformly gravelly dry soils, with the interspaces between the grass clumps filled with fallen litter, Cla- donia and Polytricum. Xestocoris nitens, Carpilis consimilis, Trapezonotus arenarius, and Lygaeospilus tripunctatus (Dallas) are seed feeding insects that frequently occur with Pseudocnemo- dus. The ground biotope, at the bases of the grass clumps or Vac- 26 VOLUME XLIV cinium low scrub, is relatively dry (2-3) and becomes quite hot (up to 48 °C. during mid-day). While these field habitats do form an early serai stage, much dry overdrained sites only slowly undergo succession. In at least one such area a population of Pseudocnemodus was present con- tinuously for at least nine years (Slater, in lift.). However, the serclimax V actinium scrub habitats of exposed rock faces forms a practically permanent habitat. The abundance of the insect is usually fairly low, in the order of 3 to 8 per square meter, more rarely as much as 16 to 20 per square meter. Hendrickson (1930) who found it in a number of climax associations, similarly did not find it numerous. The locality at Vernon, British Columbia was a site dry in summer and cool in winter, and very different from the wet and equitable coastal climate (Parshley 1919). General Biology Since all the field populations are largely brachypterous the presence of Pseudocnemodus in a mosaic of scattered favorable habitats indicates a sufficient production of dispersing maeropters. Dispersing adults are indicated by the large numbers of individuals, all macropterous, washed up on ocean beaches on Long Island (Torre-Bueno 1915, 1927). This long legged myodochine is an alert rapid-moving insect and shows little tendency to hide under the litter. Both the adults and especially the reddish nymphs display a relatively weak ant mimicking appearance and behavior. A pale lateral margin to the hemelytra and the pale humeral angles heighten the narrow shape of the insect which blends it into the dry grass background. Several populations were intensively parasitized by Catharosia tachinid flies. One population of Pseudocnemodus at a considerable density of 10 per square meter was 94% parasitized, a factor which may have led to the extinction of this population in contrast to the very long lasting population mentioned earlier where parasitism was never found over six years. The tachinids emerged after the Pseudocnemodus nymphs became adult. Only one of many pupae yielded a fly, which may indicate a pupal diapause condition, which correlates with the egg diapause of Pseudocnemodus. In the laboratory Pseudocnemodus rears readily on sunflower seeds. It expresses a strong feeding reaction to V actinium and Gaylussacia seeds. It also feeds on the seeds of Festuca rubrum, Hedeoma sp., Solidago , Aquilegia canadensis, Betida popidifolia, and Rumex obtusif olia. It appears then, on this evidence, to have a large potential seed feeding range. It also will scavenge on nearly 27 ENTOMOLOGrlCA AMERICANA dead or dead adults but no active cannabalism or predation was observed nor indicated. No seed feeding defense was observed. The insects drag the seed to more protected sites beneath loose litter. Ashlock and Lattin (1963) have described an interesting large stridulatory area (stridulitrum) along the side of the pronotum. This stridulitrum occurs also in the nymphs. The plectrum is an irregular row of small tubercules along the inside of the fore femora. This mechanism occurs in both females and males. Ashlock and Lattin note that when irritated, this species moves its femora rapidly up and down next to its body. These authors heard no sounds but assumed that if produced the sounds are inaudible. This stridulatory area may also play a role in the mating be- havior of the male. When placed with a receptive female, the male responds to the female before contacting her. The male’s response is shown by his rapidly vibrating antennae, and by a peculiar awk- ward-appearing partial spreading and closing of the fore femora. This spreading at first was thought to be a display of the shiny inner surface of the fore femora but further study showed that this maneuver effectively rubs the fore femoral plectrum over the stridu- litrum. No similar motions were observed in the female, however. If receptive, the female briefly approaches the male and also vibrates her antennae, and the antennae of the pair meet. From this posi- tion the male slowly rotates into a parallel position to the female and mounts her. Once upon the female, the male loosely taps the female’s head with his vibrating antennae. Copulation is effected and the male drops off into a reversed (end to end) position. Copu- lation lasts from 44 minutes to at least several hours. One pair mated at least five times, but a single mating is sufficient to fertilize an entire egg complement. Life History Pseudocnemodus canadensis diapauses over the winter as an egg. Both uni- and bivoltine conditions occur. The earliest first instar nymphs were found in the field May 24, but the first instars oc- curred as late as June 10, at which time third and fourth instars also were found. This variability indicates staggered hatching, which agrees with the artificially over-wintered eggs, which exhibit a very variable rate of hatching. This situation is of considerable significance, for those insects which became adult in the field shortly after mid- June went on to lay non-diapause eggs, while those which became adult after June 31 uniformly laid diapause eggs. This interesting situation is true in both field and laboratory popula- tions. Pseudocnemodus populations such as one on a cool north 28 ENTOMOLOGICA AMERICANA side of a drumlin are entirely of this late developing type, while the several populations with bivoltine females (mixed with late de- veloping univoltine individuals) are found in considerably warmer sites. Unfortunately this situation was not followed up, as the nymphs could not be forced to early maturity in the laboratory under short daylight conditions. This situation could not result from temperature conditions as the peak in field temperatures came in late July and early August, and only diapause egg laying adults are present, and the lower laboratory temperatures also yielded diapause adults. The most logical explanation is that under the solstice long day conditions, the insects are stimulated to produce non-diapause eggs. Another factor possible may be a genetic factor which allows a more rapid development and also sensitivity to long day conditions. At any rate there are two phases of diapause egg oviposition, one in summer from July to late August, another by the partial second generation from mid-August into September. Both first and second generation eggs were artificially overwintered and diapause broken at 4°C. after 5 months of cold. Diapause occurs during early anatrepsis. The total life cycle varies from a month to a month and a half, and second generation fifth instars are found as late as September 10. The overall phenology is given in Table 20. Bear in mind that local populations may differ greatly in the instar ratios due to microclimatic differences and staggered hatching, etc. TABLE 20 Phenology of Pseudocnemodus canadensis Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 24 100% — June 1 40% 40% 20% June 10 8% 44% 36% 8% 4% June 22 20% 35% 25% 15% 5% July 2 2% 16% 25% 32% 25% July 18 2% 3% 5% 20% 70% July 28 2% 8% 90% Aug. 18 4% 4% 8% 14% 10% 60% Sept. 5 2% 11% 87% Sept. 27 100% The stadia given in Table 21 are based largely on nymphs cap- 29 ENTOMOLOGICA AMERICANA tured in the field and reared in the laboratory. Because only a few populations with non-diapausing adults could be found, the egg and first instar stadia are represented by only two records each. After removal from cold the few eggs which immediately developed hatched in 15 days. The adult life span in the laboratory varies from 35-75 days. The average longevity of ovipositing females was 49 days, considerably less than virgin females (69 days) or males (59 days). TABLE 21 Stadia of Pseudocnemodus canadensis Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 5.0 7.7 6.0 8.8 40 (12-15) (6-7) (5-7) (6-9) (5-7) (7-11) (38-48) There is no summer reproductive diapause and oviposition of diapause eggs ensues after a preoviposition period of about 14 days. From 79 to 280 eggs per female were laid (mean 154). When an unsuitable oviposition substrate was provided the egg production was drastically reduced to only 1 to 25 eggs. Virgin females laid from none to only 19 eggs (mean 6.1) over a two or three month period. The eggs are of average size, cylindrical with rounded ends and densely covered with very short minute tubercules. The eggs are preferably laid deeply in fine loose dry litter debris, which sticks to the eggs, covering them up, and effectively conceals their presence. Cnemodus mavortius Say This striking large black myodochine is exceptional in that both the macropterous and the braehypterous forms lack all visible traces of ocelli. Its generic name, which means “ well-legged, ” (Grover 1876), aptly describes the general appearance of this insect. Four other species are presently known in the genus. Berg (1879, 1894) described albimaculus from Argentina and Uruguay and multifarius from Bolivia. Blatchley (1924) discerned a separate species, hirtipes, which is apparently endemic to Florida. However, the validity of the fourth species, inflatus , which was described from North Carolina by Van Duzee (1915) has been ques- tioned by Froeschner (1944). Froeschner found that populations 30 ENTOMOLOGICA AMERICANA of C. mavortius in Missouri exhibited a large range of structural variation, largely associated with pronotal changes associated with brachyptery and considered inflatus to represent an extreme form. Torre-Bueno (1946) disagreed, and considered the structural differ- ences between the two forms of rather great magnitude and as indicating species differences. My own work on Connecticut popu- lations of C. mavortius agrees with Froeschner’s observation that there is a rather striking variation in total size and pronotal shape in relation to brachyptery but there is also a sexual dimorphism in- volved, as the males are variable in size, and both mavortius and inflatus types can be distinguished. Since I have not seen the type of inflatus , this form cannot yet be synonymized. Cnemodus mavortius has a generally southern range from Georgia and Texas north to Iowa, Indiana, New York, southern New England, Nantucket Island, Massachusetts and also along the coast as far north as Maine. In New England I have found the species fairly frequently at a number of sites in southeastern New England, but not at all north of northwestern Connecticut. It appears to be uncommonly collected (Froeschner 1944, Slater 1952) . However this can be accounted for by the habitat specificity of this insect. Environment Cnemodus mavortius in Connecticut is highly specific to fully exposed old overdrained morainic sites dominated by Andropogon scoparius, especially where considerable areas of bare gravelly soil are present. Several old abandoned gravel pits with such a sparse cover of Andropogon formed a very typical habitat. In a few habitats the plant cover although short completely covered the ground. The soil is always dry (1-2), sandy or gravelly, with a sparse litter of fallen culms around the Andropogon clumps. The surface soil temperatures frequently become very high (55°C.) at these sites, and the insects at such times are frequently observed perched on bits of litter and noticeably very high on its legs as is usual with species found in such hot habitats (Cloudsley-Thompson 1962). The thermophily of this species is especially well shown in the laboratory as the insects will thickly cluster close to a source of warmth. It was never swept from plants and never found on the grass stems as is Nitheeus jaeobeae (Schill.) when it is exposed to high temperatures (Coulianos 1961). Cnemodus is found abundantly (up to 25-35 per square meter) at sites where considerable bare soil is present, and less abundantly (1-3 per square meter) in more closed over habitats. It is apparent that Cnemodus is a highly thermo- and xerophilous insect which 31 ENTOMOLOGICA AMERICANA inhabits a early succession stage on xeric old fields with poor soils. Such habitats, however, while obviously temporary are very long enduring (Blizzard 1931, Conard 1935) and may persist in such an open state for many decades. The insect is found in refuge in thick Andropogon clumps for hibernation, especially in clumps close to the margin of a field. Such an overwintering site may he meant by Blatchley (1926) when he described its habitat as ‘‘beneath dead leaves on the wooded slopes of streams. ’ ’ This is a very different sort of habitat from the xeric hot habitats where Cnemodus is found in Connecti- cut. Several authors record it as overwintering in wooded areas (Uhler 1875, Blatchley 1895, Froeschner 1944). General Biology The populations of Cnemodus studied in Connecticut are largely brachypterous, especially those in restricted populations. In the larger populations a variable percentage of macropters are found. Since the habitats described are fundamentally temporary although long enduring, it is therefore apparent that a sufficient production of macropters does occur so as to allow dispersal to new habitat types. Sometimes, however, as at a renewed gravel pit near Storrs, Connecticut a population of brachypters was but a short distance away and dispersal to this new habitat more likely occurs via the ground. Cnemodus was collected at lights (Torre-Bueno 1933) and so the macropters may disperse at night. Cnemodus hirtipes of Florida has also been collected at lights (Blatchley 1926). It would be interesting to ascertain if the absence of ocelli in any way affects the dispersal ability of this species. The prevalence of brachyptery may help explain its spotty local distribution in available habitats. In the field and the laboratory the adults of Cnemodus give an immediate impression of being huge black ants. This resemblance to ants is intensified by the jerky movements of this rapid moving insect. Also noteworthy is that the brachypters more closely re- semble ants than the macropters. Cnemodus is by far the largest (in length) of the rhyparochromines of New England and is also much larger than the largest ant ( Camponotus pennsylvanicus) and very much larger than the common black Formica. Therefore, the ant mimicry resemblance is a general one and not based on a single model. The nymphs, far more than the adults, are striking ant mimics. Instead of black the nymphs are reddish with white lateral patches along the abdominal Y-suture immediately posterior to the thorax, which helps to create the impression of a petiole as the pronotum and posterior part of the abdomen are swollen in 32 ENTOMOLOGICA AMERICANA appearance. The development of mimicry appearance is gradual, the effect heightening with each instar after the first instar which itself is yellowish with a red abdominal crossband as in other myodochine first instars. In effect, then there is a polymorphic ant mimicking population with the adults and nymphs having dif- ferent colors. Certainly, against the pale soil and dry brown vegeta- tion of the Andropogon community, these insects, especially the black adults, stand out conspicuously. Berg (1879, 1884) described similar ant mimicry in both adults and fifth instar nymphs of Cnemodus albimaculus Berg. Cnemodus is very frequently parasitized by Catharosia tachinid flies especially when the Cnemodus population density is relatively high. The parasitism appears to occur during the later instars. The parasites all emerged from the adults only. Parasitized fe- male Cnemodus can be recognized readily in the laboratory because they are non-reproductive. The parasite diapauses through the winter in the host as the parasite emergence does not occur until during the winter from the fall adults which were kept in warmth or in spring from the overwintered adults. Normal adults become reproductive in a brief time, while the parasitized adults continue on as if they were in reproductive diapause. Cnemodus feeds on Andropogon seeds in the laboratory, but was never swept from Andropogon seed heads, and probably feeds only on the fallen seeds in the field. There was some feeding on Oenoth- rea sp., Acer rubrum. and Quercus alba seeds. An especial1 v strong feeding reaction was shown to ripe Uumex acetosella seeds, but it did not feed on Panicum and Agropyron grass seed, and on various other composites. Yet it feeds readily on sunflower seeds, both hulled and unhulled. No active seed defense behavior is exhibited aside from keeping a seed out of the reach of another individual by placing its body between the seed and the intruder. The seeds are carried on the labium to more protected litter sites. The seeds are located largely by probing with the labium. When feeding the antennae are char- acterically held at right angles to the body. Although the insects were frequently observed mating, the* courtship behavior was not observed despite continued attempts.. At least it may be said that a receptive pair reacts rather slowly* to each other and no apparent reaction occurs for several hours before mating. Copulation lasts at least two hours in the usual reversed position, and may be repeated several times, but only one mating is sufficient to fertilize the entire egg complement. Two generations a year are present in Connecticut, and the adults overwinter. Copulation occurs in the field in late April 33 ENTOMOLOGICA AMERICANA and May and the fertilized females oviposit immediately on being brought into the laboratory, but the earliest nymphs were not found in the field until mid- June. The phenology at Storrs, Con- necticut is given in Table 22. Oviposition occurs over a period of several months and the second generation considerably overlaps the first. Last instar nymphs which are found later than October 10, are very frequently parasitized. TABLE 22 Phenology of Cnemodus mavortius Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 1 100% June 25 10% 30%. 20% 40% July 10 15% 15% 30% 20% 10% 10% July 27 10% 20% 70% Aug. 10 20% 25% 55% Sept, 1 5% 9% 26% 14% 9% 37%. Sept. 17 5% 10% 15% 70% Oct. 10 10% 90% Nov. 1 2% 98% There appears to be a relatively weak diapause mechanism, and in the field adults of the second generation are non-reproduc- tive. This quiescence is ordinarily broken readily and most in- dividuals become reproductively active in the laboratory in early October. However the progeny of a forced third generation was in a much stronger diapause state and did not become reproductive after three to five months. Since the general distribution of this species is to the south of New England, it may be hypothesized that this curious increase in diapause strength may represent the normal seasonal cycle further south under a longer season with a shorter late autumn photoperiod. The weaker diapause state of the second generation in Connecticut under equinoctal conditions may represent a de- veloping adaptation to north temperate conditions. It is significant that a few Cnemodus did not break diapause in October as did the great majority of the specimens. The stadia of instars 2 to 5 given in Table 23 were obtained Ly rearing field collected nymphs in the laboratory. When reared from .eggs on sunflower seeds the stadia were greatly prolonged as 34 ENTOMOLOGICA AMERICANA seen in the data on instar 1. The life span of the nondiapause adults in the laboratory was long, averaging 67 days (range 35-87). The diapause period of second generation adults averaged 35 days and the adult life span averaged 103 days. Forced third generation adults in diapause lived from 60 to 168 days (mean 97). TABLE 23 Stadia of Cnemodus mavortius Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 11.3 15.2 8.4 6.5 6.0 8.4 52 (8-14) (12-26) (4-8) (4-9) (4-9) (5-11) (39-67) The preoviposition period was 10 days in two non-diapause females. The individual fecundity in the laboratory of 19 females varied from 54 to 280 eggs (mean 128) on a sunflower seed diet. Twelve unmated females laid no eggs. The eggs are elongate and smooth without any spines or sticking mechanism. The females oviposit in dry substrates, such as litter crevices, sand, or in other loose material such as cotton and methyl cellulose. Rarely the eggs were laid in wet stoppers. Egg production is drastically reduced when a proper loose substrate is lacking. Ptochiomera nodosa Say Ptochiomera nodosa is a bizarre little species as its antennae are greatly enlarged, especially the third segment. In this respect it is most closely related to Sisamnes Dist. Formerly Barber (1928a) included in Ptochiomera various species now distributed in three other genera which are closely related ( Ptochiomera Say, Sisamnes Dist., Carpilis Stal, and Exptochiomera Barber). Pry- tanes Distant also belongs to this complex (Barber 1954). Later Barber (1935a) recognized the diversity of this group of short legged small myodochines and considered the above genera as dis- tinct, and restricted Ptochiomera to the monotype, nodosa Say. Since many early European workers used the emendation of Ptochiomera , Plociomera, to refer to myodochines in general, there are various other species now listed under the name Ptochio- mera (Slater, Catalogue). The genus is Neotropical as well as Nearctic for I have observed 35 ENTOMOLOGICA AMERICANA an undescribed species from Venezuela in the United States National Museum. Ptochiomera nodosa , itself, is distributed over the south- ern part of the United States from Florida and northern Mexico north to Iowa, Indiana, and Massachusetts. It is here recorded for the first time from Connecticut. Blatehley (1926) noted that this species was common in the southern tier of counties in Indiana, but scarce in the north counties. Slater (1952) noted it only from the southern counties of Illinois and the extreme southeast of Iowa. Froeschner (1944) stated that it is abundant in Missouri, and Uhler (1876) found it to be common in Maryland. In New England I found the species only along the extreme southern coast along Long Island Sound at Noank, Rocky Neck, and New Haven, Con- necticut. The Massachusetts record is from Woods Hole (Parshley 1917). Because of the relative scarcity of P. nodosa in Connecticut the following discussion is rather limited. Environment Nearly all my collections of this species have been along road sites and parking lots except for a few found along the margin of a lawn next to a beach at New Haven. At these sites it was narrowly confined to the dry thin litter along the sharp edge between the ruderal plants and the open road, lot, or beach. It was never found where the vegetation thickened, and formed a continuous cover, nor under dry litter in open areas, nor where the litter was at all moist. In southern Florida 1 found this species in essentially similar biotopes along the margins of open fields and roadsides. The soil was always light colored and sandy or gravelly and mixed with bits of grass litter. The temperatures at mid-day were often high and measured between 110-140° F. in the litter area but con- siderably lower than on the open ground. Of course, the litter and and ground were quite dry (1-3) at all stations. The vegetation consisted of the usual ruderal plants, chiefly Agropyron repens, Poa annua L., Digitaria sanguinalis (L.), Rumex obtusifolia L., Silene sp., Potentilla pumila Poir., Lepidium virgineum L., Bras- sica sp., with Agropyron the usual dominant species. Clearly these habitats are at best very temporary habitats which would rapidly undergo succession except for man ’s activities which cause the roadside community to conceivably be a sort of disclimax. But even among ruderal communities, this marginal biotope is especially short lasting. P. nodosa was not found for several years in succession at any sites except the one at Noank where it was found in the summers of 1957 and 1958. Some authors have mentioned P. nodosa’ s abundance in the 36 ENTOMOLOGICA AMERICANA southern states but at its northern fringe in Connecticut it is rep- resented at most sites by only a few specimens an abundance level of less than 1-2 per square meter. The only exception was at a parking lot in Noank where it was sometimes found at 20-30 per square meter. At this site the insects appeared to be distinctly gregarious and formed close groups of 10-20 which clung closely to the base of the small dried thin grass tufts. In the laboratory a similar tendency was found and the nymphs and adults would congregate under a bit of litter. The habitat notes in the literature appear to bear this observation out. Blatchley (1895, 1926) found the species beneath chunks and rubbish and at the bases of tufts of grasses along the margin of open cultivated fields and road sides. He also observed that it over- wintered at these same sites. Froeschner (1944) found it under logs, boards, rocks, and grass clumps in Missouri. Perhaps the oddest reference is that of Wirtner (1905) who said, “Klages tells me that he finds it abundantly on the mushrooms on trees.” General Biology In the brachypterous form the hemelytral membranes slightly overlap and just attain the anterior margin of tergum seven. Despite the essentially temporary nature of the habitat of this species both macropters and brachypters were always found in about equal proportions. Clearly this level of macroptery must allow sufficient dispersal capacities to colonize such temporary habitats. In this respect it is significant that whether colonies begin with brachypterous or macropterous forms both yield progeny of approximately equal proportions of brachypters and macropters. Uhler (1876) stated that the short winged form was found in the granitic and primitive areas in Maryland but was full winged in the newer and more southern portions of this region and in the southern states was always long winged. Blatchley (1926) dis- agreed and wrote that the species was fully one-half brachypterous in Florida. I have observed much the same in south Florida and Texas. But Uhler ’s observations may eventually still be significant in the relation of such macroptery and brachyptery to permanent and temporary habitats. Dispersal may occur at night as it was collected at lights (Froe- schner 1944, Barber 1953a). Glick (1939) collected this species at 3,000 feet over Louisiana. The adult coloration, with a pale coriiun and posterior pronotum, and dark head and antennae blends the species readily into the light background of its habitat. When given a choice between dark earth and dry sand litter mixture, the adults come to rest 37 ENTOMOLOGICA AMERICANA on the light surface, and generally avoid the dark surfaces. The nymphs of instars two to five are bright red and white, very similar to Sisamnes clavigera nymphs, and so would appear conspicuous to bird predators. If so, this coloration is aposematic. The gregarious habit of this species may protect it from such a predator as Geocoris uliginosus Say which actively preyed on the smaller nymphs of Ptochiomera in the laboratory and in the field at Noank, Connecticut. The Geocoris definitely showed an avoid- ance reaction when it came on a group of Ptochiomera adults and nymphs. Such an aggregation is quiet and only by vigorously disturbing it would the insects move about. This is significant for the Geocoris was found to react especially to movements. Dis- turbing and dispersing this field colony of Ptochiomera brought about predation by Geocoris adults and nymphs which were abundant at this site. The only parasite was a catharosine tachinid recovered from a male Ptochiomera collected in Georgia in December of 1961 by D. B. Leonard. It is interesting that a distinct diurnal rhythm is present. Colonies from Connecticut and Florida both exhibited this behavior. This behavior was followed daily from March 15 to April 21. It has no relation to artificial illumination, darkness, or to variation in room temperature (68-78° F.). During the day a large colony of about 200 insects kept refuge in a pile of methyl cellulose. Be- tween 4 :20 and 4 :35 P.M. the insects began to come out of the pile. By 5 :25 to 5 :45 they were out in large numbers and climbing the sides of the container very actively. This activity continued to increase and reached a peak at about 9 :30 to 10 :00 when some macropters attempted flight. Both macropters and brachypters and even the early instar nymphs were very active. At 11 :15 the numbers were noticeably decreased, at about 11 :45 only a few were out, and between 12 :15 and 12 :50 all disappeared again into the pile. This species then exhibits a crepuscular and early evening activity rhythm, but not an auroral activity period. Exptochiomera dissimilis Barber from south Florida exhibits a quite similar rhythm. Ptochiomera feeds on seeds of Rumex acetosella, R. obtusifolia , Agropyron repens, Plantago, and is especially attracted to sunflower seeds. It is reared very readily and often with little mortality on the sunflower seeds. Some colonies, however, sometimes exhibit a considerable mortality in the earlier instars. There is no scaveng- ing on dying insects even in starving individuals. Despite the xeric habitat preference it could not be kept alive even overnight 38 ENTOMOLOGICA AMERICANA without water. As with other rhyparochromines these insects move seeds about and carry or drag them into the litter. When annoyed, the insects wag their antennae rapidly. This is true both when a male at- tempts to copulate with a nonresponsive female and when an intruder disturbs Ptochiomera feeding on a seed. Copulation occurs in the usual reversed position. The court- ship behavior was not observed except that the male’s antennae quiver very rapidly on contact with a female. The heavy antennae of both Sisamnes and Ptochiomera are used to help lift the insect above the substrate and so aid the short legs in the righting response. Probably the natural sandy habitat may make righting somewhat difficult and the antennae assist such a movement. Life History Uhler (1876) noted that this species overwinters as an adult and was among the first insects to become active in the spring in Maryland. There is not much evidence on hand and the life cycle is not adequately understood in the earlier part of the year. The life cycle appears very similar to that of Sisamnes in Connecticut, and the nymphs occur late in the summer. The observed phenology is as in Table 24, but very few Ptochio- mera were found before August 15. TABLE 24 Phenology of Ptochiomera nodosa Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult July 18 100% July 25 8% 11% 81% Aug. 5 10% 23% 67% Aug. 19 16% 21% 11% 16% 36% Sept. 3 6% 25% 32% 12% 25% Sept. 11 10% 30% 10% 10% 25% 15% Sept. 30 8% 8% 19% 65% Oct. 27 11% 11% 78% Oviposition in field adults occurred from July 30 to September 11, and mating was observed in the field on July 28. The evidence for two generations hinges on the occurrence of late instar nymphs in early August and the distinct break in the 39 ENTOMOLOGICA AMERICANA nymphal ratios on August 19. Unfortunately these late instar nymphs of early August were not isolated, and it is not known whether they became reproductive or not. All other nymphs from late August on, when reared, go directly into diapause and do not reproduce. The only, and significant exception, were the nymphs and adults collected on October 20 and October 27 which became reproductive after being brought into the laboratory. This is ex- actly the same pattern of cold release (?) found in Sisamnes. In Ptochiomera, however, the second generation extends even later into the autumn. Froeschner (1944) similarly recorded in Missouri a late oc- currence of nymphs from August 11 to October 3. It is possible that this late sequence of generations is a primary limitation on the northward spread of this species. At this same time in November and December in southern Florida and Georgia, only adults were found and Blatchley (1926) observed that the adults overwintered at Dunedin, Florida. Collections from Florida became reproduc- tive at Storrs in December and January. So readily do the cold exposed insects leave diapause that, this increases the probability that an earlier generation occurs. It is possible that a long photoperiod after a diapause period brings about sexual activity. Considerably more work remains to further elucidate this life cycle. At any rate this life cycle appears adapted, in Connecticut, to a late summer production of ripe seeds. The stadia are derived from relatively few observations and given in table 25. TABLE 25 Stadia of Ptochiomera nodosa Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10 (9-12) 6 7 6-7 7 9 (8-11) 41-47 Under diapause conditions in warmth the adults live 67-235 days (average 171). It is not known whether the sexually isolated adults lay eggs or not. The fecundity varies from 66-230 eggs (average 131) including data from adults from Florida. The smooth eggs are laid in litter, in methyl cellulose, are very well concealed, but do not stick to the litter as do those of Sisamnes. 40 ENTOMOLOGICA AMERICANA Sisamnes clavigera (Uhler) The genus Sisamnes at present contains three species. The type species contractus Dist. was described from Guatemala and is also known from Arizona and Florida. S. annulicollis was described by Berg (1879) from Brazil and Argentina. The species present in New England is clearly a temperate zone species and ranges from Massachusetts west to Idaho and New Mexico and south to Texas and North Carolina. Uhler (1876) mentions S. clavigera as being more abundant in the western states. From the published records the species appears to be especially rare in eastern United States despite more intensive collecting. It was not re- corded from Connecticut and Massachusetts until 1922 (Parshley 1922). Sisamnes clavigera is apparently unique among the Myodochini for it possesses thickened scale-like body hairs much as do the other lygaeids Slater ellus and Ckauliops (Drake and Davis 1960) and Heinsius (Slater and Sweet 1963). In all these genera the “scale” appears to be a waxy exudate for it melts when heated. Also re- markable are the peculiar stiff hairs on the clavate terminal an- tennal segment of Sisamnes clavigera. Environment The only meaningful habitat note is that of Torre-Bueno (1924) who found several specimens running rapidly among dry leaves near a clump of small yellow birches in Amherst, Massachusetts. Froeschner (1944) considered the species scarce in Missouri and found it under rocks, logs, boards, and at the base of mullein leaves. My own collection records are at variance with those of Torre- Bueno, but since I have collected this scarce species only a few times near Storrs, Connecticut, I obviously cannot give an accurate picture of its habitat preferences. The site where it was first col- lected (in 1958) was in an old sand pit on a gentle, south-facing small slope. The vegetation consisted of scattered clumps of Andropogon scoparius and a few other withered plants and lichens (Cladoni-a). The grass clumps were widely spaced with a thin loose dry litter of grass culms and seeds on the exposed ground be- tween the clumps. The soil was a very dry grey-brown sand (mois- ture level 1-2). The site was obviously, from the size of clumps and the accumulation of lichens and litter, rather old and had been undisturbed for some time. Because of the over drained sandy soil, succession had evidently proceeded slowly. Temperature measurements were not taken as the site was discovered in autumn but it was evident from the open exposure that this was a warm slope. When first found on October 10 the insects were dispersed 41 ENTOMOLOGICA AMERICANA over the slope and a few were even found in an adjoining- ruderal roadside. The density was 4-6 per square meter. Later checks after a period of cold weather seemed to indicate that the popula- tion was decreasing-. However, on November 5, a cluster of 25 of the insects was found within six square inches in a small accumu- lation of litter between the grass clumps, and another smaller group of 8 insects was similarly found about ten yards away. The ap- parent population reduction probably resulted from the aggregating of the insects. While several other lygaeid species, Emblethis vicarius , Peritrechus fraternus, and Cnemodus mavortius were found hiberating within the clumps of Andropogon, Sisamnes was found in the much drier and semi-open intervals between the clumps. Unfortunately this population and site could not be further studied for the habitat Avas destroyed in late November when this portion of the sand pit was bulldozed. Although a single male specimen was found on the fringe of the area in the following May, despite continued search the population seemed to be elimi- nated until two years later in August, 1961 when a small colony was found about 125 yards from the original site. This was a much newer habitat although in general similar to the original site in soil and exposure. The vegetation consisted of a ground cover of Potentilla canadensis L. and a few scattered young clumps of Andropogon scoparius and some scattered St. Johnswort ( Hypericum ) and a few thin grasses. The ruderal aspect of this association were also indicated by the presence of Poa pratensis and Ligyrocoris diffusus Uhl. The biotope of Sisamnes was in the ground litter at the base of the plants. In this habitat the abundance of the insect was only 2-3 per square meter over a small ten square meter area. As this biotope was adjacent to open sandy biotopes with sparse litter, the litter temperatures were compared. At 1 :30 on August 24 the sandy areas where Emblethis vicarius and Geocoris bullatus occurred had litter temperatures of 45° C., while the temperature of the Sisamnes biotopes was 35° C. Unfortunately, once again, the habitat was destroyed in the follow- ing autumn. In the original habitat, Sisamnes clavigera occurred with four other chiefly brachypterous species, Geocoris limbatus , Cnemodus mavortius, Pseudo cnemodus canadensis, and Carpilis consimilis, of which the last two were much less abundant. This species is evidently rare in Connecticut. Many similar habitats were searched with no success. Along with its ecological restriction and rareness, this small insect possesses a remarkable camouflaging coloration which would render it much less frequently collected. The apparent preference of this species for dry sparse 42 ENTOMOLOGICA AMERICANA habitats suggests that the center of distribution of Sisamnes is in the drier western areas such as, perhaps Colorado, from where it was first described by Uhler (1895) . At the United States National Museum nearly all the specimens are from the western United States and include a few macropterous individuals. General Biology S. clavigera is almost entirely brachypterous and although Barber (1953) stated that the macropterous form exists, all speci- mens collected and reared in Connecticut were uniformly brachyp- terous without a trace of the membrane. Unlike Carpilis , high laboratory densities did not promote the appearance of the macrop- terous form, nor did mass rearings at the relatively high tempera- ture of 35° C. The rareness of this species in Connecticut is probably in part the consequence of its poor dispersal powers in uti- lizing the discontinuous mosaic of favorable but temporary habitats. The adult of this small, short legged species is entirely gray with a pattern of pale blotches. It blends remarkably well with the background coloration of the sparse dry litter of the area where it was collected. Indeed in the laboratory on a substrate from the natural habitat it often could not be seen until disturbed. In correlation with this effective camouflage it usually takes a great deal of agitation to induce Sisamnes to move about. When thus stimulated it only runs to the next concealing crevice and again becomes quiescent. It does not, however, exhibit a death feigning response. When given a choice between various kinds of substrate it invariably comes to rest on sand with bits of litter, often con- cealing itself beneath small leaf particles. The later instar nymphs are, in contrast, apparently apose- matically colored a contrasting red and white, in a pattern quite similar to that of Ptochiomera nodosa Say. No parasites were reared, although other lygaeid species present in the same habitat were heavily parasitized. In the laboratory, S. clavigera is easily reared on sunflower seeds. A colony was kept through four generations before it died out. Of the seeds available in its habitat, Sisamnes fed readily on Andropo- gon scoparius seeds and also fed on Hypericum and Potentilla seeds. If the seeds are placed in an exposed position as on methyl cellulose, they are dragged under the litter or substrate by the in- sects. In the supposedly congeneric Sisamnes contractus from Florida, an aggressor releases a territoriality response from a de- fender S. contractus which is very similar to the behavior observed in Pachybrachius where the forelegs are held sideways and the in- sect rears up on its hind legs. The antennae also wag alternately 43 ENTOMOLOGICA AMERICANA very rapidly. While no such elaborate display was observed in 8. clavigera, a similar annoyance waggle response is released from the defender when another insect attempts to feed on a seed “in possession” of the defender. This behavior was also observed in the fifth instars. The sexes do not appear to recognize each other until contact is made. Copulation occurs only after a considerable period (at least several hours) together and after many attempts by a male. When the male becomes excited he taps the female lightly and rapidly with his antennae and presently climbs upon her and at- tempts to mate with her. If the female is receptive she becomes passive with her antennae held low close to the substrate and she release her ovipositor. If she is not receptive, the antennae are alternately wagged rapidly back and forth at a high angle. Life History While I do not have definite field evidence on whether this species has one or two generations a year, there is enough supporting evidence to indicate that there are very probably two generations a year. The observed phenology at Storrs is as in Table 26. TABLE 26 Phenology of Sisamnes clavigera Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult Aug. 26 8% 15% 47% 15% 15% Sept. 16 Oct. 14 Nov. 1 May 16 10% 4% 45% 20% 45% 76% 100% 100% It may be added that Froeschner (1944) in Missouri found adults on April 22, July 14, and September 18, and a fifth instar Septem- ber 24. All the adult females when collected in the field were in repro- ductive quiescence or diapause. Unfortunately the few adults collected in August and May were all males. Dissection of several females from the field in October showed immature, non-active ovaries. Furthermore when two replicates were reared in warmth under autumn (normal) short daylight conditions no reproductive activity occurred, and the adults died without copulating or laying 44 ENTOMOLOGICA AMERICANA any eggs (40-74 days). In contrast, those reared under long day laboratory illumination (over 15 hours) began ovipositing (in 4 replicates) between 18 and 21 days. This is only slightly longer than the normal preoviposition period of 13.6 days (12—15 days). Unfortunately, it must be emphasized that the role of cold was not taken up. Those reared under normal daylight had not been exposed to cold. Those reared under long daylight had been ex- posed to the considerably cooler temperatures of October 14 and after. Therefore it is not known whether long photoperiod or cold was the releasing factor. The species may be reared continuously under long photoperiod conditions. It is probable that short photoperiods bring about reproductive diapause. From the short preoviposition period, the rapid release from diapause and the developmental period in the laboratory, it would appear that ample time is available for two generations a year, especially as the apparent second generation develops so late in the year that fifth instars appear after August 26. The laboratory stadia in days are as given in Table 27. The salient aspect is the relatively slow egg development. This development period is long enough to explain the late appearance of nymphs of the second generation. If it may be assumed that only the adults can overwinter at the northern limits of the species, it is probable that the northern extent of the distribution of 8. clavigera may be determined by a warm season long enough to allow it to pass through two generations successfully. The mean adult longevity in the laboratory is 90 days and varies from 41 to 148 days. TABLE 27 Stadia of Sisamnes clavigera Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 16.3 6.3 6.5 7.8 7.3 8.6 53 (14-22) (5-7) (5-8) (6-11) (5-9) (6-12) (41-69) As already mentioned, the preoviposition period is 13.6 days with a variation of 12 to 15 days. As these insects are slow to react sexually, it is difficult to determine the precopulatory period. When the male and females were left together it was clearly less than 13 days, but when daily two hour trials of introduction of males to virgin females were used, frequently the females would lay 45 ENTOMOLOGICA AMERICANA a few infertile eggs before copulation. That is, the precopulatory period appeared longer than the preoviposition period. Perhaps a more sensitive test is the male’s response to a female. Twice, a male made no recognition response to a virgin female less than 6 days old but became excited when exposed to a 7 day old female. Egg productivity is 3-4 eggs per day with mean total produc- tivity of 130 and varies from 78-235 eggs. These figures appear low compared to the longevity of the insects and the number of eggs laid in small cultures. Virgin females in this species lay very few eggs (2-6) , or none, in their life span. While the slender eggs are not beset with hairs they are adhesive when first laid and become closely covered with dust and sand grains which effectively conceal the eggs. The eggs are laid singly mainly into the soil and crevices. Very few are laid in methyl cellulose or wet cotton. Carpilis consimilis Barber The genus Carpilis, with three known species, is entirely Nearc- tic in distribution. The type species ferruginea was described by Stal from Texas and later recorded by Uhler (1876, 1894) from New Mexico and Lower California. It was not until 1953 that Barber discerned that the northeastern population represented a separate species, consimilis. The third species, barberi, was described by Blatchley (1926) from Florida. The northeastern species, C. con- similis, has a relatively restricted known range from southern Que- bec south to Long Island (New York) and northern New Jersey. As comparable ecological conditions exist along the Appalachians it seems entirely probable that consimilis will eventually be found as far south as the grassy mountain balds of the southern Appala- chians. Nevertheless, the three known species are apparently wide- ly separated from one another. It may be that the quite similar eastern and western species evolved from populations isolated dur- ing Pleistocene times. The third species, barberi, is endemic to Florida, but is not the usual Florida endemic (c.f. Hubbell 1961) for it apparently occurs in south, not north, Florida. It should be added that Carpilis is very closely related to Pry fanes Dist. (Bar- ber 1954) which includes a Cuban species along with two described by Distant (1993) from Central America. Environment As is generally true of brachypterous species, populations of Carpilis usually persist in the same locations from year to year, and 46 ENTOMOLOGICA AMERICANA do not readily invade new pioneer habitats. Nevertheless, most of the habitats where consimilis was collected were basically open tem- porary habitats eventually to yield in succession to forest vegetation. Therefore, it would appear that this species violates the restriction to climax habitats of brachypterous species (c.f. Southwood 1962). The important element, however, is the time factor involved; Carpilis is restricted to environments which become invaded by woody plants relatively slowly because of the overdrained nature of the soil. These environments may be considered to be of two types. One includes overdrained xeric slopes, usually morainic structures, which undergo succession relatively slowly. The other consists of dry wind-swept mountain balds where the available soil is much too thin and dry to give sustenance to any but scattered trees. This is a very long lasting habitat, which should be considered a ser- climax somewhat resembling a savanna woodland (Dansereau 1957). Such serclimax habitats were found on Canaan Mountain, Con- necticut and Mt. Greylock, Massachusetts. The scattered, stunted trees of these habitats are predominantly of Quercus ilicifolia Wang, Q.. prinus L., with some dwarfed white pine ( Pinus strobus L.), quaking aspen ( Populus tremuloides Michx.) and low shrubs of Vaccinium, angustifolium Ait., V . vacillans Kalm, Gaylussacia baccata (Wang.), and Aronia melanocarpa (Michx.) are present. The biotope of Carpilis consimilis in this habitat is in the litter beneath the low plants around and in the crevices of the exposed rock outcrops. The low vegetation consists of Vaccinium angustifolium Ait., the graminoids Festuca rubra L., F. capillata Lam., Danthonia com- pressa Augt., Panicum spp., Bromus kalmii Grey, Juncus greeni 0. and T., a few forbs such as Solidago squamosa Muhl., and some coarse mosses, mostly Polytrichum piliferum, and Cladonia lichens. Carpilis is often found in this environment with several other lygaeids especially Xestocoris nitens and Lygaeospilis tripunctatus (Dallas) but it is the least abundant of these, and occurs in abundances of 1-4 per square meter. The other habitat type is found on dry overdrained morainic sites which support a definitely pioneer type vegetation of sparse clump-forming grasses with semi-open interstices among the clumps. Such overdrained sites with poor gravely to sandy soils are usually found on slopes of eskers and driunlins. With such conditions, the ground litter is quite dry (2-3) . It was noted repeatedly that Carpilis is rarely found on very hot exposed slopes, but more frequently on the cooler, but dry, north facing slopes and in the temperature gradient of ecotones 47 ENTOMOLOGICA AMERICANA and margins of occasional trees and shrubs which were establishing themselves on south exposed sites. As this old field type is particu- larly favorable for the ecesis of Pinus strobus L., Carpilis is fre- quently found in a particular edaphic zone around this tree. But the relation is apparently only an edaphic one as Carpilis is simi- larly found around Betula populifolia Marsh, B. papyrifera Marsh, Populus tremuloides Michx., Vaccinium corymbosum L., Spirea latifolia (Ait.), Quercus, and others. The herbaceous vegetation of such habitats usually consists of the grasses Festuca rubra L., Festuca elatior L., Danthonia spicata (L.), Panicum sp., and Andropogon scoparius L. which form tussocks, producing interstices with a dry litter of seeds and debris and often with a considerable growth of xeric mosses and lichens. The forbes Veronica officinalis L. and V. serpyllifolia L., Potentilla canadensis L., and Rubrns villosus Ait. are frequently present. These interstices form the biotope of Carpilis. The habitat described by Barber (1928b) at Indian Lake in the Adirondacks seems quite similar. The tempera- ture preferences in the field could be effectively studied at sites where the preferred biotopes was limited to a narrow ecotone. Such an ecotone at Canaan, Connecticut was found between a pine wood lot and a close-cropped poor pasture on a drumlin. The litter temperature readings (average) were: shaded pine forest edge litter, 66° F. ; ecotone biotope favorable to consimilis , 95° F. ; exposed pasture, 120° F. While moranic structures are common throughout southern New England, Carpilis was not found near the Long Island Sound coast and becomes progressively more common northward. This distribution parallels that of white pine in contrast to pitch pine ( Pinus rigida Mill.) which predominates on such sites near the coast. Habitats of this sort are relatively abundant across central New England and northern New York, an area frequently called the white pine region (Bromley 1935), and where Carpilis was collected at Warrensville, New York ; Canaan, Connecticut ; Great Barrington, Massachusetts ; Northampton, Massachusetts ; and other sites. In these temporary but long lasting habitats I have often found the species in abundances of 10-15 per square meter. By no means should this species be considered rare, but rather as an ecologically restricted species whose habitat renders it infrequently collected by conventional means. The northern and especially the western limit of the range of this species are rather poorly known. It was not found in the present study on high cool but also humid elevations at Mount Washington, New Llampshire ; Mt. Greylock, Massachusetts; and Mt. Monadanock, New Hampshire; nor in the relatively moist 48 ENTOMOLOGICA AMERICANA environments of the lowlands of the Green Mountains of New Hampshire and along the coast of Maine. It was, however, col- lected from Maine at Orono by Parshley (1917b) and by Procter (1946) on Mount Desert Island. As the species was collected in southern Quebec by Provancher (1886) and Moore (1944), it would appear that the species might be found west of Michigan and central Canada along the southern margin of the Laurentian shield where equivalent ecological conditions occur (Monroe 1956). Of especial interest is that Torre-Bueno (1912) collected it in a marsh by sweeping at Yaphank, Long Island, New York, for I have never collected the species in or near a marsh, or by sweeping, or in the pitch pine community which dominates on Long Island. Despite the great predominance of brachypterous forms, this species must evidently disperse enough to invade and persist in the dry old field habitats that undergo succession rather slowly. Eventually, as I was able to observe in several habitats, sucession does proceed and eliminate the Carpilis colonies. General Biology In this small species the macropterous form is very rare, and most field populations yield entirely brachypterous samples. Only two macropters were found in five years of fairly intensive collect- ing. It is therefore interesting that there is partial evidence suggesting that the occurrence of the macropterous form is related to high densities. In 41 cultures of low density rearing, all progeny were brachypterous both when reared from field nymphs or overwintered eggs, except in the one relatively high density cul- ture reared from overwintered eggs of brachypterous parents. In the culture, of 25 insects, 3 (two male, one female) or 12% were macropters. This hypothesis would seem supported by the occur- rence of the two field macropters among an exceptionally dense field population (Slater in lift.). However this may be simply due to chance in large populations. Rearing C. consimilis under differ- ent temperature regimes (15° to 30° C) at low densities did not produce any macropterous forms (see discussion of wing poly- morphism) . The habitus of Carpilis is rather unlike that of most myodochines as it is a short legged, sublitter inhabiting species which moves relatively slowly in contrast to the many long legged myodochines. The nymphs are protectively colored dark to light grey for con- cealment, and are not ant mimics. The darkness of the abdominal cuticle varies with the sizes of the abdomen, becoming lighter in swollen nymphs about to molt. In the first instar, however, the abdomen is yellow with a thin red transverse band like other 49 ENTOMOLOGICA AMERICANA myodocliines. The adults might seem to be conspicuous in nature as their white hemelytra contrast with the otherwise black body, but in the light grass, seed litter, and debris, the insects appear to blend in readily when quiet. No parasites have been recovered in insects captured in the field, nor have any predators been observed. Carpilis feeds on many available seeds in its habitats. It feeds on Betula, Panicum sp., and Festuca seeds, but does not feed on pine seeds, nor on Andropogon scoparius seeds. It was reared on sun- flower seeds (Sweet 1960) and Betula populifolia and Veronica spp. seeds. Survival from the egg is much enhanced with the in- clusion of Veronica seeds, while a much higher mortality results when it is fed on sunflower seeds alone. The species survives only about 6-12 hours without water. In their relatively dry habitats water is presumbly acquired from plants and dew. Carpilis , like Ptochiomera and Sisamnes , uses its antennae to raise the body in the righting response, which may suggest the functional significance of the short stout antennae. No seed terri- toriality or any brandishing of the fore femora was observed. Betula , Panicum , and Veronica seeds are transported from ex- posed positions to sites under debris or methyl cellulose. The seeds are dragged by the labium and several times when a seed was caught in the litter, the fore legs were used to dislodge it, but whether the tarsi or femora were used could not be determined. In the males the fore tibia is armed with a strong spine but no function could be discerned for it. It may have some mating function. At no time are the males highly reactive to the females and there is no pronounced courtship. The female responds to the presence of a male by rapidly “ wagging” her antennae. When very receptive, she sometimes actively advances on the male. The male climbs upon the female from any angle, grips her firmly with his fore tarsi and presses the apices of his antennae against her head as she continues to wag her antennae. If receptive, copulation ensues ; if not, the male is dislodged by the female with rapid, convulsive side to side jerks of her body. Copulation lasts from three-quarters of an hour to four and a half hours during which time the pair usually conceals itself under the debris. The coiled vesica of the aedeagus is very long (Ashlock 1957). Never- theless, it entirely enters the equally long spermathecal duct. In separating, the insects walk away from one another, pulling at the substrate. Because the straightened vesica is three times as long as the male’s body it frequently becomes caught in debris especially when a pair is startled while separating. Perhaps because of its great length the vesica does not snap back into place as is usual, 50 ENTOMOLOGICA AMERICANA but recoils slowly. The male tilts forward upon antennae and fore and middle legs, and uses the hind tarsi to help settle the coils back into place. Later the male may mate again. Sometimes, however, the aedeagus does not recoil and it is dragged along behind the male and eventually becomes entangled which results in the death of the insect. Life History Carpilis consimilis has a univoltine life cycle and overwinters as an egg with a strong obligative diapause. Evidently, the life cycle of the species is adapted to a cold temperate climate with a rela- tively short summer. The entire life cycle is passed in the same habitat. Because of the relatively infrequent occurrence of Carpilis around Storrs, Connecticut, only a rough phenological outline can be drawn. Table 28 represents a summary of data from Canaan, Connecticut. Adults were observed copulating in the field as early as July 24 in Canaan. It appears evident, as compared with other egg-diapause species, that eclosion in Carpilis occurs relatively late in spring. This probably results from the early onset of diapause in egg develop- ment, which necessitates that egg development occur largely during spring after the ground temperatures have risen. Under the warmer laboratory conditions such eggs hatch much earlier in the spring and the life cycle is then accelerated. TABLE 28 Phenology of Carpilis consimilis Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult J une 1 33% 67% June 10 60% 40% — June 13 10% 90% June 21 50% 50% — July 1 100% July 7 50% 50% July 12 — 33% 67% July 18 9% 86% 5% July 26 15% 85% Aug. 5 7% 93% Aug. 18 100% 51 ENTOMOLOGICA AMERICANA The eggs of C. consimilis possess a strong diapause as none of the eggs develop spontaneously under room temperatures. While the eggs may survive at least - 15° C. and obviously much colder temperatures in the field, only moderate cold temperatures are required to break diapause. A detailed analysis of diapause re- quirements was not attempted, but the following conditions were noted. Diapause was broken at 0° C. and 12° C. with cold ex- posures of 199 to 250 days. Shorter cold periods of 28 to 127 days were unsuccessful except for one of 51 days where 9% of the eggs broke diapause, but only after abnormally long developmental periods of 38 to 50 days as compared with the normal (average) egg development period of 17 days. There is no pronounced reproductive pause, and oviposition follows directly after maturity is reached. It appears conceiv- able that the austral limits of the species are in part determined by the ability of the eggs to survive prolonged exposure to warmth, for the longer the warm season, the sooner the adults mature and begin egg-laying. The crucial question then becomes whether enough eggs will survive to ensure the next annual generation. Of course the cold requirement must be met, but selection may probably vary this need. The obligative diapause, univoltine seasonal cycle relationship is more demanding. Several tests were made at 60 and 90 day pre-cold periods with cold periods of 80, 127, and 199 days. None of these eggs hatched, and the best survival when the eggs were placed in cold shortly (2-20 days) after being laid. In fact, many of the old eggs died in room conditions before being placed in cold. Eventually all die if left too long in warmth. This relationship then very well might form the major factor limiting the austral distribution of this species. Clearly this situation warrants more detailed study. Under laboratory conditions the stadia in days are as in Table 29. TABLE 29 Stadia of Carpilis consimilis Egg* Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 17.5 7.0 8.3 9.0 7.6 12.1 61.5 (6-26) (7) (5-12) (6-13) (5-12) (7-23) (48-83) * period after return to warmth 52 ENTOMOLOGICA AMERICANA As usual, development occurs more rapidly in the laboratory than in field populations. No significant difference was found in the longevity of males and females or whether mated or not. The average longevity was 54.4 days and varied from 23-65 days. Some field adults survive in the laboratory until early Decem- ber, but the majority die during October. The precopulatory period of the male and female is not pre- cisely determined but is less than nine days. The preoviposition period is eleven days. Egg productivity is markedly less than in other rhyparochro- mines and varies from 40-147 eggs (mean, 70 eggs) in mated fe- males. The variation results from differing longevity, as on a per day basis the productivity ranged from 2.2 to 3.3 (mean 2.6). Sexually isolated females lay from none to 21 eggs. Such females become very swollen with eggs. Withdrawal of seeds caused a cessation of reproduction, so oviposition in the field must be also related to the availability of seeds. When a substrate for oviposi- tion was not provided, egg production was greatly curtailed, and two fertile females laid only 26 and 38 eggs. The smooth narrow somewhat pointed eggs are laid singly deep into the soil, or into crevices as the axils of seeds of Veronica, in hollow stems, often pressed into soft pith, into heavy tomentosity and other plant debris. In the laboratory Carpilis also oviposits into methyl cellulose, and cotton both wet and dry, but prefers dry substrates. The eggs do not cling to the substrate as in many other rhyparochromines. Considerable time and care is spent by the fe- male selecting individual oviposition sites. Such care may be re- lated to the long diapause state of the eggs, for the eggs must be laid in safe locations. Exptochiomera nana Barber This unique and apparently very rare species was described from a damaged specimen which was collected December 13, 1913 by Mr. C. A. Frost in Framingham, Massachusetts. Mr. Frost indicated (in lift.) that he had indeed collected this specimen at Framingham, but discussed at length another new species Blissus breviusculus Barb, which together with Exptochio- mera nana he had sent to Dr. Walther Horn in Germany, who had then sent the specimens on to Mr. Barber. Unfortunately, at this juncture, Mr. Frost died, before I could take further advantage of this eminent coleopterist ’s clear memory. Despite continued search at Mr. Barber’s urging, Mr. Frost had been unable to obtain any 53 ENTOMOLOGICA AMERICANA other specimens. Among the Myodochini this species is unusual because the an- terior lobe of the pronotum, instead of being smoothly rounded, is delicately carinate. The sole known specimen is macropterous. Barber expressed some doubt whether this species actually belonged in Exptochiomera. Slater (in litt.) has examined the type and remarks that it may be related to the oriental genus Suffenus Dist. Even whether it is a myodochine is open to question as the spiracle positions are not known. It is a possibility then that this species may be adventive in Massachusetts, perhaps introduced from the Orient or possibly from the Neotropical region where the large genus Exptochiomera is con- centrated. Its occurrence in the field in December, however, would seem to indicate that at least some adaptation to cold was present. Kolenetrus plenus Distant As it is presently understood, this species and monotypic genus has a discontinuous known distribution in the mountainous areas of North America. It was originally described from the mountains of Guatemala, and has been recorded from the Huachuca Mountains of Arizona, northern New York, New England (Barber 1918b), Quebec (Moore 1950), Ontario (Criddle 1922), and recently from British Columbia (Scudder 1961). Father C. V. Reichart (in litt.) has collected the species at Chateau, Montana. I have collected this species from the Great Smoky Mountains in North Carolina. However, a species complex may be involved. I have examined material collected in Guatemala (by Champion), from Arizona, Utah, and from eastern North America, and these populations although certainly closely related, appear quite different, and the whole complex should be carefully restudied. As noted by Distant (1893) K. plenus males from Guatemala have black femora, tibial bases, and first antennal segments and are brachypterous with the membrane barely reaching tergum seven. In the eastern popula- tions the sexes are dimorphic in color, much as is Trapezonotus. In the male, the black areas mentioned are yellow-brown (ochrace- ous) in color, and the membrane almost reaches the apex of the abdomen. In the female these black areas are distinctly infuscated, but not as dark in the Guatemalan males. Moreover, the propleural punctures are fewer and larger in the Guatemalan specimens. It is apparent, then, that this discussion may refer to quite another species or subspecies than Distant’s plenus. In the northeastern United States this species becomes more 54 ENTOMOLOGICA AMERICANA common northward toward the mountainous areas, especially in the Adirondacks. Its distribution, however, is very spotty, and I consider it one of the more uncommon species in New England. Environment In New England this species is ecologically quite restricted and is predominantly found in relatively xeric, but cool sites. Since the more boreal habitats are also often more moist, this species was infrequently found in northern New England. In this respect its distribution parallels Carpilis, but it is even less frequently found. However, it does occur in northern New England. Barber (1918b) records it from Mt. Washington, New Hampshire, and Moore collected it on Peaks Island, Maine, in Portland Harbor Bay (Parshley 1920). It was found most abundantly at a site near Warrensburg, New York, in the southern Adirondacks which may combine the favorable ecological aspects of other sites where it occurred less abundantly. This was an old dry field partly invaded by white pine. Along one side of the field ran an esker moraine which had only a thin scattered vegetation of the grass Andropogon scoparius along with a few small pines which had been planted. There were considerable interspace areas of bare dry gravelly ground between the grass clumps. Clearly this was a pioneer type habitat, but judging from the planted pines and the accumulation of Cladonia lichens and litter around the clumps, the habitat had persisted for a long time. At one side a clump of Spirea latifolia (Ait.) had established itself, and part of the clump had been cut down and piled by the landowner. In this habitat the biotope of Kolenetrus was in the thin layer of pine needles mixed with a short grass which formed a fringe or ecotone between the white pine dense litter and the open areas. It was also found in the shade of the dry pile of Spirea stems and at the margin of a clump of Betula populifolia. Here it coexisted with Carpilis consimilis, Xestocoris nitens, and Trapezonotus arenarius. In this habitat its abundance reached as high as 15 per square meter. It was found about Storrs in far fewer numbers, less than 1-2 per square meter, in dry old north facing fescue fields slowly being invaded by woody plants such as sweet fern ( Comptonia perigrina (L.) ), Vaccinium pennsylvanium, and Quercns spp. It is perhaps significant that this species although it was frequently found with Carpilis, was never collected in bald-type mountain habitats except for one specimen collected at 5,000 feet on the edge of Andrew’s Bald, Clingman’s Dome, North Carolina. On this bald, which was dom- inated by sedges and fescue grasses, it was found on the margin of 55 ENTOMOLOGICA AMERICANA Rhododendron catawhiense E.L.Br. litter. At the Great Smoky National Park it was also collected along a gravelly roadside near Snco Bald at 4,500 feet. The essential requirements of this species would appear to be xeric (2-3) habitat which is cool and partly shaded, as an old field ecotone, a combination which was not commonly found. These re- quirements in large part explain the scarcity of this species. Scudder (1961) collected K. plenus in a quite different habitat among J uncus tufts at the edge of Westwick Lake, Cariboo, British Columbia. Scudder further notes that Kolenetrus appears very similar to Acompus which in Europe occurs in this sort of habitat. Torre-Bueno (1922), it should be added, collected Kolenetrus in a bog in Massachusetts. The specimens from Guatemala were collected by Champion from the Quiche Mountains and at Quezaltango between 7,000 and 9,000 feet (Distant 1893). This altitude is above the cloud forest (4,500-7,000 feet) at a temperate level variously dominated by temperate-type coniferous or broad leaved forests (Standley and Steyermark 1946). General Biology All the specimens seen from eastern North America are macrop- terous (female) or submacropterous (male). Actually the male’s wing is proportionally as long as the female’s, but tergum seven is exceptionally long in the male Kolenetrus which causes the wing to appear submacropterous. The specimens appear capable of flight. According to Parshley (1917) this species was freshly washed up in ocean litter on Beach Bluff, Massachusetts, and be- cause of the on shore wind he thought it clearly had to be flying over the water at that time. K. plenus appears to possess disruptive coloration, with a clear contrast between the black-bronzed head and pronotum, and the half pale, half fuscous hemelytra ; but when moving it is quite conspicuous. The nymphs have the abdomen cross banded in a pattern similar to that of Stygnocoris pedestris, which is frequently found in similar habitats. No predators or parasites are yet known. The natural food seeds of this species are not known. It does feed readily on sunflower and Betula popidif olia seeds. It feeds very little during the essentially aestival diapause period of late summer which complicated attempts to elucidate feeding prefer- ences. No seed defense behavior was observed. Mating behavior was not observed except that the copulating pairs are secretive and 56 ENTOMOLOGICA AMERICANA take refuge under litter. Observed copulations last from 1 hour 43 minutes to at least 3 hours 30 minutes. Life History The life cycle of this insect is not clearly resolved. It has one generation a year with the phenology in table 30 based on very small samples. TABLE 30 Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 7 June 22 20% 40% 40% 33% 67% July 7 14% 29% 57% July 17 July 28 50% 50% 100% No adults were collected from September 1 on. Therefore the method of overwintering could not be directly observed in the field and correlated with the field populations brought into the labora- tory. The laboratory populations clearly were in an obligative state of reproductive inactivity in late summer in so far as oviposi- tion was concerned. However, on July 27 in the field and on August 5 in the laboratory copulation was observed. But no eggs were laid. When left at room temperatures the insects came out of diapause in late November, laid a few eggs, and died. More- over, these eggs did not go into diapause and most hatched readily. On this basis it would appear that the insect overwinters as an adult as this situation frequently occurs in overwintering adults which may complete diapause in warmth. However, in no other rhyparochrominc species with a single generation and which obligatively overwinter as an adult was copulation observed in the field or laboratory in late summer. But a somewhat similar pre- oviposition copulation period exists in Stygnocoris and Drymus wdiich overwinter as eggs. A replicate of seven Kolenetrus adults were placed in cold (3° C.) for 18 hours on September 23. One male and a female were removed. The other adults were left in cold for a longer time but died. The female after four days laid 14 eggs which were put in the cold room on October 15 and re- moved April 16, 1961 after 183 days. Of the 14 eggs, eleven de- veloped, and one hatched, which at least indicates that the eggs have ENTOMOLOGICA AMERICANA the capacity to overwinter, even if in a quiescent rather than dia- pause state. The best interpretation of this abnormal laboratory data, and the field evidence is that the eggs overwinter and the adults, like Stygnocoris and Drymus need a cold exposure to in- itiate oviposition. The stadia of table 31 are based again on rather limited samples, the earlier instars on laboratory hatched eggs, the later instars on field nymphs reared in the laboratory. None were reared com- pletely through the life cycle. TABLE 31 Stadia of Kolenetrus plenus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 16 9 10? 7 7 8 57? The slow development of the eggs is noteworthy. Longevity of the adults averaged 125 days (range 82-164) in the laboratory. In one female in which the first copulation was ob- served, the eggs were laid after a preoviposition period (?) of eight days. As indicated above, the data on reproduction is quite abnormal. The females which came out of diapause laid few eggs (average 18.5, range 14-21) over 14-20 days. The poor fecundity may be the result of metabolic wastage from keeping the adults in warmth for so long a period, as is the case in Stygnocoris and Drymus. The eggs are smooth and slender and have the anterior ends flattened. Most are laid in the tight crevices in the litter and into methyl cellulose. Several are pressed into the tight wet cotton stopper. 58 ENTOMOLOGICA AMERICANA TRIBE PLINTHISINI Plinthisus americanus (Van Dnzee) P. americanus (V.D.) is here raised from synonymy with P. compactus (Uhler). Barber (1918d) carefully compared the types of P. americanus described by Van Dnzee (1910) from Tyngsboro, Massachusetts and P. compactus described by Uhler (1904) from La Vegas, New Mexico and considered them to be conspecific. They are very similar, but a striking sexual dimorphism suggested that the species were distinct. In the eastern population, while the males have a nearly nude hemelytra, the females have the hemelytra densely investitured with long erect hairs, similar to the pronotum. In the western population, however, both sexes have nearly nude hemelytra. The species can be separated as in the following couplet which may be added to Barber’s 1918 key (1918d). 1. Hemelytra of female nude; male with a nearly feature-less internal surface of tergum seven ; paramere with basal lobe small, nearly pointed ; paramere apex long and slender ; spermathecal duct with only a narrow rim of sclerotization just beneath the bulb ; female with sutures on tergum 8 diverging cephalad and not meeting compactus (Uhler) la. Hemelytra of female densely pilose ; male with a circular apodeme on inner surface of tergum seven which receives genital capsule at rest ; paramere with a large rounded basal lobe ; spermatheca with a broad sclerotized band on the duct just proximal to the bulb ; females with sutures on tergum 8 converging and meeting cephalad americanus (V.D.) The disjunct presence of such closely related species in eastern North America and southwestern United States probably indicates a long isolation (Ross 1962). The other three known American species are recorded from the western United States and Ashlock informs me (in litt.) that ac- tually a large complex of Plinthisus exists there. So far americanus is known only from Quebec and Ontario south to northwestern Connecticut. It will probably be found southward along the Ap- palachians. The southwestern form is also found in Arizona (Banks 1910) and was collected in a Neotoma nest in California (Torre-Bueno 1946). The genus Plinthisus itself is nearly world wide, and the only major zoogeographical region without any recorded species is South America. There is a heavy concentration of species (43 to 62) in the Palearetic (mostly Mediterranean region) but recent work on 59 ENTOMOLOGICA AMERICANA the Ethiopian fauna rather suggests that the distribution of Plin- thisus may more closely parallel the presence of hemipterists. Nevertheless only a few of the species are found in north temperate areas. It is therefore of interest that P. americanus is adapted and restricted to a north temperate climate. Environment P. americanus is a forest species found in the litter biotope beneath hemlock ( Tsuga canadensis ), hemlock-birch ( Betula papy- rifera), spruce-fir ( Picea rubrum-Abies balsamea) forest associa- tions, but rarely in white pine litter. The heaviest densities (20-25 per square meter) are found in Tsuga litter. Litter that is rela- tively loose and on a slightly drier than average slope is more favorable than the tightly packed litter common to coniferous forests. The litter is always a typical mor type and the soil a grey podsol. In favorable biotopes the litter has a mesic moisture level of 4-6. Plinthisus is absent from wet sites with considerable accumulation of mosses, typical of many spruce forests and also is rare in very dry red spruce litter on mountain tops. It is fre- quently found with Eremocoris ferus and S colop ostethus diffidens, especially where a considerable admixture of Betula papyrif era L. is present. The herbacious level, as is typical of coniferous forests, was absent or insignificant. Within the wide spread forests of this type, Plinthisus displays a discontinuous ecological distribution in relation to the variable litter and slope types. In northern New England, it is more abundant in valley forests than in the mountain forests. Its abundance is frequently quite low, 2-4 per square meter, but occasionally up to as much as 20-25 per square meter. General Biology Macropters are very rare in this species, and only two were found in the course of the study. This correlates with the permanence of its climax forest habitat. The only specimen of Plinthisus col- lected by Ashlock {in litt.) was a macropterous male. This becomes significant for it was collected in white pine litter which is appar- ently a rather unfavorable habitat for Plinthisus, and the macrop- ter may represent a dispersing individual. Like all brachypterous species of Plinthisus, P. americanus is a small, oval, shiny brown insect. It is a sublitter form, keeping to cover under the debris, and it usually remains quite still when the litter is disturbed and the insect must be directly disturbed before it moves to seek cover elsewhere. It does not feign death, however. Its color blends well into the background litter and it requires careful search especially where the densities are low. Plinthisus feeds very readily on the fallen seeds of Tsuga 60 ENTOMOLOGICA AMERICANA canadensis , Betida papyrifera, B. populifolia, and Picea rubra. It also feeds readily on sunflower seeds. Like many forest rhyparo- chromine species it can survive only about 4-6 hours in a dry en- closure. Plinthisus actively moves the seeds into interstices in loose litter for feeding*. Since Plinthisus shows a marked sociability in the field and often aggregates in a particular litter clump in the labora- tory, the seeds are often found grouped together. Frequently these litter mounds were bound together with ( ?) mycelia strands. On several occasions Plinthisus was observed moving the large (compared to the insect) Tsuga seeds. It first feeds on the seed and then proceeds to drag the seed with its labium. When the seeds became caught in the litter, the insect places its fore femora over the seed, dislodges the seed, and resumes dragging. When en- closed with Eremocoris ferus and Scolopostethus diffidens, this be- havior becomes much more accentuated, as the Plinthisus endeavors to protect the seeds from the Eremocoris which displays an equal readiness to feed on the Tsuga seeds. In the mating behavior, the male leaps suddenly on the female and apparently senses her without any prior contact. Copulation lasts from 5 to 8 hours and is often repeated. Life History Plinthisus americanus has a univoltine life cycle with an obliga- tive egg diapause. The earliest nymphs were found in mid-June in the third instar, and fifth instars were found as late as July 27. This evidently indicates a late eclosion period, which would correlate with the fool forest floor environment. There apparently is a fairly long preoviposition period of 17-21 days as oviposition begins in early to mid-August. Laboratory conditions, however, may greatly curtail egg production. When hemlock seeds were in- cluded, the fecundities were more uniform and higher. The eggs are laid from late summer until winter intervenes, and diapause in early anatrepsis. I was unable to completely break the strong diapause. However, after six months exposure to mod- erate cold (4° C.) egg development to katatrepsis occurs, but no hatching. Fecundities of from 45-97 eggs per female (mean, 67) are recorded. Four sexually isolated females lay no eggs. The lon- gevity of the adults in the laboratory varies from 30 to 101 days (mean 62). The eggs are laid into the forest litter, pressed into small crevices in the substrate. The eggs are narrow, somewhat pointed on the posterior end, the anterior end is flattened and the sides are slightly fluted. 61 ENTOMOLOGICA AMERICANA TRIBE ANTILLOCORINI Antillocoris minutus (Bergroth) The tribe Antillocorini as recently established by Ashlock (in press) includes a number of genera of minute Lygaeidae. The species of Antillocoris themselves are about 2 mm. in length. The genus includes 6 species, 3 of these occur in Central America or the West Indies and one reaches South America. Many more species will probably be described when the Neotropical fauna is better known. The other three species are found in the eastern United States, each with a different latitudinal distribution (Bar- ber 1952). A. discretus Barb, was recently (Barber 1952) de- scribed from the Gulf Coast area from Florida to Texas with one isolated record from southern New Jersey. A. pilosulus is found throughout the southeastern United States north to New Jersey and Missouri. The third species, A. minutus, is a northern species, occurring from eastern Canada south to New Jersey and Missouri. Southern records of minutus refer to the two southern species (Bar- ber 1952). While the two southern species are entirely macropter- ous, A. minutus is pterygopolymorphic. Environment Antillocoris minutus is a common although infrequently col- lected inhabitant of forest litter. It is often abundant in litter beneath gray birch ( Betula populifolia Marsh) and white birch ( Betula papyrifera Marsh) where it may attain densities of 50-80 per square meter. A. minutus is also found in lesser abundances (up to 20 per square meter) in hemlock ( Tsuga canadensis) litter, and in V accinium-V iburnum- maple litter. In all of these forest habitats it is frequently associated with the rhyparochromines Scolopostethus spp., Eremocoris ferns and Drymus unus (see Com- petition discussion). However, the species was never collected in oak-hickory forests and rarely in very mesophytic deciduous forests. Occasionally it is found in various marginal type habitats but in low abundances (1-5 per square meter) as in vole runs in heavy grass meadows, or at the base of sedge clumps in marshes. In spring after the post- hibernation flight it frequently occurs in various habitats, then apparently suitable, but which dry out later, and the species dis- appears. In the favored biotopes of A. minutus , the litter is well-drained but relatively mesic (moisture 5-8) and the temperatures vary from 71° to 82° F. at midday in July. The litter layer varies from 2 to 10 inches deep and contains many small seeds. Sometimes the 62 ENTOMOLOGICA AMERICANA litter accumulates between clumps of a fine leaved grass. The soil is usually a mull litter on a sandy loam or in hemlock forests, a grey podsol. In nearly all habitats its ground layer biotope is com- pletely shaded from direct illumination. In southern Connecticut A. minutus occurs frequently on cool north-facing slopes which are also favorable to Betula- hemlock communities. The few habitat notes in the literature largely confirm a wood- land habitat selection. Torre-Bueno (1924, 1929a) sifted it from moss in dense woodlands, and from sphagnum and leaves near a swamp edge (Torre-Bueno 1925). Barber (1923) says it is a com- mon species among dead leaves in damp situations. A. minutus occurred at several forest sites for at least five years in succession and overwintered in these same habitats. General Biology In marginal habitats only macropterous specimens are collected. These probably result from post-hibernation migration flights. In favorable habitats where it occurs year after year the large majority of the population is braehypterous. At one area for two years in succession the percent of macropters diminished sharply from early spring to late summer. There are no good records of A. minutus at lights. A. minutus is unique among New England rhyparochromines in possessing two distinct braehypterous forms. In one, the bra- chypter, the hemelytra are merely abbreviated as is usual, but in the other, the subbrachypter, the membrane is absent and the margin of the corium is truncated. Since the species is univoltine, little could be done to resolve the nature of this interesting wing poly- morphism. From one large population with an exceptional number of macropters present the spring females were isolated. The fe- males were already mated and only the resultant progeny were examined. The results are interesting as shown in. Table 32. TABLE 32 The wing development of the progeny of Antillocoris females Parental Females Progeny 5 subbrachypters 134 subbrachypters 4 brachypters 120 brachypters 8 macropters 191 brachypters 1 macropter 37 brachypters, 8 macropters 63 ENTOMOLOGICA AMERICANA Since this is the same population, random mating was probable. The data on the progeny permit a few deductions. First, that the laboratory environment did not affect the appearance of the subbrachypterous phenotype since the conditions were nearly iden- tical. But all the macropterous females but one yielded only bra- chypters and the brachypters, only brachypters. The meaning of these data is unclear: a maternal factor may be involved and/or a switch gene for brachyptery and maeroptery as in Gerris (Brink- hurst 1959). This certainly warrants more work. A. minutus is one of the least conspicuous of all the New Eng- land lygaeids and its abundance must be considerably underesti- mated. The adult is dark brown in color and characteristically moves slowly and deliberately like many woodland lygaeids. When touched it runs rapidly for a short distance then suddenly resumes its deliberate gait. Its small size allows minutus to conceal itself in small crevices such as in rolled up birch leaves. The nymphs however are very conspicuous. They are a distinct pale pink color. Both nymphs and adults possess a peculiar scent gland odor resembling Tapinoma ant odor. The nymphs of Antillocoris lack the anterior scent gland. The scent gland secretion is released only under very rough treatment. The predators or parasites of A. minutus are not known. In the laboratory Antillocoris feeds readily on seeds of Betula populifolia , Betula papyrifera, Tsuga canadensis and probably other seeds as its feeds on sunflower seeds. It can be reared very easily on a sunflower-birch seed mixture. No seed defense behavior was observed. Beyond the male vibrating his antennae at the approach of a female, the mating behavior was not observed. Life History A. minutus is univoltine and has an obligative adult diapause. Its seasonal cycle begins in mid- June and is completed by late July as shown in Table 33. Adults collected in late April and early May lay eggs in the laboratory within 2 to 5 days and many adults are already fertilized by this time. Since the nymphs do not appear until much later it is probable that under cool and shaded litter conditions in spring the eggs develop slowly as in Xestocoris and Scolopostethus diffi- dens. A long photoperiod stimulus is apparently not necessary as in Myodocha since A. minutus matures under a 12 hour photo- period. The diapause condition is a strong one and very few indi- viduals complete diapause development under warm conditions. In one culture a female broke diapause after a long dormant period of 254 days but laid only 19 eggs. This is unusual, for most adults 64 VOLUME XLIV live under warm conditions only until December to February and die without reproducing. Photoperiod seems to have no effect on the seasonal cycle and diapause is completed under cold conditions. The stadia given in Table 34 are based on cultures reared on a gray birch — sunflower seed mixture. Poor growth is made on sun- flower seeds alone. TABLE 33 Phenology of Antillocoris minutus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 8 100% June 18 5% 20% 75% June 28 10% 30% 25% 5% 30% July 9 10% 20% 40% 20% 5% 5% July 18 25% 45% 30% July 26 16% 42% 42% Aug. 13 3% 7% 90% Aug. 26 2% 98% Bept. 10 100% TABLE 34 Stadia of Antillocoris minutus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 11.0 7.0 6.0 7.1 7.8 10.4 53.0 (9-15) (5-10) (5-8) (6-9) (7-10) (9-12) (46-62) The egg productivity of field collected adults ranges from 34 tn 158 eggs with an average of 94 eggs per female and the eggs are laid at a rate of 2 to 6 a day. Eight spring collected virgin females laid no eggs whatever. At least one field female was non-reproduc- tive as late as June 7, and oviposited only after fertilization. The eggs are oviposited singly in wet substrates such as the stopper of the water vial or damp litter. The eggs are smooth, unadorned, rather short and thick, and slightly curved on the ventral side. They do not stick to the substrate. 65 ENTOMOLOGICA AMERICANA Antillocoris pilosulus (Stal) This species reaches the northern limit of its range in southern New England and Barber (1952) cited it as only extending north to southern New Jersey. However Barber (1923) recorded it from Massachusetts. The species of Antillocoris are similar, and litera- ture records must be used with care. I was able to collect only 3 specimens of what are apparently A. pilosulus in Connecticut. These were collected July 28, 1957 at Noank, Connecticut which is on the Long Island Sound coast. The habitat was an open area heavily invaded by shrubs. The precise biotope was in thick litter beneath the bayberry Myrica pensyl- vanica Loisel. In the Great Smoky Mountains of North Carolina, 8 adults were collected on June 6 and 7, 1960. Three specimens were found in the litter of the shrub, Rhododendron catawbiense at 5,600 feet at Andrew’s Bald, a grassy bald on Clingman’s Dome, North Caro- lina. The other five specimens were collected in leaf litter along a mesophytic woodland edge in the valley at 2,000 feet near Chero- kee, North Carolina. These habitats were relatively moist and shaded much like the habitat of Antillocoris minutus. In the literature A. pilosulus was swept from grasses in a dry cranberry bog in New Jersey (Torre-Bueno 1912) and Blatchley (1926) collected it from beneath stones in a pasture in Indiana. Wray and Brimley (1943) found it in the pitcher plant ( Sarracenia purpurea L.). It has been collected at lights in Missouri (Froesch- ner 1944) and Glick (1939) found what was probably A. pilosulus up to 5,000 feet in the air over Louisiana. This aerial collection included two nymphs so the occurrence might be accidental due to strong winds and the insect’s small size. The adults collected in Connecticut in late July, like minutus, were in a state of dispause, did not become reproductive in the laboratory and died in October. The insects feed readily on sunflower seeds. The adults from North Carolina also fed on sunflower seeds and two surviving females laid a total of 37 eggs. Only 2 adults, a male and a female, were reared through and the did not become reproductive. In southern Florida at Naranja, A. discretus Barb, was collected In mesic leaf litter under an old overgrown avocado tree. This in- dicates a considerable similarity in the general habitat of these three species of Antillocoris. 66 VOLUME XLIV TRIBE LETHAEINI Cryphula trimaculata (Dist.) According to Scudder (1962) the correct name of the single species of Cryphula in northeastern North America is trimaculata (Dist.) not par allelo gramma Stal. C. par allelo gramma, the type of the genus, is a small dark species known from Texas and Arizona. C. trimaculata was described by Distant (1882) from specimens collected in the highlands of Guatamala. If these widely separated populations represent one and the same species, trimaculata has a wide distribution from southern New England west to Colorado and Texas and south to Guatemala. However it should be under- stood that this discussion may refer to a new species (or subspecies) closely related to Cryphula trimaculata of Guatemala since a Guate- malan male specimen before me differs from New England speci- mens in size, puncturation and color. The genus Cryphula includes seven described species of which five occur in the United States and three in South America as well as Central America. However specimens from various collections indicate that species of Cryphula range throughout the Neotropical Region and this genus is quite probably Neotropical in origin. In New England I find Cryphula limited to the southern portion from Cape Cod to southwestern Connecticut. Although it is rela- tively abundant at Storrs, Connecticut I could not find the species in northwestern Connecticut and northward. Environment Cryphula trimaculata is a ground litter inhabitant of long lasting old fields vegetated with perennial bunch grasses such as Andropogon scoparius, Festuca rubra L., Panicum spp., a plant association described under Cnemodus mavortius with which Cryph- ula frequently occurs. Its habitat range extends to woodland margins and grassy glades in relatively xerophytic oak-hickory forests, but it is not a woodland species like Ozophora picturata and it nearly always occurs on the field side of woodland ecotones. This plant association dominates exposed areas on dry over- drained gravelly slopes for considerable periods of time. The bio- type of Cryphula is in ground litter at the bases and in the inter- stices between the grass clumps. The surface soil is usually quite dry (1-2) sometimes, in edge areas relatively mesic (4—6). The biotope temperatures at midday in August range from around 28° to 35° C. much less than fully exposed soil nearby (excess of 50° C.). 67 ENTOMOLOGICA AMERICANA The abundance of the species where it occurs is usually about 6-10 per square meter and in favorable habitats under Panicum sp. it reaches 30-40 per square meter. Blatchley (1926) reported sweeping adults and nymphs of this species from the flowers of redhaw ( Crataegus punctata Jaeq.). Barber (1923) noted Cryphula as another species found in sifting dead leaves or beneath stones and sticks on the ground. Barber (1918c) also collected C. abortiva Barber and C. nitens by shifting among dead leaves. In general, Cryphula trimaculata appears ecologically similar to the other lethaeine species, Xestocoris nitens. While Xestocoris also is an inhabitant of semi-permanent grass habitats, it is rarely found with Cryphula. Cryphula may ecologically replace Xesto- coris in southern New England where, perhaps in competitive exclu- sion, Xestocoris occurs on north-facing slopes on dry Festuca habi- tats as described under Carpilis. Cryphula, on the other hand, is much more frequently found on south-facing slopes, and more mesic margins of woodlands than is Xestocoris. And perhaps as a consequence of this, Cryphula is frequently found with Pachy- brachius basalis under Panicum spp. in mesic habitats where Xesto- coris is rarely found. More work is needed both to the north and to the south to see whether these species are really ecological equivalents and if any character (ecological) displacement is occurring where the ranges of the species overlap. With Xestocoris, the ecological amplitude appears greater in more northern habitats than at Storrs, Connecti- cut but this may simply reflect more restricted microclimatic condi- tions in southern New England. In southern Florida at Naranja, I collected a small rather in- fuscated form of this species in short grasses in a palmetto — slash pine habitat. The general aspect of this open association is similar to some habitats of Cryphula in southern New England. General Biology Like Xestocoris, Cryphula trimaculata is wing polymorphic and roughly 10% of the field populations are macropterous, the per- centage varying from one population to another, but with the brachypter always the much more abundant form. This wing con- dition correlates with the relatively long-lasting dry habitat of this species. The presence of the macropter correlates with the fre- quent occurrence of Cryphula in the scattered mosaic of favorable habitats. There apparently are no dispersal records for this species. However, field observations show a larger proportion of Cryphula along woodland margins in autumn than in midsummer 68 VOLUME XLIV which may indicate some small scale movements to more protected hibernacula. Dowdy (1955) collected this species overwintering in oak-hickory forest in Missouri. Cryphula is a small brownish insect blotched with pale lines and areas. This procryptic coloration blends very well with the dry grass litter background. This insect is short-legged and flattened and does not run far when provoked, but scurries to the nearest crevice to hide. It frequently takes direct contact to provoke the insect, but Cryphula does not feign death. In the laboratory it conceals itself under bits of leaves and avoids white substrates. The function of the remarkable iridescent sheen of the integument and the iridescent patches on the vertex of the head is not under- stood. There is a very low percentage (less than 2%) of parasitism by an undescribed species of Catharosia. The parasite overwinters in the diapausing host. Cryphula feeds readily on sunflower seeds, oviposits abundantly, and can complete its development on this seed, although the mor- tality is high. However, it definitely prefers to feed on seeds of Panicum spp. and is easily reared on this seed. It also feeds on seeds of fescue, millet, and strawberry and probably feeds on other seeds as well. Under hunger stress this species will scavenge on dead and dying Cryphula specimens. The seeds of Panicum are dragged about to sheltered locations for feeding. The seeds are impaled on the labium and dragged about under the insect. When the seed catches on the litter the insect lifts the seeds over the obstacle with the combined action of its fore and middle legs. In its mating behavior, the male approaches a receptive female which it apparently recognizes before actual contact. The anten- nae are alternately moved up and down with a quick flick at the end of each beat. After contacting the female the male jerks his body from side to side with a very brief pause after each movement. He climbs on the female, flicking her head with these antennal movements and accomplishes copulation in an end to end position. It may be relevant that in all eight observed copulations the female was feeding on a seed. Life History Despite its southern distribution, Cryphula has an univoltine seasonal cycle with an obligative adult diapause. Dowdy (1955) noted that this species overwinters as an adult. The nymphs do not appear until rather late in the season at Storrs, Connecticut as shown in Table 35. 69 ENTOMOLOGICA AMERICANA TABLE 35 Phenology of Cryphula trimaculata Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 20 100% July 3 25% 30% 15% 30% July 20 11% 28% 22% n% 11% 17% Aug. 6 4% 16% 32% 48% Aug. 22 7% 7% - 26% 60% Sept. 5 25% 75% Sept. 25 9% 91% Oct. 10 100% As early as May 26 some females are already fertilized. Active mating and oviposition were observed in field popnlations at Storrs, Connecticut on June 18 and 19, 1962. To test for photoperiod response a population of early spring adults not yet reproductive was divided into two groups and cul- tured under photoperiods of 12 and 15 hours. Under both long and short photoperiods the cultures became reproductively active at the same time. Moreover the progeny of each culture went into reproductive diapause, showing the obligative nature of the dia- pause. The duration of the diapause state was also not affected by these photoperiodic conditions. Most cultures which are kept in the laboratory in autumn under warm conditions do not become reproductive. The few that do are all collections which were exposed to cool field conditions of early November. However, only certain individuals are released from diapause and these after diapause periods varying from 16 to 98 days, after which the ovipositing females live 28 to 42 days. These females lay very few eggs (13-28 eggs) at a rate of less than one a day. The normal fecundity of Cryphula in spring is from 84 to 128 eggs (mean, 98). Adults collected in late April begin ovipositing in about a week. Unmated females lay from none to 41 eggs (mean 26). The longevity of diapausing adults ranges from 97 to 219 days. The stadia which are given in Table 36 are not completely worked out and for the later instars only the range is given. The egg development given is for 80° P. (room temperature) and is relatively slow. Eggs incubated at 67° F. developed even more slowly, taking from 34 to 39 days to hatch. The late occurrence of 70 VOLUME XLIV nymphs is probably best accounted for by prolonged egg develop- ment under spring ground temperatures. The eggs are rather short and thick with a smooth chorion which does not stick to objects or to other eggs. The eggs are oviposited singly in cotton and under bits of litter. TABLE 36 Stadia of Cryphula trimaculata Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 26.3 (16-39) 9.2 9.0 (7-12) (8-10) 9.4 (8-10) (10-12) (10-12) (60-62) Xestocoris nitens (Van Duzee) Xestocoris is a small genus of three species. One, X. collinus Dist. is known from Guatemala and Panama. The second, uhleri (Dist.), is known from Grenada, West Indies. The third, nitens, has a northern distribution from Quebec west to Iowa and south to Massachusetts and New York (Slater, Catalogue), and I have collected the species in the high elevations of the southern Appala- hians in the Great Smoky Mountains National Park, North Caro- lina. X. collinus has on several occasions been intercepted on orchids, Oncidium sp. and Cattleya bouringiana from Panama (Swezey 1945). From the literature, Xestocoris would appear to be an ex- tremely rare species, but again such a “ rareness” is more the re- sult of its habitat choice and ground layer biotope. It was only recently found west of New York in Illinois and Iowa (Slater 1952). Environment Xestocoris is a characteristic species of dry (2-3) overdrained morainic old fields with a long persisting sparse, low vegetation of clump forming grasses such as Andropogon scoparius , Festuca rubra L., F. capillata Lam, and Danthonia spicata (L.). In south- ern New England it is usually found on north-facing exposures. Similarly it is common on mountain balds in the sparse vegetation among rock outcrops. It is frequently found in dry edge habitats between a field and forest, especially on the margin of the dry soil indicator, white pine ( Finns strobus). It is not found in very barren open hot habitats with a considerable amount of bare soil, 71 ENTOMOLOGICA AMERICANA but only where the vegetation forms a closed ground biotope of litter and lichens between the grass clumps. Its abundance in some lowland fields is as much as 25-30 per square meter and on moun- tain sites, 15-20 per square meter, but it is usually found in den- sities of about 5 per square meter. I never found Xestocoris in wet habitats from where it was reported by Barber (1923) and Blatchley (1926). Torre-Bueno (1929a) collected it by sifting grass piles in Massachusetts. It hibernates in situ, and frequently forms large hibernating aggregations at the base of a single grass clump which is no way distinguishable from other neighboring clumps. This tendency to aggregate is also shown in the laboratory for a group of insects will cluster together in one corner of the rearing dish, especially under cool conditions. General Biology The great majority of the specimens are brachypterous which correlates with the semi-permanent habitat perference of Xestocoris. The resultant low dispersal rate, however, must be sufficient to readily colonize such areas. Dispersal apparently occurs in the spring, as is indicated by the nearly complete disappearance of macropters from several habitats through the spring. Movement from the localized hibernacula occurs in early spring and this also actually constitutes an active dispersal movement but not by flight as most individuals are brachypterous, Xestocoris is a small insect with a generalized body shape and short legs. It keeps to cover at the base of grass clumps in the fallen grass litter and lichens. When disturbed the insect’s imme- diate reaction is to conceal itself again. Its dark shiny coloration against the light litter does not appear to be especially procryptic except when it is concealed in dark crevices at the base of the plants. The nymphs are pale pink in color, and the late instars are quite conspicuous in the field. No predators or parasites were found. Its chemical defense should be interesting as this species in both nymphs and adults (like Antillocoris and Crypkula) has a heavy odor very similar to the odor of the ant Tapinoma. There is a possibility that Xesto- coris is avoided by ants as the odor of Tapinoma repels other ants (Wheeler 1910). This is one of the few species which suffers a heavy mortality in the shift from field diet to sunflower seeds. It does not do well on this seed and could only be reared by adding Festuca seeds. It feeds readily on Festuca rubra, F. capillata, Andropogon scoparius, Paspalum muhlenbergia (Nash) and Dantkonia seeds. 72 VOLUME XLIV It does not feed readily on Vaccinium or various composite seeds. While it will feed, if forced by starvation, on non-grass seeds such as sunflower seeds, it strongly prefers grass seeds. Its water needs are large, despite the xeric environment preference and it dies quickly without water. Although the seeds are moved about to safer sites, no seed defense was observed. Nor could the mating behavior be observed beyond a male responding to a receptive female by rapidly vibrat- ing his antennae. Life History Xestocoris nitens has a univoltine seasonal cycle and an obliga- tive adult diapause. As shown in Table 37 the phenology in Con- TABLE 37 Phenology of Xestocoris nitens Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 1 100% June 20 100% July 1 22% 78% July 17 12% 25% 25% 38% July 29 17% 32% 17% 17% 17% Aug. 15 18% 55% 18% 9% Aug. 30 6% 22% 50% 22% Sept. 15 2% 18% 80% Oct. 7 4% 96% Oct. 25 — 100% necticut is late. The early instar nymphs are not found in Storrs, Connecticut until early July, and fifth instars are found from mid- August to early October. Oviposition begins at least by mid- June and continues until early August. However, oviposition apparently is not correlated with a photoperiod response since long or short day photoperiods do not affect oviposition time in the laboratory, and oviposition occurs under short (12 hour) photoperiods. The stadia as given in Table 38 are based on laboratory rearing on mixed fescue-sunflower seeds. The life cycle is especially characterized by a relatively long egg development period in the laboratory of 16 days which may in part explain the late occur- rence of nymphs. As already indicated the apparent preoviposition 73 ENTOMOLOGICA AMERICANA period after winter ends is long, at least two months. However, the precopnlatory period is much shorter as adults collected April 27 were found mating May 1st in the laboratory. The late cycle is probably related to a late maturation since maturation is speeded by bringing the overwintered insects into the warm laboratory. This late seasonal cycle closely coincides with the major seed food plant of Xestocoris, the fescue grass, which in Connecticut in early July ripens its seeds. Thus the nymphs develop during the period when the host plant seeds are abundant. The diapause is a relatively strong one and is broken only occasionally under warm laboratory conditions after a long dispause development of three or four months. The life cycle from egg to adult takes from seven to eight weeks in both the field and labora- tory. The fecundity in the laboratory is relatively low. On a mixed sunflower-Hesttma seed diet, only 40 and 62 eggs were laid by two females. These eggs were oviposited at a rate of 2-4 eggs a day. Sexually isolated virgin females lay no eggs. A substrate factor is apparently involved as egg production is greatly reduced (to 10- 20) when a suitable oviposition substrate is unavailable. The eggs are smooth, and thickly cylindrical with rounded ends. They are laid in crevices between culms, between the culm and a grass stalk, and into loose dry litter. They may also be laid in dry methyl cellulose, but are not laid on wet surfaces. TABLE 38 Stadia of Xestocoris miens Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 16.0 (14-17) 9.3 (8-11) 10.0 (8-12) 10.3 (8-12) 9.2 (8-10) 11.5 (10-13) 61.0 (49-72) 74 VOLUME XLIV TRIBE OZOPHORINI Ozophora picturata Uhler 0. picturata is the only representative in New England of the largely tropical tribe Ozophorini. The essentially Neotropical genus Ozophora contains 21 known species, 18 apparently restricted to an area approximating the Carribean Province of Good (1953). Only two species are known from South America, but this probably indicates once again our very poor knowledge of the South Amer- ica lygaeid fauna. Two species, 0. reperta and 0. trinotata ap- pear to be endemic to Florida. Not only does 0. picturata represent the northern limit of the genus, but it is also the most widely distributed species if the literature is to be taken at face value. It occurs north to Massa- chutts and Ontario, west to Iowa, Arizona, and perhaps California, and south to Mexico, Guatemala, Dominica, and Grenada (Slater, Catalogue). Some doubt, however, should be reserved for the Car- ribean records. The Ozophora species are closely related and need careful study. According to Slater (1952), while 0. picturata is a common insect in the southern states it is very uncommon in Iowa and Illinois. There is some diversity of opinion about its abundance, which may be linked to its distribution. Uhler (1876) remarks that it “is very rare near Baltimore, in spots with rank growth late in July.” Barber (1923) and Torre-Bueno (1925) said that it is frequently taken in New York by a sifting among dead leaves in late fall or sheltering under boards. Blatchley (1926) described it as “scarce throughout Indiana under logs, low marshy places, or herbage.” He also collected it as far south as the Everglades National Park, and considered it the least common species of Ozophora in Florida. In New England, I found 0. picturata com- mon along the coast of southern Connecticut at Noank and New Haven and fairly abundant at Storrs, Connecticut. But at Canaan, Connecticut in the highlands it is either absent or at least so scarce that I was unable to collect it. Slater (in litt.) collected a specimen at Camden Hills, in southern Maine. Environment In Connecticut, 0. picturata shows no evident seasonal change of habitat. It is found in the same habitats from season to season and year to year. This species is strictly an inhabitant of forest floors, ranging from climax oak-hickory forests to shrubby con- solidation seres. It is never collected in open habitats, but only where the litter layer is shaded. 75 ENTOMOLOGICA AMERICANA At Storrs, Connecticut Ozophora occurs in the thick loose litter of well-drained mesophytic oak-hickory forests, usually on moderate slopes. In relation to the life history of the insect this community forms a relatively permanent habitat. It appears to be of some interest too, that this rhyparochromine is the only seed feeding lygaeid found in the oak-hickory forest community which forms the climax community in much of the eastern United States (Braun 1950). In such habitats in southern New England, however, its density is never high, usually about 1-3 per square meter. This average holds quite constant over large tracts of oak forest in eastern Connecticut. Forests with maple, grey or black birch admixtures do not usually alter the counts. In this habitat the species is found most abundantly in the drier, well drained duff while depressions and spots with wetter litter rarely harbor the species. Ozophora is much more abundant in several shrub communities at Noank, Conn. The plant community of this sere consists of a few Acer rubrum L., with a dense shrub understory of V actinium, corymbosum (L.) and Viburnum dentatum L. and with a herb layer largely of Aster spp., Rhus toxicodendron L. and the fern, Dennstaedtia punctilobula (Michx). In the mesic leaf litter under the above plants 0. picturata, is found in a density of 20-25 per square meter along with the rhyparochromines Eremocoris ferus, Scolopostethus atlanticus, and Antillocoris minutus. At Noank, in the extensive dense sumac community ( Rhus typhina L., R. glabra L., R. toxicodendron L., Smilax spp.) so typical of the shrub sere of old fields along the Connecticut coast, there is a sparse population especially beneath occasional scattered black cherry trees ( Prunus serotina Ehrh.). On Ram Island at Noank in dry litter beneath a pure strand of bayberry ( Myrica pensylvanica Loisel.), first and second instars were found along with adults of Eremocoris ferus. In addition to occurring in a closed oak-hickory forest Ozophora is also found in the more open white oak forest with considerable short grass growing between the trees. It occurs here in abundances of 1-2 per square meter. The chief factors which distinguish the litter of the foregoing habitats is a general dryness of about 4-5, considerable shade, and a litter depth of at least two inches, usually more. The texture of the litter is important as Ozophora is always found in coarse, loose mull litter which characterizes the litter formed by oaks in contrast to most other trees. Both V actinium and Myrica also form a fairly loose but finer litter. Dry oak-hickory slopes are frequently found on well drained south slopes of moraines in southern Con- 76 ENTOMOLOGICA AMERICANA The Committee on Publications of the Brooklyn Entomological Society regrets to announce that it is compelled to suspend publication of Entomo- logica Americana with the present volume. The editor wishes to take this occasion to thank contributors and readers who have been so coopera- tive and pleasant during his tenure as editor. It is the hope of your editor that this old and distinguished journal will be reactivated in the fu- ture so that it can once again serve those interested in the Science of Entomology. J. A. Slater VOLUME XLIV necticut. The soil is usually of overdrained sand and gravel. But where the litter is thick and the uppermost layers dry, the soil may vary considerably in moisture. Forest litter temperatures are moderate in summer with re- corded litter temperatures never over 75° F. Since the forest com- munity inhabited by picturata extends much farther north than does picturata and the relative abundance of the insect drops very rapidly north from the coast in Connecticut, it is probable that temperature determines the northern limits of the distribution of this species. To explore this hypothesis, the resistance of picturata to cold stress must be studied. As already mentioned, populations of Ozophora in Connecticut tend to be thin, usually 1 or 2 per square meter, and occasionally up to 20 per square meter. There is apparently little tendency to aggregate, and the populations are dispersed fairly evenly over large areas. In 38 square meter counts on an oak-hickory hillside at Mansfield, Connecticut an average abundance of 2 per square meter was obtained with extremes of 0-4 per square meter. This density pattern was repeatedly observed in oak-hickory associa- tions. Dowdy (1947) collected ten specimens in a 10.8 square feet sample on August 14 in Missouri in oak -hickory litter. Available habitat notes largely confirm the forest preference of Ozophora. Torre-Bueno (1912) beat the species from oak at Yap- hank, Long Island, New York in a pine-scrub oak roadside area. Froeschner (1944) collected it from May 11 to August 2 under rocks and logs in Missouri. Dowdy (1947) recovered it from forest litter during an ecological study of an oak-hickory forest at Jeffer- son, Missouri. Other habitat notes are as follows. Gaines (1933) collected it in a trap in a cotton field in Texas, and Hussey (1954) caught the species in a baited malt trap in a cotton field in Georgia. Linsdale and Trevis (1951) reported collecting the species in Monterey Co., California with 0. depicturata Barb, in the nest of Neotoma fuscipes, the dusky footed wood rat. General Biology 0. picturata often inhabits a climax association which is surely a permanent habitat. Yet these insects are entirely macropterous as are all known members of the genus Ozophora. Several aspects of the behavior of these insects may explain the importance of macroptery. This fast moving rhyparochromine which is extraor- dinarily difficult to catch, may frequently “flit,” to use Southwood’s term (1961) to evade capture. These insects are known to fly at night and come to lights (Blatchley 1926). Torre-Bueno (1912) captured three by lantern on Long Island on September 24, 25, and 78 ENTOMOLOGICA AMERICANA I captured three adults August 20, 1958 at Noank, Connecticut and September 10, 15, 1961 at Storrs, Connecticut. Other species of Ozophora also have been captured at lights. According to Barber (1954) 0. atropicta was collected at lights in Puerto Rico by Ramos (1946) and 0. pallescens was taken at lights in Puerto Rico by Wolcott (1941). Barber (1939) records specimens of 0. quin- quemaculata taken at lights, and Blatchley (1926) collected 0 . trinotata at lights in Florida. Dispersal flights may be an essential part of the life cycle of this insect, but with little change of habitat. Myers (1926) found Ozophora in beach drift litter in Massachusetts. The selection value of dispersion and protection, then, may be factors which outweigh the advantage which can result from brachyptery. The coloration of this insect, both adults and nymphs, which is a light brown mottled with off-white areas, blends it very closely into its leaf litter habitat. Like many other members of the genus, 0. picturata has a white ring around the basal one half of the terminal antennal segment. The nymphs also have the last segment entirely pale. Against the dark leaf litter, these white rings are very conspicuous as the resting insect slowly moves its long anten- nae to and fro. It seems quite apparent that such a white antennal segment serves to distract the attention of a predator such as a bird from the bug’s exact location (Cott 1940). No predators or parasites are as yet known from this species. Like other rhyparochromines 0. picturata is a seed feeder (Sweet 1960). It feeds very readily on hulled sunflower seeds and the adults will pierce the hard shell to reach the kernel. It also feeds in the laboratory on seeds of Aster cordifolius L., A. lateri- florus (L.), Aster spp., Betula populifolia L., B. lenta L., on blue- berry seeds (Yaccinium corymbosum L.), and on grass ( Panicum ) seeds. It does not feed on acorns, of either white or red oaks, even if hulled. It then appears that the sparse distribution of Ozophora in oak forests may reflect the rather poor herb layer which develops on the deep litter of the dry oak slopes. I was able to rear this species on sunflower seeds through four generations. With each generation, however, as is commonly the case, the mortality be- comes greater. No pure food rearing was attempted with Aster seeds. Occasionally, when starving, the insects will scavenge on dead and dying individuals, but very inefficiently. Ozophora in its normal activities moves slowly and deliberately much like Myodocha and Antillocoris. It constantly taps the ground with its long antennae as it walks. When excited, as Torre- Bueno noted (1912), it runs very rapidly and with great agility. If caged in overdense numbers the insects become very excited and 79 VOLUME XLIV active and no mating or sustained feeding was observed. This was true both during day and night. Under these conditions mortality became very high and reproduction low. Seed defense behavior was not observed in this insect. I have not been able to observe the entire sequence of mating in this species. I have observed, however, the initial response of an excited male to the female. The male showed no response to the female until he touched her. He turned, approached the female, and touched his vibrating antennae against hers. Then suddenly the male lunged for the female, and the insects tumbled actively with antennae and legs moving rapidly, and just as suddenly stopped and ignored one another. They mate more than once, and were found mating both by day and by night. Copulation lasted for 1% to 3% hours in the laboratory. Life History 0. picturata overwinters as an adult. Blatchley (1895) collected it in Indiana on December 3 in a log on a sandy forest hillside. Torre-Bueno (1925) collected it by sifting leaves in winter. Dowdy (1955) collected the adults hibernating in oak-hickory litter. 0. picturata hibernates in the same habitats in which it completes its life cycle. There appears to be very little change in population density from late fall to early spring in oak-hickory habitats which would suggest that winter mortality is low. The species, however, becomes quite scarce in late May and early June, but enough col- lections were made to show that the species in the field does not lay eggs until relatively late in the summer. The phenology given in Table 39 begins then on July 1 at Storrs, Connecticut. Only adults are found earlier. At Noank, Connecticut while last instar nymphs are found as early as July 17, 1958, much later in the year on September 7, there were 20% fourth, 30% fifth, and 50% adults. This may rep- resent a partial second generation at Noank since a few third instars were found September 20 at Storrs (see below). This late seasonal cycle may reflect the same southern distribution of this species as is illustrated by Pachybrachius albocinctus. Adults collected after mid- July remain reproductively inactive for two to three months. After this period in the laboratory at room temperature the insects spontaneously come out of this weak reproductive diapause, and no cold treatment is required. The field data indicates that there is apparently only one prolonged generation at Storrs with the females laying eggs over a consider- able period, as is shown by the occurrence of third instars as late as August 15. This reproductive pattern is similar to that of 80 ENTOMOLOGICA AMERICANA Cryphula trimaculata and Xestocoris nitens. Similar to Cryphula, the cycle may be hastened by “forcing” earlier in the laboratory. But while Cryphula remains in obligate diapause, the first genera- tion 0. picturata in the laboratory becomes sexually mature in about 18 days during June, and can be mated with field over- wintered males of the previous generation. This contrasts with adults reared later in the year from nymphs collected in July and August and the second generation and laboratory adults. These late summer adults remain in reproductive diapause for at least several months. The only significant apparent difference between the two sets of laboratory populations is day length. It may be hypothesized than that shorter day lengths bring about a facultative reproductive diapause in 0. picturata which may be broken by cold exposure or several months time in warm temperatures. This ex- planation seems to best fit the available data. It is apparent, too, that a relatively high ambient temperature must be attained for reproduction to occur, for the insects may be forced in April, yet nymphs are not found in the field at Storrs until July 1. The slower spring increase in ground temperatures of woodlands com- pared to field (Geiger 1950) would also delay development. It is highly probable that two generations a year occur in southern parts of its range and perhaps a partial second at Noank, Connecticut, (Froeschner (1944), in fact, found late instar nymphs on June 21 in Missouri). The stadia in Table 40 represent cultures at room temperature (75-80° F) on a sunflower seed diet. TABLE 39 Phenology of Ozophora picturata Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult July 1 July 9 100% 100% July 13 40% 60% — July 17 25% 25% 50% July 25 50% 50% Aug. 6 10% 10% 20% 50% 10% Aug. 15 10% 25% 40% 25% Aug. 24 5% 60% 35% Sept. 1 15% 85% Sept, 10 5% 95% Sept. 15 100% 81 VOLUME XLIV TABLE 40 Stadia of Ozopkora picturata Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 12.5 (10-17) 8.5 (6-10) 8.2 (5-10) 5.6 (5-8) 8.3 (6-11) 8.0 (6-11) 51.1 (27-63) The longevity of the adults, especially the female, depends on time of generation and whether mating had occurred or not. Mated females live 53-65 days in a forced generation while two unmated females lived 130 and 145 days. A normal generation, however, overwinters and lives nearly a year. One spring female lived 60 days in the laboratory. Overwintered adults are found as late as June 21 at Storrs, Conn. Sexually isolated females lay no or very few eggs. Of thirteen virgin females, only three produced a few eggs (average 5.7 eggs). In contrast, mated females in the laboratory lay 88-135 eggs. As there is one generation a year in Connecticut, the normal precopula- tory and preovipositional periods are difficult to determine. How- ever in the forced early summer culture the preovipositional period is 18 days. Some of the winter forced cultures are very irregular in oviposition time and rates. Starvation effectively halts egg production. 0. picturata in the laboratory oviposits in dry material such as fine litter, methyl cellulose, and similar materials and in wet sub- strates only if no other is available. The cucumber-shaped eggs are covered with fine knobbed hairs which effectively fasten the eggs to fibrous and flaky substrates. 82 ENTOMOLOGICA AMERICANA TRIBE STYGNOCORINI Stygnocoris Douglas and Scott The eleven species included in the genus Stygnocoris are re- stricted to the western Palearctic, except for two, S. rusticus and pedestris which have been introduced into North America. These species were probably introduced through ballast dumped by ship- ping vessels (Lindroth 1947). This is evident because both species show a spread pattern, first appearing in northeastern North Amer- ica, and later being found at more southern and western localities. Of the European rhyparochromines, these two species are un- usual in having a winter egg diapause (Pfaler 1936, Putshkova 1956). Diapausing eggs seemingly have a greater probability than do adult insects of surviving transport from Europe to North Amer- ica, because adult rhyparochromines at least in New England show little tolerance to water stress. It may also be significant for the establishment of the species that in contrast to European rhyparo- chromines, many rhyparochromines in New England overwinter in the egg state. Eyles (1963b) found S. fuligineus to be the most common of the English species of Stygnocoris at the Imperial Field Station, Sun- ninghill, Berkshire and intensively studied the biology of this spe- cies. The other species S. pedestris and S. rusticus were apparently rare in the area and Eyles suggested that their habitats did not overlap with that of fuligineus. Stygnocoris rusticus Fallen This eastern Palearctic species shows a definite spread pattern. It was first collected in northern New York by Torre-Bueno (Heide- mann 1908) and in Quebec, Canada (Horvath 1909). Despite in- tensive collecting, Parshley (1917b) did not find this species in New England until Moore collected it in 1918 in Maine (Parshley 1920a, 1922). It was collected in Nova Scotia at Truro in 1913 (Barber 1918c, Parshley 1923), and in British Columbia (Downes 1924). Barber (1948b) reported that Esselbaugh collected the species at Duckabush, Washington, and Slater (1952) collected the species in northern Illinois. The present study indicates that S. rusticus is now fairly abun- dant throughout northern New England. Its distribution south- ward may be limited by climatic factors for there is a rather sharp change in abundance from common in the cooler highlands of northwestern Connecticut to scarce at the lower elevations at Storrs 83 VOLUME XLIV and it is either absent or very rare along the southern coast of New England. These localities are separated by distances of only 40 to 70 miles and this pattern showed no significant changes for at least six years. Environment In New England 8. rusticus inhabits mesic open field habitats supporting an abundance of tall forbs such as Saponaria officialis L., Solidago canadensis, Solidago spp., Daucus carota L., Geranium maculatum L., Ranunculus acris L., Silene spp., and Potentilla fruc- ticosa L. This association is characteristic of a later state of arable succession on mesic loamy soils. Normally the biotope of 8. rusticus, especially during nymphal development, is on the ground. Under the tall forbs, the ground is shaded and relatively mesic (4-6) in midsummer. In early spring before the herbs have grown up, this biotope is more exposed to iso- lation. In late summer, the species is frequently found on the ripen- ing seed heads of certain plants, especially Tanacetum vulgare L. At any one time, however, most of the population is on the ground. In such a habitat 8. rusticus is found in its greatest abundance, at- taining 25-30 per square meter. It is found in much lesser abundances in a variety of open habi- tats more dominated by grasses, such as in new roadside associations, and is taken infrequently along woodland margins. 8. pedestris is frequently found with 8. rusticus and in the laboratory will feed on the same seeds, but 8. pedestris is least frequent in the tall forb old field habitat and is more abundant along wooded margins and rud- eral sites. The pattern indicates that while the biotopes of these species overlap they do not coincide. The abundance of these two species of Stygnocoris varies from year to year. In 1957 both spe- cies were abundant, but in 1960 both species were rather scarce, and in 1962 both were relatively common again. In the extensive European literature on this species, there are various brief habitat notes. It was collected from and under heather ( Calluna, Erica) (Scholtz 1847, Saunders 1892, Stichel 1925), from furze (Halbert 1935), from Pulicaria dysenterica (Douglas and Scott 1865, Butler 1923), from Tanacetum (Prohaska 1923, Smre- zynski 1954, Forster 1955), under Thymus and Potentilla (Duda 1885, Stichel 1925), from Achillea millefolium (Beer et al. 1935, Feige and Kuhlhorn 1938). Barber (1948b) also recorded this species as abundant on Achillea at Montreal, Canada. Woodroffe (1955) collected rusticus on cindery ground in large numbers with nymphs under Cerastium vulgatum. Nearly all of these plants from which 8. rusticus has been col- 84 ENTOMOLOGICA AMERICANA lected are meadow or open habitat species. Piasecka (1960) con- sidered the species characteristic of meadows in Czechoslovakia. Hedicke (1942) said that the species occurs in Switzerland under vegetation hummocks at edges of fields. Southwood and Leston (1959) characterized its habitat as dry sandy places with plenty of flowers and stated that 8. rusticus is now rarer in Britain than it was 50 years ago. In North America, Barber (1922) took it in “moss” in New York. Procter (1938, 1946) collected it near bogs and water on Mt. Desert Island, Maine. General Biology S. rusticus differs from 8. pedestris in being pterygopolymorphic but its habitat selection in New England is for the temporary habi- tats described instead of long persisting habitats inhabited by most indigenous pterygopolymorphic rhyparochromines. It is however dangerous to speculate on the natural habitat of an introduced spe- cies which may be exploiting an open niche in New England. More- over, the persistence of the species is enhanced by the overall con- tinuation of such forb habitats by agricultural activities. Its oc- currence in the long persisting heath habitats as well as tall forb habitats in Europe may indicate a different habitat selection. Again, 8. rusticus originally may be a savanna species of eastern Europe (Lindroth 1957). Stehlik (1952) suggests that the macropter is more abundant at high elevations in Czechoslovakia. Weber (1930) quotes Sahlberg (1868b) as believing that the brachypters are less sensitive to cold and penetrate further into the North than the macropters. 8. rusticus is a medium sized rhyparochromine, generalized in shape with short legs. It runs actively but not rapidly, and shows no special tendency to hide in narrow crevices. Its uniform dark grey coloration blends well into the dark substrate of its tall forb habitat. The nymphs however have bright red abdomens like nymphs of Drymus and seem very conspicuous in comparison with the adults. When disturbed while feeding on a seed head the in- sects feign death and drop to the ground. After a few seconds the insects recover and crawl away. No parasites are as yet known for this species. Thomas (1955) stated that the damsel bugs Nabis major , N. minor, and N. mirmi- coicles, the centipede Lithobius, and the ant Formica rufa prey on 8. rusticus. At least some of the forbs mentioned earlier from which rusticus has been swept are probably food plants. 8. rusticus feeds on seeds of bitter buttons ( Tanacetum vulgar e L.), yarrow (Achillea mille- 85 VOLUME XLIV folium), Solidago spp., and several undetermined fallen seeds. Woodroffe (1955) found it in large numbers under Cerastium vul- gatum and surmised that this is a host plant. Its significance is not clear, but Waddell (1951) stated that rusticus survived a testing period on a sweet cherry cover crop. It is definite that in the field the nymphs feed on ripe fallen seeds from the previous year and the insect does not climb on the plant to feed on the seeds until early autumn when oviposition takes place. The nymphs are readily reared to adults on sunflower or Tanacetum seeds. S. rusticus sometimes carries and drags small seeds about in the petri dishes to more protected locations, but no seed defense be- havior was observed. In the mating behavior the male senses the female well before actual contact, raising his antennae stiffly toward the female. When near enough the male springs suddenly on the female. He taps the females’ head with a slow alternate slapping movement. If receptive, the female becomes quiescent, and exposes her ovipositor which is then gripped by the male’s parameres. The male slides off the female into the normal end to end copulation posi- tion. Several females at this point become cataleptic for several minutes. Later the female recovers and actively runs about, the male running backward in unison. When the female is not receptive to the male she wags her an- tennae rapidly back and forth directly over her head while trying to dislodge the tapping male. One male, after leaping on the female, gripped one of her antennae with both forelegs before tapping her head. This occurred twice in succession and again when the excited male leaped on another male. This is the only time the swollen fore- legs were ever observed being used during mating behavior by any rhyparochromines. Since this species frequently lacks one of its an- tennae, this may be the reason. Copulation lasts for a long time in the laboratory, often for 10- 12 hours and is repeated frequently. In the field rusticus indi- viduals are very frequently found mating both on the Tanacetum seed heads and on the ground and running up and down the stems while en copulo. Even when disturbed by collecting, the mating pairs rarely separate. Life History S. rusticus has a univoltine seasonal cycle with an obligative egg diapause (Pfaler 1936). Several authors (Scholtz 1847, Stehlik 1952, Massee 1960) state that this species overwinters as an adult but these records were probably late autumn adults as the insects are cold hardy and have been collected as late as December 3 in New England. 86 ENTOMOLOGICA AMERICANA The eggs evidently hatch in late May and the observed phenology at Canaan, Connecticut is given in Table 41. TABLE 41 Phenology of Stygnocoris rusticus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 28 80% 20% June 5 30% 65% 5% June 11 40% 50% 10% June 20 5% 45% 42% 7% i% June 30 8% 20% 27% 45% July 5 10% 30% 60% July 15 5% 95% July 20 100% Mating occurs in the field continually from late July to October. Oviposition, however, does not begin until after mid-September. It is clear then that there is an extended period of nonreproduction. Such adults collected in the summer and kept in the laboratory re- main in this state, dying in midwinter. Adults collected after mid- September are normally reproductive. Exposing the insects to short photoperiods (12 hours) does not affect the condition. When two cultures were exposed to cold room temperatures (36° F.) in mid- September oviposition was stimulated. However one culture which was exposed to cold for 18 days in late October did not become re- productive. The adaptation of S. rusticus to cool conditions is shown by its moving around and mating under cold room (6° C) conditions. The egg diapause is strong, and does not terminate under warm laboratory conditions and the eggs die. Three aliquots of eggs kept under moderate (36° F.) conditions for four months also did not hatch. The eggs diapause in the katatreptic embryonic phase as a well-developed embryo. It appears probable that the southward distribution of S. rus- ticus is limited by the capacity of this insect to survive such a long summer nonreproductive period and then to oviposit vigorously in autumn. The stadia given in Table 42 are based on field collected nymphs reared to adults on sunflower seeds. Since only three to five records per stadium are available only the range is given. The longevity of 87 VOLUME XLIV adults in the period of reproductive quiescence varies from 36 to 130 days (mean 84). The precopulatory period was not determined. The oviposition period lasts about two months from mid-September to November. Thomas (1955) stated that only a few batches of 10 eggs are laid. This undoubtedly results from depriving the insects of seeds. The reproductive rate of rusticus is actually much higher. The fecund- ity in the laboratory of 54-128 eggs (mean 98) is also probably low. TABLE 42 Stadia of Stygnocoris rusticus Instar 2 Instar 3 Instar 4 Instar 5 6 5-7 8 10-13 The eggs are relatively short and smooth with no spines. The eggs are coated with a cement layer that sticks the eggs to the sub- strate and to each other. Michalk (1935b) described and figured the eggs and referred to them as the horizontal agglutinent type. The eggs are laid singly in loose moist substrates as wet cotton and moist soil, and sometimes a group of eggs stuck together resulted, perhaps forming the “groups” mentioned by Putshkova (1956). Such a “batch” however is not laid at one time. Stygnocoris pedestris (Fallen) The history of 8. pedestris in North America parallels that of 8. rusticus. It has a similar western Palearctic distribution, and very probably has been introduced into North America. Gibson (1917) first recorded this species from North America from specimens col- lected in 1913 at Truro, Nova Scotia. It was a little later recorded from upper New York State and Cape Breton Island (Barber 1918, Parshley 1919). Parshley (1917b) however did not record the species in New England in his check list despite his intensive col- lecting. Scudder (1961) recorded this species from British Co- lumbia. It is possible that 8. pedestris has been introduced twice into North America by ballast, once each on the east and west coasts of North America. Although there are no further records of it in the literature, 8. pedestris is very abundant throughout northern New England. 88 ENTOMOLOGICA AMERICANA Even more so than in 8. rusticus, there is a pronounced change in abundance toward the southern coast of New England. In the northwestern highlands of Connecticut, 8. pedestris is extremely abundant, one of the most common lygaeid species. In eastern Connecticut at Storrs, it was very rare and only 2 individuals were collected in six years, and along the southern coast, none. Environment In northern New England this species has a rather wide eco- logical distribution and is the only introduced species that has in- vaded natural New England environments. The other three pre- sumably introduced species 8. rusticus, Megalonotus and Sphragis- ticus inhabit disturbed areas only. The habitat of S. pedestris ranges from woodland margins to disturbed roadside areas. In open fields with a greater dominance of grasses and with fewer tall forbs it frequently occurs with 8. rusticus. 8. pedestris inhabits relatively mesic litter (moisture 3 to 6) on the edge of forests of Tsuga canadensis, Pinus strobus, Junip- erus virginiana, Fagus grandentata, Betula spp., Acer spp., and in general along northern hardwood forest margins as well as in open fields and roadsides. It also occurs along the margin of Picea rubens in both highland forests in New Hampshire, and coastal Picea for- ests in Maine. It is not, however, a forest species and does not penetrate far into forests and is usually found on the field side of a forest-field ecotone. It occurs in low abundance of 3-5 per square meter in a few Vaccinium balds on Mt. Grey lock, Massachusetts and at Laconia, New Hampshire. It is never found in the xeric sparse grass habitat favored by many New England rhyparochromines, nor in oak-hickory forests. While its normal biotope is on the ground and the nymphs are never found off the ground, the adults like those of 8. rusticus sometimes climb up the plants in autumn to feed on the ripe seeds. It was swept from pearly everylasting, Anaphalis margaritacea L., 8 pirea tormentosum L. Tanacetum vulgare L. Beneath Anaphalis and in the margin of juniper litter it reaches an abundance of 50 to 80 per square meter. It is found in lower abundances of 10 to 20 per square meter in the other habitats. Over the five year study, its numbers did not fluctuate as much as did those of 8. rusticus but in the same, years that 8. rusticus was rare, 8. pedestris was also less abundant. The European literature indicates a similar habitat choice. 8. pedestris is very abundant in England (Butler 1923, Southwood and Leston 1959), although Eyles' (1963b) found it scarce at the Imperial Field Station at Sunninghill, Berkshire, England. It has been frequently found in leaves and moss litter (Lethierry 1869, 89 VOLUME XLIV Saunders 1892, Marchal 1898, Nicholson 1935, Leston and South- wood 1961), in Calluna-Erica heath (Scholtz 1847, Sahlberg 1868, 1920, Becker 1886, Stichel 1925, Stys 1960). Piasecka (1960) noted it in meadows near woodlands. Lambertie (1906) found it on Salix, but this is probably accidental. Krogerus (1960) found it numerous in southwest Finland in moss, in Ledum-C alluna heath and as ex- tending north to 62° 5". Southwood and Leston (1959) recorded 8. pedestris in England as being found on dry sand chalk or light soils with good vegetation cover. Lindberg (1958) recorded 8. pedestris on Newfoundland under stones on grass verges, on shingle overgrown with grass and on the seashore. As mentioned under 8. rusticus the biotope of S. pedestris over- laps on that of S. rusticus. However, it is not clear whether either of these species is displacing any native rhyparoehromines unless it be species like Myodocha , Heraeus, Zeridoneus, and Pachyb rachius basalis. These are all myodochines of relatively mesic habitats, but all are biologically quite different from the species of Stygnocoris in their different seasonal cycles, distributions, food plants, and behavior. General Biology S. pedestris, unlike 8. rusticus, is entirely macropterous (South- wood and Leston 1959). It is probable that this wing condition correlates with the wide ecological distribution of S. pedestris and the occurrence of the species in temporary roadside habitats. There appear to be no records that can definitely be attributed to dispersal. 8. pedestris is one of the smaller rhyparoehromines, being only about 2.7 mm. in length, considerably smaller than 8. rusticus. Its procryptic coloration is black and dark reddish brown ; the legs are inconspicuously pale. It is generalized in shape with short legs and is not a rapid runner, but conceals itself in the litter debris of stems, leaves, and grass of its biotope. The nymphs are conspicuous with the abdomen banded, pale and red and are quite different in coloration from the nymphs of rusticus. Frequently when disturbed, it feigns death. On Tanacetum even when mating pedestris will feign death and fall to the ground. After about 10-15 seconds it recovers and conceals itself. During this death feigning period it does not respond to a touch stimulus. Eyles (1963a) found an overwintering individual of 8. pedestris parasitized by Alophora pusilla Mieg, a parasite also known from the hemipterons Chilacis and Cydnus. 8. fuligineus, that Eyles (1963a) investigated intensively, is attacked by the fungi Poeci- lomyces and Entomophthora. No predators are known. 90 ENTOMOLOGICA AMERICANA 8. pedestris has been found in nests of the ants Myrmica sca- ~brinodis and M. miruginodis (Donisthorpe 1927) and of Formica rufa (Reclaire 1932). These records are probably accidental, re- sulting- from the ground layer habitat and abundance of the species. S. pedestris feeds on the seeds of Tanacetum vulgare L., Ana- phalis margaritacea (L.,) Spire a tormentosum L. and S. latifolia (Ait.), Aster novaeangliae L., Solidago spp., Betula populifolia, and a number of other unidentified fallen seeds. When starving the insects will scavenge on dead and dying insects but cannot sub- sist on this diet and soon die. No seed defense behavior was ob- served in this species. In the mating behavior, like 8. rusticus, the male senses the female usually before actual contact. When a female is near, the male springs suddenly on the back of the female, turns into a posi- tion parallel to the female and begins to tap her head with alternate rapid movements of his antennae. The tapping is more rapid and has a smaller arc of movement than in 8. rusticus. Under the tapping the female quiets and releases her ovipositor and copula- tion begins. In three cases after the male swings off the female to assume the end to end position, there is a cataleptic pause affecting both sexes which lasts about 30 seconds. After moving to the end to end position there is some pulling and pulsating between the male and female until abruptly the pair move together with the male running rapidly backwards. Copulation lasts from 3 to 11 hours and is repeated frequently over a period of three months from July to October. Many copu- lating pairs are found in the field at this time which correlates with the long copulation period in the laboratory. Life History Like S. rusticus, S. pedestris has an univoltine life cycle with an obligative egg disapause (Pfaler 1936). There is also an ex- tended adult reproductive diapause in late summer. Eclosion occurs in May, and the earliest nymphs are found in late May. The observed phenology at Canaan, Connecticut is as given in Table 43. In northern New England a few fifth instars were found as late as August 20. The long preoviposition period of the Canaan population lasts until late September although copulation occurs from at least late July and through the autumn oviposition period. Adults which are collected after September 26 actively oviposit. Females brought into the laboratory do not oviposit as do the field adults but remain in a nonreproductive condition until death in late autumn and early winter. A few of these fertilized females laid a few (2-17) 91 ENTOMOLOGICA AMERICANA eggs, but such ovipositing soon ceased. The long prereprodnctive period is not affected by photoperiodic conditions. Adults reared in the laboratory under short, long, normal (decreasing), and even increasing photoperiods, remained in diapause. TABLE 43 Phenology of Stygnocoris pedestris Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 28 83%' 17% June 7 55% 35% 10% — — June 15 21% 40% 33% 6% June 22 5% 25% 43% 17% 10% July 4 3% 12% 85% July 15 2% 98% July 20 — 100% Exposure of three cultures of adults to repeated cold (6° C.) exposures of 7-24 hours each in August did not stimulate oviposi- tion. However, exposing adults of four cultures collected in sum- mer to low temperatures in September did stimulate oviposition. These colonies normally would not have become reproductive. How- ever, it is noteworthy that the oviposition rate fell off rapidly in the laboratory except in one colony that was reared in a cool office. The rearing room temperature may have been too high for optimal oviposition. It is possible that both cold and short photoperiods are required. There is also a definite acclimation because adults exposed to 6° C. in August are immobilized but adults in late November move actively about at temperatures barely above freezing (2°— 3° C.). The records of 8. pedestris overwintering as an adult (Scholz 1847, Butler 1923, Murray 1936, Michalk 1938b, Massee 1960) undoubt- edly are late autumn collections reflecting the cold resistance of this species, or perhaps may represent a confusion with 8. fuligineus which does overwinter as an adult (Pfaler 1936). Although 8. fuligineus overwinters as an adult, Eyles (1963b) states that it does not have a diapause state. Yet his data indicates that 8. fuligineus is not reproductive (immature) for periods of two to three months in summer which suggests that a reproductive diapause is involved in this species. 92 ENTOMOLOGICA AMERICANA The eggs diapause either in early anatrepsis or late katatrepsis. Some eggs kept in warmth will develop to late katatrepsis, but no further. Normal diapause eggs will not hatch kept at room temp- eratures. Both egg stages overwinter at 6° C. and complete dia- pause development at this temperature. Non-diapause eggs were obtained from two females collected on November 27 and which had been exposed to low field temperatures. The eggs which were placed in the cold room remained undeveloped, but the eggs left in warmth developed and hatched in 16 to 23 days (mean 19 days.) This indicates that as in L. diffusus the dia- pause-producing mechanism was broken in the female before ovi- position. However, the egg development period of these eggs was considerably longer than in diapause eggs which have completed diapause development. The stadia of immature forms are given in Table 44. The shorter stadia figures apply to the more rapid development of the nymphs collected in the field and reared in the laboratory. The longer stadia apply to nymphs reared in the laboratory from eggs. The egg incubation period indicates the time from the day the eggs were moved out of the cold room until they hatched. This is proba- bly not comparable to a normal egg development period. TABLE 44 Stadia of Stygnocoris pedestris Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10.3 5.7 4.8 6.0 5.9 10.0 42 (8-12) (3-9)' (3-9) (3-10) (4-10) (4-15) (28-63) The longevity of nonreproductive adults in the laboratory ranged from 52-144 days (mean 77). Only a single exceptional female lived 144 days. The mean represents the usual longevity. In the field at Canaan, Connecticut, adults were found as late as November 28. The laboratory longevities are consistently shorter than in the field probably due to warmer temperatures. Cooler summer temperatures also probably prolong the adult longevity in northern New England. Since the nonreproductive period in the field is approximately 60-70 days in Canaan, Connecticut (in southern New England), and only about 30 days in northern New England, the sudden drop off in abundance toward the Connecticut coast becomes more under- 93 ENTOMOLOGICA AMERICANA standable. The southward distribution of 8. pedestris is most likely limited by its capacity to endure a long summer nonrepro- ductive period. Adults forced from eggs hatched in early April failed to survive the summer. The precopulatory period was not determined but is no longer than two weeks. Copulation occurs freely from mid- July on. The preoviposition period is largely determined by the environment. Due to the interference of reproductive diapause, the fecundity figures of ovipositing adults is sketchy, ranging from 20 to 81 (mean 48) which is probably much too low, judging from oviposi- tion rates for a few days (5-7) and the length of the oviposition period in the field (late September to mid-November). The eggs are laid singly, preferably into moist substrates as the cotton stopper of the water vial or damp loamy soil. The eggs like S. rusticus are relatively short and smooth, and covered with a cement layer that sticks the eggs to soil particles and litter and sticks the eggs together. 94 ENTOMOLOGICA AMERICANA TRIBE DRYMINI DRYMUS Fieber This well-named genus is characteristic of forest habitats both in North America and in Europe (Leston and Southwood 1959.) Two Nearctic species are known, D. units and D. crassus. Recently the genus has been divided into two subgenera, the typical sub- genus, and Sylvadrymus (Le Quesne 1956). Despite having, at most, only a sparse pubescence instead of long erect hairs on the tibia, the remarkable expanded parandria of the genital capsules place both D. units and D. crassus in the typical subgenus. More- over, and unlike the European Drymus sensu stricto, the punctura- tion of the anterior lobe of the pronotum is only slightly finer than the posterior lobe’s and hardly so in D. crassus. As remarked under the distribution discussion, although Dry- mus is found throughout the Palearctic with 17 species, the two Nearctic species are restricted to eastern North America. Since a number of the species are not yet placed, the distribution of the subgenera cannot be discussed except to say that both occur in Europe, and only Drymus sensu stricto in the Nearctic. Although all the European species are recorded as over-winter- ing as adults (Pfaler 1936, Southward and Leston 1959), both of the Nearctic species, at least in New England, overwinter as eggs. However Putshkova (1956) states that D. brunneus lays winter eggs in Russia, although in the key she notes “eggs in spring.” Under the discussion of D. units the seasonal cycle of D. brunneus is dis- cussed further in light of the recent observations of Eyles (1963). Drymus contains both macropterous and pterygopolymorphic species. Woodroffe (1963) found that the English species of Dry- mus and Sylvadrymus form two parallel series based on wing de- velopment. In Sylvadrymus, sylvaticus F. is macropterous, ryei (D.S.) is submacropterous and brunneus is brachypterous and all three species may be found together. Similarly in Drymus ( sensu stricto ), pilipes (Fieb.) is macropterous, latus largely submacrop- terous and pilicornis brachypterous and these species are also found together. Drymus unus (Say) D. unus is known from Quebec and Iowa south to North Carolina and Texas, which approximates the extent of the eastern deciduous forest biome. Uhler (1895) also mentions this species from Colo- 95 ENTOMOLOGICA AMERICANA rado and lower California, but these records must be further con- firmed, especially as there are several western species of Eremocoris which bear a slight resemblance to Drymus, and there are no sub- sequent records. Like most forest floor Rhyparochrominae there are few records of this species in contrast to its actual abundance in New England. Indeed, this appears to be the first published record of D. unus from Connecticut. Slater (1952) noted it as scarce in Iowa and Illinois. Environment D. unus is a common and characteristic inhabitant of the forest litter of light mesic woodlands. While it does occur in climax forests, it is much more abundant in light subclimax forests. It is abundant where black birch and red maple are associated with oak and hickory, but rare in the thick litter of dry oak slopes or in climax sugar maple-beech forest. In the heavy deciduous forest D. unus is replaced by D. crassus. D. unus is also apparently more abundant in edge habitats. Where abundant, it may even be found on the field side of forest edges, providing the substrate is mesic and shaded by rank herbs. Frequently it is found beneath isolated trees in open fields. In such favorable habitats its abundance may attain 40 per square meter. It occurs in intermediate abundance of 5-10 in both birch and hemlock forests associated with Scolopostethus diffidens , and Eremocoris ferus, and also in Vaccinium litter with Scolo- postethus atlanticus. A few specimens were collected at one ex- ceptional habitat, a hummock in an open brackish salt marsh. In the biotopes of D. unus the herb layer is thin, averaging a foot in height, and usually includes several species of Aster, Mitella diphylla L., Smilacina racemosa Morang., Anemone quinquefolia L., Solidago bicolor, Dryopteris spinulosum, other herbs and tree seedlings. D. unus is abundant in biotopes of fine friable mull lit- ter as beneath birch and maple, and rare in dry loose oak litter. The soil is nearly always a dark humus and mesic (moisture, 5-6). The chief limiting factors of this species are its mesic re- quirements. It is never found in habitats with ground tempera- tures over 26° C. In one area where it invaded the rank edge of a small field, the ground biotope was 24° C. while the air temperature above the herbs was 36° C. When exposed for a few minutes to this higher air temperature, Drymus quickly succumbed. D. unus is also infrequent in very wet habitats such as swamps and ex- tremely mesophytic forests. D. unus is closely restricted to its litter biotope and only rarely is swept from plants, these woodland Compositae in late autumn. 96 ENTOMOLOGICA AMERICANA It frequently remains in a habitat for several years, and in one black birch oak habitat it was continuously present for at least nine years. In more marginal sites, such as along ecotones, its occurrence is only temporary. It was collected by sifting under alders (Smith 1910), from grass piles (Torre-Bueno 1929a), and under stones and boards on Mt. Desert Island, Maine (Procter 1946). Dowdy (1955) collected D. unus in an oak hickory woodland, and Blatchley (1895, 1926) found it in shelter under a rail near the border of an upland wood. General Biology It would appear that the entirely macropterous I), unus in- habits forest areas which are habitats as permanent as any of the habitats of the xeric-adapted brachypterous lygaeids. However, it is especially abundant in edge and glade habitats and it readily colonizes isolated patches of forest. Full macroptery and ready dispersal is probably a definite advantage in a mosaic of subclimax woodlands or, in a more homogeneous forest condition, it would allow the ready invasion of new habitats such as river bottoms and glade openings in forests. The adults of Drymus are a deep dark red-brown and dull black which effectively conceals the insect on the forest floor. This pat- tern is especially effective since D. unus readily feigns death. In death feigning, the antennae and the legs are rigidly held close to the body. When it is disturbed on a low herb, it feigns death and falls directly to the ground. Shortly afterwards it quickly seeks refuge under bits of litter, but does not run rapidly as do the long- legged lygaeids. The nymphs, in contrast, are a very bright red against the forest floor which would make them conspicuous to bird predators such as towhees or catbirds probing the leaf litter floor. D. unus does not feed on acorns, ash, willow, or maple seeds, but rather strongly prefers the seeds of small composites, especially the woodland asters. It could sometimes be found feeding on the dried seed heads of this plant in autumn as late as November 18 with the air temperature at 6° C. It also feeds on the seeds of Solidago spp. and Spirea latifolia and is readily reared on Betula seeds as well as sunflower and Aster seeds. The nymphs feed on the fallen ripe Aster , etc., seeds of the previous year while the oviposit- ing adults feed on the newly fallen seeds. Thus the seasonal cycle of this insect correlates with the fall seed production of the host composites. When hungry the insects readily carry or drag the seeds about. When disturbed during feeding, the insect endeavors to protect its position by maneuvering itself between the aggressor and the seed 97 ENTOMOLOGICA AMERICANA and by wagging its antennae. No further seed defense behavior was observed, except that the defender, with its hind legs, may push away the aggressor. In courtship, the male recognizes the female apparently only on contact. In every instance observed, the male advances on a female which is feeding. This may simply be an occasion when the female remains still. The male’s excitement is shown by its rapid agitation in moving around and in climbing upon the female. The female frequently dislodges the male by jerking her body sharply from side to side, or earlier she may keep him away with her hind legs. If the female is passive, the male vibrates his antennae on the female’s head. The male’s antennae which are held parallel and close together with the apical joint bent slightly down, are vibrated very rapidly with a barely discernible plane of movement. Life History D. unus has a univoltine seasonal cycle and diapauses over the winter as an egg. The phenology at Storrs, Connecticut is given in Table 45. This is a pooled phenology from summer of 1957 to spring of 1963. The first instars were found as early as May 19, but could also be found as late as June 15 at which time third, and fourth instars were present. This probably indicates that eclosion is staggered over a long period. TABLE 45 Phenology of Drymus unus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 19 85% 15% — May 30 25% 63% 12% — June 10 12 %■ 40% 42% 6% June 19 4% 36% 53% 7% June 27 3% 52% 45% July 5 16% 78% 6% July 18 1% 38% 61% July 27 — 2% 98% Aug. 4 100% Copulation occurs from August to October. However, oviposi- tion does not occur until after the advent of cold weather. In the 98 ENTOMOLOGICA AMERICANA laboratory under both long and short day photoperiods and in warmth (ca. 25° C.) there is either no or a greatly delayed and diminished oviposition in most adults captured before mid-Septem- ber. In most adults collected after this time oviposition in the laboratory is rapid and plentiful. This is probably in response to cold nights. Repeatedly exposing several summer reared colonies to cold room (5° C.) conditions stimulated increased oviposition. The cold room was continuously illuminated to simulate long day conditions. However, photoperiod cannot be ruled out, short days may further promote the basically cold stimulated oviposition. D. unus is remarkably adapted to cool conditions and even ovi- posits under cold room (5° C.) conditions. It was observed on November 6, at dusk, feeding and mating on seed heads at an air temperature of 6° C. Ovipositing females have been collected in the field in late autumn as late as December 24. This late oviposi- tion period and cold hardiness explains the over-wintering records in the literature, as these are all late November or December records (Blatchley 1895, 1926, Torre-Bueno 1925, Barber 1928d, Dowdy 1955). At this point it is important to compare the seasonal cycle of D. unus with the European D. brunneus. Eyles (1963b) suggests that in England, D. brunneus overwinters in three stages, as adults, eggs or nymphs. However I see no clear evidence that the nymphs overwinter. Even so if D. brunneus overwinters in two stages as either immature females or eggs then there must be two reproduc- tively isolated populations as the species is obligatorily univoltine. This would imply that two sibling species are involved with quite different seasonal cycles and the corresponding physiological adap- tations. Perhaps a more probable interpretation of the data is that D. brunneus like D. unus is cold hardy, oviposits in autumn, and per- sists into winter. Perhaps a few survive to also oviposit in the early spring. This would be similar to the pattern in D. unus. More- over the nonreproducing adults of D. brunneus in Eyles ’s insectary parallel the reaction of D. unus which did not oviposit under warm laboratory conditions outlined earlier. The oviposition period is long, and in the laboratory was con- tinued in some normal, cold exposed and fall collected specimens until early January. Since Drymus unus ranges as far south as North Carolina and Texas, it appears entirely possible that in the more southern areas the normal oviposition period may fall during the prevailing mild winter period. The laboratory evidence indi- cates that this is within the insect’s physiological capacity. Sum- mer reared Drymus kept in warmth frequently begin laying a few 99 ENTOMOLOGICA AMERICANA eggs after October 15. This continues sporadically through the autumn and winter and in one case continued until as late as March 10. If this seasonal cycle is present further south, it follows that the southern limit of the distribution of D. unus may be determined by the capacity of the insect to survive a very long summer re- productive diapause. Drymus nymphs which are forced to early maturity in the laboratory by early June rarely survive past late September, which would tend to support this hypothesis. The egg diapause occurs in early anatrepsis in most eggs, but some attain katatrepsis and remain in that state. As in Ligyrocoris diffusus, when the females are exposed to considerable cold, i.e. in adults collected during late November, from 10% (November 5) to 40% (November 26) of the eggs do not enter diapause, but hatch directly in the laboratory. This indicates that cold exposure affects the diapause forming mechanism in ovarian egg development. Such late eggs, then, must normally “hibernate” over the winter. The entire nymphal life cycle takes about a month and a half in the field, and is shortened to a little more than a month in the laboratory. Part of the explanation for the apparently variable ecolsion and development rate in the field probably resides in the changing microclimate with the ground layer exposed and sunny in early spring, but shaded when the canopy develops in late May. TABLE 46 Stadia of Drymus unus Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 7.6 6.0 6.8 7.8 9.4 (5-9) (5-9) (5-9) (6-9) (7-13) The stadia given in Table 46 are derived from rearing field collected nymphs. Egg development period is not available. The first stadium is based on laboratory hatched first instars. These could not be reared past the third instar. As mentioned earlier, the adults are long-lived and the average longevity in the laboratory of adults reared in the laboratory ranges from 62 to 238 days (mean 116 days). There is no marked difference between mated and un- mated females or males. Egg production in the laboratory is highly variable. In normal cold exposed field collected females 120-250 eggs (mean 131) are recorded. About half of the females which were not exposed to 100 ENTOMOLOGICA AMERICANA cold, later laid a few fertile eggs, (6-38 eggs (mean 24) ). The eggs are laid singly in the laboratory at a rate of 2-7 eggs (mean 3.2) in normal females. Three sexually isolated virgin females which were reared in mid-summer when given a cold exposure laid no eggs in contrast to the control fertilized females which laid 36-85 eggs. The eggs are laid singly after some probing for an oviposition site. The eggs are laid on wet or dry substrates and are deeply imbedded. The eggs are relatively short and fat and covered with fine nap like those of D. crassus. The eggs stick to each other and to the substrate by a cement secretion. Eggs which will diapause become quite dark in comparison to the few non-diapause eggs. Drymus crassus Van Duzee This is a much larger species than D. unus. It is recorded from a slightly more restricted range which corresponds closely to the deciduous forest biome. It is known from North Carolina to Texas north to South Dakota, Iowa, and New York (Slater, Catalogue). I have collected it in Connecticut and also in Maine in the coastal Acadian spruce zone. Environment Drymus crassus has the most mesic habitat preference of the various rhyparoehromines. It is only found in mature leaf mold sites beneath mesophytic forests, especially on north exposures. It is not found in oak-hickory forests or younger subclimax associa- tions, but is found in maple-beech, hemlock-white birch-maple, and red spruce (coastal Acadian) litter. In New England it accord- ingly does not exhibit a southern type distribution, but rather re- flects the distribution of mesic habitats. It has been infrequently collected, probably because of this habitat choice, and partly because of its low density (2-6 per square meter) in such dense mull litter. Its extreme mesic requirements demand even greater care in collect- ing than D. unus to avoid exposing the insects to sunlight. The litter, while very mesic in type, is at 8-9 moisture level, not water- soaked or hygric. Most noteworthy is that D. crassus is rarely found with D. unus despite the presence of large populations of D. unus nearby. Blatchley (1926) reported finding it on a densely wooded slope in Indiana, a habitat suggesting those reported here from New England. General Biology Drymus crassus at first appears macropterous but it actually displays an unusual type of wing polymorphism. In the flightless 101 ENTOMOLOGICA AMERICANA form the corimn is elongated and the membrane although it attains the apex of the abdomen, is narrowed, and only partly overlaps the other membrane. The hind wing is considerably reduced in length and barely reaches past the end of the commissure. As the corium appears thick and heavily sclerotized, the wing takes on a distinctly coleopteroid appearance. This condition forms a logical inter- mediate to the remarkable tropical coleopteroid forms. Only one macropter was seen among 52 specimens. This largely flightless condition correlates well with the perma- nent climax forest habitat of D. crassus. The presence of the macropter must have enabled the spread of this species into mesic habitats in areas like Canaan Mountain, Connecticut which was com- pletely deforested at one time. As in D. unus, the dark coloration of the adults form a procry ptic coloration, but the nymphs are bright red and conspicuous on the leaf mold. Is this a warning coloration for bird predators ? When disturbed in the field the adults, like D. unus, frequently feign death. Care must therefore be taken in searching for these insects as the adults blend in very well with the background. These insects feed on Aster spp. seeds, birch ( Betula papyrifera L.), and very readily on sunflower seeds. Mating occurs in the usual reversed position in late summer and autumn, but courtship was not observed. However there appar- ently is complete reproductive isolation between the Drymus species as three Drymus unus females enclosed with 5 Drymus crassus males were not inseminated. Life History The seasonal cycle is similar to D. unus, with one generation a year and an obligative overwintering diapause in the egg state. The phenology is rather sketchy: on July 14, at Storrs, one fourth, and three fifth instars were collected; July 26, seven fifth instars, three adults ; August 4, one fifth instar and seven adults ; and from August 15 on, all were adults. The stadia are not known. The diapause situation appears to be very similar to D. unus. The population collected in late July mated readily but produces no eggs whatever throughout autumn in the laboratory under short day periods. Adults collected later in August laid a few eggs, but from September 14 on, diapause eggs are laid readily. The eggs diapause in early anatrepsis. None could be released from diapause by moderate cold (4° C.) exposures of five months. Fertile eggs are easily told from unfertilized eggs because the egg shell becomes dark and horn colored in fertile eggs. There are, then, two sorts of diapause states operating as in 102 ENTOMOLOGICA AMERICANA D. unus, a late summer oviposition inhibition and an overwintering egg diapause. It is probable that the very late autumn “overwin- tering” insects reported by Torre-Bueno (1925) and Barber (1928d) represent cold resistant insects as in D. unus. It may ovi- posit throughout the winter further south. The eggs are laid at a relatively low rate of 2-4 per day, and total egg productivity ranges from 49-93, which appears to be a little low, but may correlate with the permanent habitat, as this is a relatively large rhyparochromine. The large thick eggs are preferably laid on moist substrates, and a loose substrate is preferred. Like the eggs of D. unus , the eggs of D. crassus are beset with minute spinules. Scolopostethus Fieber This Holarctic genus includes 20 species of which 16 are known from the Palearctic, 4 from the Nearctic, (8. ihomsoni is found throughout the Holarctic region). Horvath (1893) thought that 8. thomsoni may have been introduced into North America. It is more probable that this species occurs naturally in North America. In the eastern United States 3 species occur, one, 8. atlanticus, is restricted to the eastern United States. In the western United States are two other species, one of which, 8. tropicus extends sohth to Guatemala from where it was described by Distant (1882). The species of eastern North America are adequately distin- guished by Barber’s (1918e, 1923) keys to the species, but care must be used for some populations of 8. thomsoni resemble 8. at- lanticus in the coloration of the hind femora and the reduction and absence of the proximal spines on the fore femora. Eyles (in litt.) has crossed several of the English species and obtained progeny which were sterile. I have been unable to cross 8. diffidens with S. atlanticus or with S. thomsoni (Car ex popula- tion). Eyles (1963b) has studied the life histories of four species of Scolopostethus, 8. thomsoni Reut., 8. affinis (Schill.), 8. decor atus (Hahn) and 8. grandis (Horv.) and concluded that the species are univoltine and have no diapause, overwintering in a quiescent state. However, it must be noted that the new adults remained immature for a rather long period, especially 8. decoratus, suggesting that a reproductive diapause may be involved. 103 Scolopostethus diffidens Horvath 8. diffidens has a boreal distribution across North America and extends south to British Columbia and northern California. It is much more abundant in northern New England than southern New England and apparently does not occur along the coast between Cape Cod, Massachusetts and New Haven, Connecticut. Environment S. diffidens is a sylvan species typical of northern birch-conifer- ous forests. It is restricted to and abundant in the litter beneath grey birch (Betula populifolia Marsh), white birch ( Betula pa- pyrifera Marsh), hemlock ( Tsuga canadensis L.) and red spruce ( Picea rubens L.). 8. diffidens is found in either pure or mixed stands of these trees. Its presence in southern New England largely parallels the presence of this forest assemblage. The forest herb layer is frequently absent and is of no apparent significance in the ecology of this insect, except occasionally, as an edaphic factor. The spruce and hemlock forests form climax assemblages but the birches, especially gray birch, are subclimax. However, these are long persisting subclimaxes in terms of this insect’s life cycle. These trees often form extensive stands, and the edaphic effects of the varying forest floor conditions are readily observed. Where the litter is thin and tightly packed, as commonly occurs under pure coniferous stands, 8. diffidens is scarce. In rougher spots where small twigs and herbs create a looser litter, the species is more abundant. The same effect occurs when the litter is of mixed conifer-birch leaves, or completely of birch. Slightly drier spots also favors diffidens and very wet litter is unfavorable. The very dry litter of some of the mountainside Picea forests is also a rather unfavorable habitat for 8. diffidens. In general, thick loose dry-mesic (5-7) litter with many fallen seeds from the trees forms the most favorable biotope for this species. Its abundance is frequently extremely high in such bio- topes, attaining at least 130 per square meter. 8. diffidens fre- quently occurs in clusters which indicates a definite sociability effect which is apparently unrelated to edaphic factors. Temperature conditions are moderate in the litter because of the shade, rarely exceeding 26° 0. at midday, even in sun flecks. Al- though gray birch is abundant in southern New England it is only on north or east exposed slopes under old stands that 8. diffidens occurs. On such slopes the snow cover lasts until rather late re- tarding the warming of the ground biotope. There are no observ- able seasonal population movements. 104 ENTOMOLOGICA AMERICANA Populations of 8. diffidens were found year after year in the same habitat and at three sites for six years in succession. As dis- cussed under Competition, diffidens is one of an assemblage of seed feeding rhyparochromines in this habitat. Despite the abundance of 8. diffidens in its habitat, there are very few records of it in the literature. Both Blatchley (1926) and Barber mention it as being collected under dead leaves. General Biology In correlation with its climax or long persisting habitat, 8. diffidens is pterygopolymorphic with only about 10% of the popula- tions macropterous. Although there are no definite dispersal rec- ords, during the spring there is a definite decline in the abundance of the macropterous form. Brachypter x brachypter matings yield a few macropterous as well as brachypterous progeny. These small rhyparochromines are brightly marked with white, tan and black with conspicuous white spots in the lateral corners of the hemelytral membranes. The nymphs of the later instars also have their reddish abdomens laterally marked with white areas. Although these color patterns do not appear very ant-like, when a cluster of these insects are disturbed the sudden boiling up of the insects and their rapid jerky running produces a very definite ant- like effect. This interpretation of the protective coloration and behavior may also explain the sociability effect mentioned earlier. Such an ant mimicry effect has also been noted by European work- ers in 8. affinis Schill. (Wassman 1889, 1894, Guide 1937), S. pictus (Marchal 1898, Butler 1923) and 8. pseudograndis (Singer 1952). Despite its existence in large populations, no insect parasites have been reared from 8. diffidens. Several fifth instar nymphs had red mites (near Thrombidium) attached to them in the field. No predators are as yet known. 8. diffidens is readily reared on the seeds of the trees mentioned earlier, white and gray birch, hemlock and red spruce. It also feeds readily on sunflower seeds but suffers a very high mortality. The seed defense behavior of this species is easily elicited by presenting seeds after allowing a colony to become hungry. The insects clash over seeds by pushing and flailing at each other with their antennae and fore tibiae. Neither the fore femora nor the beak are used in this behavior. After repelling the aggressors, the insect pierces the seed, and drags it with its beak to a more pro- tected site for feeding. The mating behavior was not observed in detail, but a male after touching the female climbs or leaps quickly on her and begins vibrating his antennae on her head. Mating lasts for at least 2 105 ENTOMOLOGICA AMERICANA hours and the adults move about readily in the end to end position, with the male running- backwards. Life History Scolopostethus diffidens has an univoltine seasonal cycle with an obligative adult diapause. Although the females are fertilized in early spring, some as early as March 26, the nymphs do not appear until June. The phenology of 8. diffidens at Storrs is given in Table 47. These insects are quite cold hardy and are found actively moving about in the field directly after the snow melts. Such adults are not yet sexually active and the females are un- fertilized. TABLE 47 Phenology of Scolopostethus diffidens Date Instar 1 I ns tar 2 Instar 3 Instar 4 Instar 5 Adult June 1 — — 100% June 11 20% 80% June 19 45% 15% 5% 35% June 27 20% 30% 25% 20% 5% July 9 3% 7% 25% 35% 25% 5% July 17 5% 10% 25% 35% 25% July 25 5% 15% 80% Aug. 7 7% 93% Aug. 21 100% Climatic effect on the phenology is clearly apparent. On Au- gust 10, 1960, on the cool northside of Canaan Mountain in the northwestern Connecticut highlands, the instar ratios were IV : 5% V : 20%, and adult : 75%. In the White Mountains of New Hamp- shire, the instar ratios on August 12, 1961 were III : 5%, IV : 20%, V : 50%, and adult : 25%. The late occurrence of nymphs is evidently a temperature effect affecting egg development because females dissected in early April have fully developed eggs in their ovaries. Females collected in late April and early May lay eggs in the field in the specimen bottles shortly after being captured. Photoperiod control is not involved. Colonies collected in early March and kept in 12 and 15 hour photo- periods became reproductive simultaneously. The adult reproductive diapause condition is relatively strong 106 ENTOMOLOGICA AMERICANA and only a few individuals live long enough to spontaneously break or complete diapause development under warm laboratory condi- tions. Again photoperiod conditions have no apparent effect on the diapause state. The longevity of the diapansing adults under warm laboratory conditions varied from 91-220 days (mean 175 days). Diapause is not broken until January or February after about 190 days of adult diapause and such few adults lived only 15 to 25 days. Exposing the insects to brief one month cold ex- posures of 6° C. did not terminate diapause. TABLE 48 Stadia of Scolopostethus diffidens Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 9.7 4.3 4.6 4.4 6.0 7.6 37.0 (9-11) (3-5) (3-7) (3-6) (5-7) (6-10) (34-46) The stadia in Table 48 are derived from spring cultures reared on mixed sunflower — gray birch seed mixture. Development is very rapid in the laboratory, definitely faster than in the field, probably because of cooler field conditions. Such forced spring cultures become adult by late May and early June. The diapaus- ing adults of these cultures begin dying in the laboratory in late August. It appears possible that the southern limits of this species is determined by its inability to survive the long summer in the diapause state. The spring precopulatory period is very short but in cultures collected in March after the snow melts the pre-oviposition period in the laboratory is about 7 days. The normal fecundity in the laboratory ranges from 94-184 eggs (mean 132 eggs). The eggs are laid at a rate of 4-5 a day. Very few eggs (15, 18, and 27) were laid by three females released from diapause in midwinter. Three sexually isolated females laid 24, 43, and 49 eggs. The smooth, relatively stubby cucumber-shaped eggs are laid loose in dry substrates and do not stick to the substrate. Wet sub- strates are utilized only when no other is available. Scolopostethus atlanticus Horvath I am using Barber’s written concept of atlanticus (Barber 1918, 107 VOLUME XLIV 1923) despite some question as to the status of the species (Lattin, in lift.). Horvath (1893) probably had this clearly marked eastern species in mind when he described atlanticus. In practice however Barber himself at the United States National Museum frequently and erroneously named specimens of 8. thomsoni and 8. diffidens as S. atlanticus. For this reason many if not most of the distribution records of atlanticus are incorrect. \ have seen definite specimens of atlanticus only from New Jersey, Long* Island, New York, south- eastern Connecticut and on Cape Cod at Wood’s Hole Massachu- setts. This is a typical pine-barren type distribution. The speci- mens of “atlanticus” seen from middle and northern New York (Torre-Bueno 1917, Barber 1923), and from British Columbia (Parshley 1919) are all referable to 8. thomsoni and should be deleted as records of 8. atlanticus. 8. atlanticus is clearly distinguished from 8. thomsoni by sev- eral characters in addition to those used by Barber. In the male of 8. atlanticus the large fore femoral spine is oddly bent rather than straight, and the parameres of S. atlanticus are only slightly curved as in 8. diffidens rather than strongly curved as in all populations of thomsoni seen. In both sexes of atlanticus the adult scent gland evaporatorium covers about 3/5 of the metapleuron similar to 8. diffi- dens while in 8. thomsoni the evaporatorium covers less than % of the metapleuron. It appears that 8. atlanticus may actually be most closely related to 8. diffidens. Environment The only habitat note in the literature that is definitely referable to 8. atlanticus is Barber’s collection of it under huckleberries in New Jersey (Blatchley 1926). These same specimens are in the U.S. National Museum. I have been able to find only one good station for atlanticus. This is a habitat at Noank, Connecticut. A few specimens were also collected at Storrs, Connecticut with S. diffidens. The Noank habitat is a mixed Vaccinium corymbosum — Vibur- num dentatum tall shrub community with an emerging overstory of Acer rubrum. The herb layer consists mostly of a heavy growth of the hay-scented fern Dennstaedtia punctilobula, Aster spp., and a few undetermined grasses. The biotope of S. atlanticus is in the shaded mesic litter beneath the V accinium and Viburnum where it coexists with Ozophora pic- turata, Eremocoris ferus, Antillocoris minutus, and Drymus unus. The litter varies from relatively moist (7) to mesic (4) and atlanti- cus is concentrated in the better drained and drier spots. As the ground layer is shaded, the litter temperatures are moderate. In 108 ENTOMOLOGICA AMERICANA late summer in 1957 8. atlanticus was the third most abundant species with 20-30 insects per square meter, but in successive years as the maples shaded out the shrubs, the abundance of the other rhyparochromines dropped leaving 8. atlanticus as the most abun- dant species but with only about 10 insects per square meter. 8. atlanticus was present in the same biotope for six successive years. General Biology Although Barber (1918e) thought 8. atlanticus was entirely macropterous, the population at Noank is polymorphic and con- sisted mostly (75%) of brachypters. Not enough is known about this species to evaluate the relation of its wing condition to its nor- mal habitat, except to note that its six years persistence at the Noank habitat is typical of pterygopolymorphic species. This brighly marked species behaves like 8. diffidens in the field and appears to be an even better ant mimic than S. diffidens. The nymphs unlike either 8. diffidens or any of the populations of thom- soni also have the tips of their fourth antennal segments pale white as in the early instars of Eremocoris ferus, which probably serves to distract predators. 8. atlanticus feeds on the seeds of Vaccinium and Viburnum and also a number of small unidentified seeds in the litter from the field. It probes the litter with its labium until a seed is sensed and then proceeds to pierce the seed. It frequently drags the seeds with its labium to more protected locations for feeding. Like S. diffidens, when hungry, the adults of atlanticus clash over the seeds, flailing at each other with their legs and antennae. Life History Like 8. diffidens, 8. atlanticus is univoltine with an obligative adult diapause. Despite the warmer coastal climate at Noank than at Storrs, the seasonal cycle is no earlier. The observed sketchy phenology is given in Table 49. Adults which are captured in spring have a long oviposition period in the laboratory which correlates with the late presence of nymphs and the overlap of old and young adults in July in the field. These new adults remain in reproductive diapause in laboratory w^armth until midwinter. The late occurrence of first instars has apparently no relation to photoperiod. Adults collected in late March and allowed to become mature under 12 and 15 hour photoperiods do so simul- taneously. As in 8. diffidens, a temperature response is probably involved. Bringing the adults into the laboratory forces the sea- sonal cycles several months ahead of the field population’s cycle. The diapause state is a moderately intense one and only a few 109 VOLUME XLIV adults survive under warm laboratory conditions to spontaneously complete diapause development in January. In cultures forced to maturity several months ahead of the field populations the few surviving adults become reproductive in November instead. This is a diapause period of five months. Such adults which complete diapause development in warmth live only a month as compared with spring adults which live two or three months or until as late as the middle of July. The stadia are inadequately worked out and only the single figures available are given in Table 50. The adult longevity is at least a year as atlanticus is a univoltine species and the generations overlap. TABLE 49 Phenology of Scolopostethus atlanticus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 25 100% June 9 20% — 80% June 20 25% 20% 10% 45% July 16 5% 15% 25% 25% 20% 10% Aug. 5 10% 15% 35% 40% Aug. 18 10% 90% Aug. 28 100% TABLE 50 Stadia of Scolopostethus atlanticus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10 5 4 4 8 12 43 Some females are already fertilized by April 22, but eggs are not laid until after 6 days in the laboratory. A sexually isolated unmated female laid only 4 eggs. The fecundity in the laboratory is largely due to the extended oviposition period. One female laid 208 eggs at an average rate of 5 eggs a day over about 40 days. The eggs are similar to those of 8. diffidens and are laid at ran- dom or pressed into crevices in the litter. The eggs do not stick to each other and are often laid into wet substrates. 110 ENTOMOLOGICA AMERICANA Scolopostethus thomsoni Renter Such is the biological and morphological diversity of the New England populations of thomsoni that this study is inadequate and a more detailed study will be needed to understand this species. It is not clear whether the various populations represent different species, ecophenotypic groups, or genetic aggregates separated by ecological barriers. No attempt was made to study mating reactions among the different populations. Eyles (1963, in litt.) also has found considerable diversity in the habitats of 8. thomsoni in Europe. He has intensely studied several populations of this species in England. In North America thomsoni has a generally northern distribu- tion and is known in high altitudes south to New Mexcio. I have collected it on a cool grassy bald at 6,000 feet on Roan Mountain, North Carolina. There are specimens from Anchorage, Alaska in the United States National Museum and thomsoni is recorded from Lappland and Siberia in the Palearctic as well as from Algeria and Morocco. This wide spread distribution and its far northern ex- tension along with the biological diversity of 8. thomsoni in North America makes it seem likely that thomsoni is one of the few lygaeids with a Holarctic distribution. It is conceivable that both native and introduced populations may be present in New England. Environment The population studied in New England may be separated into three or four groups. The first includes the common Scolopostethus found around sedge “ stools’ ’ (Carex stricta Lam.) in marshes sometimes with Ligyrocoris caricis and Pachyhrachius albocinctus. (See under L. caricis for a description of this habitat at Storrs, Connecticut). This habitat agrees closely with that described by Torre-Bueno (1917) for his “atlanticus.” He considered this the most common Scolopostethus in New York and swept it in large numbers from sedges. His records undoubtedly refer to this pop- ulation of 8. thomsoni. This is an unusually wet habitat for a Scolopostethus species and sometimes individuals are seen running across water on bits of litter between the sedge clumps. This form was collected in large numbers in flood wash left by high water at Mansfield Center, Connecticut. This area contains extensive stands of sedges under which occurs this Scolopostethus. It was also col- lected in marshes at Norfolk and Canaan, Connecticut. This is the population that may key down to 8. atlanticus because the hind femora are pale and the proximal small spines on the fore femora are often reduced or absent. Although it is commonplace for northern species to occur at 111 VOLUME XLIV their southern limits in marsh or bog habitats (Allee et at., 1949), there are no equivalent Car ex habitat records in the European literature. The remaining populations of 8. thomsoni are collectively re- ferred to as thomsoni “B” in the general discussion. Actually it too consists of at least two groups here termed “ B ” and “ C. ’ ’ The second group “B” is found in northern New England in mesic roadside habitats, often among rank forbs and along woodland margins and sandy flood plains. This habitat probably corresponds to the collections of 8. thomsoni in roadside trash near Buffalo, New York (Van Duzee 1894) and resembles the typical habitat in Eu- rope where thomsoni is found abundantly on nettles ( TJrtica ) (Southwood and Leston 1959). However, there are no records of thomsoni on nettles ( TJrtica ) in North America, nor have I found thomsoni on or about TJrtica in New England despite specific search. S. thomsoni “B” is the most contrastingly colored population, and is the most clearly “thomsoni.” However some individuals have the proximal fore femora spines reduced, especially in several pop- ulations found in Connecticut on a sandy flood wash plain. Pop- ulations in northern New England fall definitely into Barber’s concept of thomsoni. In a habitat at Laconia, New Hampshire 8. thomsoni was found abundantly with Ligyrocoris diffusus and Trapezonotus arenarius. In a transect at Laconia from a ruderal roadside to a white birch stand the numbers of 8. thomsoni “B” dropped off sharply and were completely replaced by 8. diffidens in birch litter. The third population “C” is a little larger and lighter in color and was found at Gorham, New Hampshire, in grasses under and near birch trees but not in thick birch litter. The nymphs of this form are also lighter in color. This habitat is similar to the grass- birch habitat found for thomsoni in Oslo, Norway by Eyles (1963b, in litt.). Several other habitat notes bring out the diversity of the habi- tat of 8. thomsoni in Europe. In Switzerland Hedicke (1942) notes it in wood and field edges. In England, Southwood and Leston (1959) list it from damp meadows, wastelands, woodland clearings, parks and gardens. Stys (1960) records it in Czechoslovakia on Calluna, in birch litter and Sphagnum as well as TJrtica. Lindberg (1958) records it in Newfoundland from many localities in sifting among leaves and dry grass, on vegetation, in humid and shady locations in woods and on shores. In Canada, thomsoni was col- lected in humus under pine trees (Brown 1934). One point clear about the habitats of S. thomsoni is that although they are usually open, they are wet sites with an abundant herbaceous growth. 112 ENTOMOLOGICA AMERICANA General Biology All populations of S. thomsoni studied are pterygopolymorphic, but none have the low number of macropters (4.2%) reported by Eyles (1963b) for the nettle population of 8. thomsoni. Forster (1955) also notes that the macropters are very rare. There is some variation among the New England populations but these all have at least 10% macropters. The meaning of the flightless condition is not apparent in those populations existing in rather temporary habitats, but the population of thomsoni “A” is certainly inhabit- ing a long persisting serclimax type of habitat along pond edges. The presence and abundance of S. thomsoni on nettles, a forb of temporary habitats, seems to be an exception to the relationship between macroptery and temporary habitats. Of course if the net- tles are very abundant only a small dispersal rate would be neces- sary for migratory purposes. Southwood (1960) noted that S. thomsoni in England is probably a day flier, and Eyles (1963b) col- lected only two females in air suction traps. Although the other two species of Scolopostethus are definite ant-mimics the field appearance of 8. thomsoni and its smoother, less jerky run, does not appear ant-like. The coloration pattern appears disruptive, especially the macropter, in which the mem- brane is almost entirely white. Like E. ferns some populations of 8. thomsoni possess a sweet aromatic scent gland odor quite unlike the usual “buggy” odor. This odor is present in the sedge Scolopostethus and in a Laconia “B” population. However population “C” from Gorham, New Hampshire does not possess this odor nor does a “B” habitat type form from Camden Hills, Maine. Some populations studied earlier were not tested for this characteristic. This variation underlines the diversity of “8. thomsoni.” No predators or parasites were observed in this study. Accord- ing to Thomas (1955) in England S. thomsoni is the prey of Nobis major, N. mimicoides, Lithobius sp. and Formica rufa L. S. thom- soni is parasitized by a tachinid (Southwood and Leston, 1959) and Eyles (1963a) found 7 specimens parasitized by Cirochira atra Zett. which also parasitized 8. decoratus. 8. thomsoni is also attacked by a fungus, Entomophthora sp. (Eyles 1963a). In the European literature 8. thomsoni is repeatedly reported to feed on nettles (Puton 1896, Butler 1923, Brown 1925, Murray 1935, Feige and Kulhorn 1938, Hedicke 1942, Franz 1943, Shaw 1945, Forster 1955, Southwood and Leston 1959, Stys 1960, Eyles 1963b). Eyles definitely showed that the insects feed on the seeds of the nettles. 8. thomsoni has also been collected from Mentha sp. (Puton 1894) and Tanacetum blossoms (Prohaska 1923). Eyles 113 VOLUME XLIV (1963b) found that the larger part of the population is on the net- tle plant, not on the ground. Although Torre-Bueno (1917) swept thomsoni 11 A” on sedges, I found less than 10% of the total popula- tion ever actually on the sedges, and the other populations are re- stricted to the ground. The feeding habits of the New England 8. thomsoni reflect the population diversity already seen. The sedge population feeds on Car ex and Scirpus seeds to the exclusion of other seeds offered it. Nymphs reared from eggs on a Carer-sunflower seed mixture fed exclusively on Car ex seeds ignoring the sunflower seeds unless starv- ing. In contrast populations of “B” thomsoni feed readily on a larger variety of seeds, including sunflower seeds. A “B” popu- lation at Canaan, Connecticut feeds on seeds of Oenothera, Bumex and Mentha. Populations from northern New England feed on Budbeckia, Tsuga and Betula seeds. Specimens of 8. thomsoni clash intraspecifically over the seeds as do the other two species, and drag the small seeds about or suspend them under their body on the beak. The Canaan population also somehow attaches the Oenothera seeds to the sides and top of the glass rearing dish. On at least ten occasions a thomsoni which was not feeding on the seed drove away another Scolopostethus ap- proaching the seed. Several times the fore femora were raised at the appearance of an aggressor. This behavior pattern was not ob- served in the other populations. Life History Although Eyles (1963b) detected only one generation a year in 8. thomsoni in England, the populations in southern New England are bivoltine. Although population “C” from Gorham, New Hampshire probably overwinters as adults and late instars as does 8. thomsoni in England (Forster 1955, Eyles 1963b), the only overwintering form found in populations “A” and “B” in Con- necticut is the adult. No phenology table can presently be given for 8. thomsoni be- cause no one population could be sampled through the entire year and the different populations are not ecologically or climatically equivalent. Some comparative notes can be given, however. The nymphs of thomsoni “A” population at Storrs do not appear until early June, but in the thomsoni “B” population of the sandy flood plain at Canaan, Connecticut, first instar nymphs are present in the middle of May and become adult in late June. The new adults of the thomsoni “A” population appear in the middle of July. This difference may result from the different insolation levels of these habitats. 114 ENTOMOLOGICA AMERICANA Instead of remaining “immature” as does the English 8. tliom- soni, the first generation of Connecticut populations both in the laboratory and in the field become reproductive after a preoviposi- tion period of 8 to 10 days. This is also true of a Laconia popula- tion “B” in late July and early August and a Gorham, New Hampshire population “C” in mid-August. The second generation of both “A” and “B” populations of 8. thomsoni remains nonreproductive or immature. This however is not true of the late August Gorham “C” population in which the new adults in September become reproductive in the laboratory and produce a third generation. In contrast, late August adults of Camden Hills “B” population do not become reproductive. Therefore only population “C” lacks a reproductive diapause mechanism and overwinters in a quiescent state. That the other populations have a facultative reproductive diapause and not merely an ‘ ‘ immature ’ ’ period is shown by the short preoviposition period of the first generation in contrast to the long reproductive diapause of the second generation. Because this occurs in the laboratory under warm conditions the diapause condition is prob- ably a photoperiod response. This would account for a few ap- parently first generation adults which go directly into diapause. Since the English climate is cooler, the univoltine condition may not be obligative but facultative, the single generation becoming adult in late summer. The diapause state under warm laboratory conditions is broken by only part of the population in January and so lasts at least four months. The table of stadia given here (Table 51) is based on the Scolo- postethus “A” population on Car ex seeds. Under laboratory con- ditions development occurs more rapidly than in Eyles ’s work prob- ably because of the higher room temperature (78° F.). TABLE 51 Stadia of Scolopostethus thomsoni Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 10.0 5.8 5.0 4.7 5.3 6.8 39 (8-12) (4-7) (3-7) (4-5) (4-7) (5-10) (31-50) As already mentioned the preoviposition period of the first generation is 8-10 days. After being brought into the laboratory 115 VOLUME XLIV in spring on April 14, the flood plain population “B” females are already fertilized and oviposition begins immediately. Females from the Scolopostethus “A” population are not fertilized at this time. As in S. thomsoni in England virgin females of the Car ex popu- lation laid a few eggs (7-20, mean 16). The fecundity varied from 104 to 250 eggs laid at an average rate of 6 a day in the thomsoni “B” population. The eggs are similar to those of the other Scolopostethus. They are laid in substrata available. Population “A” definitely prefers to lay the eggs in wet substrates while the other two populations lay eggs in both dry and wet substrates. The wet substrate pref- erence correlates with the wet habitat of thomsoni “A.” Eyles (1963b) found eggs of the nettle population laid in the perianth of the nettle seed heads. Eremocoris ferus (Say) The genus Eremocoris includes 30 species, of which 11 are Nearctic, 16 Palearctic, 2 Oriental but from high elevations in India, and 1 in the Ethiopian region. This is in general then a Holarctic distribution. One species angusticollis Jak. from Siberia was judged by Horvath (1883) to be close to E. ferus. In the eastern United States there are three species recorded of which two are southeastern in distribution. E. ferus has the widest distribution and occurs completely across North America from Newfoundland and Hudson’s Bay to British Columbia and south to California, Texas, Louisiana, and South Carolina. (Slater, Cata- logue). There appear to be no records from the extreme south- eastern United States. Nevertheless, this is a very broad distribu- tion indeed. Unfortunately it was not realized until the present moment of writing that Blatchley’s (1926) Barber’s (1928c) and Torre-Bueno’s (1946) concepts of Eremocoris ferus are composite and ferus ac- tually includes two distinct species. One species is northern, the other southern. The two species can be easily distinguished. The northern species has the hind tibia nearly nude except for two rows of small moveable spines ; the body, including the abdomen is almost nude ; and the labium attains only the metasternum or the mesocoxae. The southern species has the hind tibia densely covered with many fine erect long setae ; the body is rather pilose, especially the ab- domen ; and the labium attains the hind margin of the metacoxae, or 116 ENTOMOLOGICA AMERICANA the abdomen. The last instar nymphs may also be told apart by the setae of the hind tibia and the length of the labium. The color patterns of these two species are very similar, and the southern species is slightly smaller. Within the study area the northern species occurs south to Canaan and Storrs, Connecticut but not to Noank and New Haven, Connecticut. The southern species occurs at Noank and Storrs but not in Canaan and farther north. Specimens before me from Quebec are all the northern species. This is not a subspecies pat- tern because the two specific forms coexist as separate populations with no evident overlap at Storrs, Connecticut. Indeed, the southern ferns in Connecticut differs only slightly from specimens of ferns from Texas. Since ferns is one of Say’s species the types of which were de- stroyed and a lectotype has not been selected for E. ferns, it is not certain which species should carry the name ferns. Although Say gave “the Union” for the locality of E. ferns, the austral species is probably the one that he described, and the boreal form is the new species. The name borealis Dallas presently a synonym of ferns may refer to the northern species. Environment In the discussion on wing polymorphism E. ferns is noted as having a very wide ecological distribution. The resolution of E. ferns into two species is reflected in their more restricted ecological distributions, although the variety of habitats exploited by each is still considerable. The two species are similar in that both inhabit shaded or cool habitats and are restricted to loose ground litter except during spring dispersal flights. The northern species, like Scolopostethus diffidens is a common inhabitant of the litter beneath the northern forest trees hemlock ( Tsuga canadensis ), spruce ( Picea rnbens), and birches ( Betnla papyrifera, B. Intea Michx. and B. popidifolia) . Drake’s (1922) record of it under yellow birch {Betnla lutea) probably refers to this species. It is also found in open but cool habitats at high ele- vations. Adults were collected on the alpine area on Mt. Washing- ton at 6200 and 6000 feet. A little lower, at 5500 it was collected in V accininm heath with Trapezonotns arenarins. At lower elevations it is frequently found in forest edge habitats. At Storrs, Connecti- cut it is found in several cool ravines where hemlock stands exist, and is much less abundant and widespread than farther north. The ecological distribution of the southern species is more heterogenous and difficult to characterize. It occurs at Noank, Con- 117 VOLUME XLIV necticnt in the litter under V actinium corymbosum (L.), Viburnum dentatum L. with Scolopostethus atlanticus (see Ozophora for a description of this habitat), under the litter of arbor vitae ( Thuja occidentalis L.), and under bayberry (Myrica pensylvanica Loisel). Under gray birch ( B . populifolia) at Storrs, it is much less abun- dant than the northern species is under this same subclimax tree at Canaan, Connecticut. In small scattered groups it occurs in light mesophytic woodlands of Acer rubrum L. and Betula lenta L. with Drymus unus. It is even occasionally found along shaded beach strands, or the litter under roof eaves. The species does not occur in open dry grassy habitats, or among rank forbs, or in xeropyhtic oak-hickory forests. The litter of austral ferus is usually drier (4-8) than the boreal ferns which sometimes, as on Mt. Washington, New Hampshire occurs in completely water-soaked (10) sedge-sphagnum environ- ments. The driest habitats of the boreal ferus are some well drained hemlock slopes (4). For the northern form the soil and litter varies from the gray podsol soil — mor litter under hemlock and spruce to brown mull soil with a deep leaf litter under birches. For the southern form the litter is usaully loose and well drained, soil variable. In both the presence of field colonies corresponds to small edaphic changes, but as in Scolopostethus a degree of gregariousness is apparent. Both species are occasionally found in great abundance within their preferred habitats. The northern ferus occurs in a density up to 50-60 per square meter in Picea-Betula litter, and the south- ern form in a similar abundance in V accinium-V iburnum litter. Most of the available habitat notes, judging from their distribution, refer to the austral species. Torre-Bueno (1929a) collected E. ferus around a marsh under cattails. The species was found under cover in upland woods and in the summer on the ground in sandy places (Blatchley 1926). Froeschner (1944) noted that f erics is uncommon in Missouri and found in weedy fields. General Biology Both species are completely macropterous. At least the south- ern form flies readily in dispersal flights. There is a definite spring migration phase which takes place the first warm days in early spring. This dispersal occurs during the day and can be easily sampled by inspecting sheets hung up to dry at homes near wood- lands. The dispersing Eremocoris are attracted to the large white sheets. Other records demonstrate that Eremocoris disperses. Torre-Bueno (1914, 1915) and Myers (1926) found Eremocoris in 118 ENTOMOLOGICA AMERICANA large numbers washed up in beach litter. Glick (1939) collected E. ferns in the air at 200 feet over Louisiana. Eremocoris ferns is not only found very frequently with Scolo- postethns, but like it, displays a “flash” type ant mimicry when disturbed in the field. However, Eremocoris is twice (6 mm.) the size of Scolopost ethns. The boreal form is found only with S. diffi- dens, the austral with S. atlanticns and S. diffidens. The northern form is much more definitely associated with Scolopost ethns. The nymphs of instars one to four are also ant-like because the red ab- domen has a transverse white band across the anterior abdominal segments. The fifth instar is dark colored. E. ferns feigns death for a brief period of 10-20 seconds when disturbed directly. The scent gland odor of Eremocoris is not the usual “buggy” odor but a strong sweet aromatic odor which is readily released at slight disturbances. This odor is similar to that found in some pop- ulations of S. thomsoni but not in the Scolopost ethns with which Eremocoris is often associated. The nymphs have the usual “buggy” odor. As yet no parasites or predators are known. Ashlock (in litt.) has reared a Catharosia from an undescribed western species of Eremocoris. There are many references in the European literature relating Eremocoris abietis (L.) (Puton 1876, Wassmann 1889, 1894, Guide 1937, Franz 1943, et al.) to ants. I have found no unusual relationships between the eastern Eremocoris and ants. Both species are usually found in litter devoid of plant material, and feed on the fallen seeds from the trees or shrubs. The northern form feeds on seeds of Betnla spp., Tsnga, and Picea. The south- ern form feeds on seeds of Myrica , Vaccininm , Gaylussacia and Thuja, and in the laboratory also on the seeds of Tsnga and Betnla. The insects actively drag seeds about in the rearing dish to pro- tected sites for feedings, so that the seeds are frequently brought together in a small area under a bit of litter where the bugs aggre- gate. The seed possession behavior is well developed in the adults and a feeding Eremocoris endeavors to keep the seed away from other insects. Several times an adult would even climb to the un- derside of the top of the cage and suspend the seed on the end of its labium. The nymphs are more tolerant and several may feed together on the same seed. Only data on the mating behavior of the southern species is avail- able. The male, on sensing by contact a receptive female will slowly approach her. The apical three antennal segments are held at a right angle to his body and parallel to the floor, and the male vibrates his antennae very rapidly. The antennae vibrate in a small horizontal plane on the joint between the first and second 119 VOLUME XLIV antennal segments. The antennae are vibrated in bursts while the male climbs on the female and effects copulation. The male drops off into the usual end to end position. When approached by a male a female easily decamps or merely holds the male off with repulsing movements of her hind legs. Copulation is accomplished most fre- quently while the female is feeding. Copulation lasts for at least several hours. Although it may be repeated, copulation does not occur with the frequency observed in several rhyparochromines such as in Stygnocoris. Although no deliberate mating experiments could be run, by accident two occurred during experiments to determine prerepro- ductive periods. A boreal ferws male from Canaan, Connecticut was placed with a 25 day old austral female from Storrs, Connecti- cut. After 28 days the female died after having laid only four infertile eggs. This small fecundity is typical of unmated females. This same male was then placed with a teneral virgin austral female and after 20 days only six infertile eggs were produced. An austral male was then supplied and normal oviposition ensued. In the other case a boreal male from Laconia, New Hampshire was placed with two unfertilized austral females from Storrs, Con- necticut. As before, one female produced only 7 infertile eggs. The other female however, produced a total of 65 eggs of which 42 did not develop. The 23 hybrid progeny could not be reared past the third instar. Since I failed to recognize that two species were involved at the time, these peculiar results were merely dismissed as due to abnor- mal males, despite the males’ healthy appearance and behavior, and despite normal reproduction in the source cultures. These results need substantiating, but as they stand, indicate a significant degree of reproductive isolation between the boreal and austral forms. Life History Most of my data refers to the southern species, but enough is available on the northern species to show that the life histories are similar. Both are bivoltine, have no diapause mechanism, and overwinter in a quiescent state as adults, and in a few cases, as nymphs of the last three instars. Cool temperatures in autumn bring about a cessation of reproductive activities in late autumn although nymphal development continues. As a result early instars are not present in the population at the advent of cold weather. As usual in drymines, the adults are quite cold hardy and move about and feed under 6° C. conditions. It is not known whether or not the females may survive the winter already fertilized. Two females collected on March 12 and isolated, laid fertilized eggs, but 120 ENTOMOLOGICA AMERICANA copulation begins very early in spring so this is inconclusive. The phenology given in Table 52 is for the population of the southern species at Storrs, Connecticut. The northern species at Canaan shows a similar although later phenology but is incomplete and so not given here. The two generations completely overlap in August and September. Different small populations of the austral ferus vary completely in their instar composition in late summer during the second generation because of differences in the coloniz- ing of new habitats and the maturation of the females. The figures then represent a pooling of the data for the populations. When brought into the laboratory any time from spring to autumn the adults become or remain reproductive. E. ferus can be readily reared on sunflower seeds but suffers a very high mortal- ity especially in the early instars (Sweet 1960). Mixing birch or hemlock seeds with sunflower seeds greatly reduces the mortality in the early instars. Under high densities there is a clear-cut fall off in oviposition and frequency of copulation and increased mortality, despite food availability. TABLE 52 Phenology of Eremocoris ferus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult Apr. 15 i% 4% 95% May 15 2% 98% June 1 20% 5% 1% 74% June 15 30% 20% 15% 35% June 25 5% 30% 45% 15% 5% July 7 10% 20% 20% 30% 10% 10% July 20 10% 20% 50% 20% Aug. 15 20% 20% 15% 10% 10% 25% Sept. 7 15% 10% 15% 20% 20% 20% Sept. 21 10% 5% 10% 10% 15% 50% Oct. 7 5% 5% . 15% 75% Oct. 22 3% 7% 90% The stadia given in Table 53 again is for the austral form only, and represents the first generation reared in the laboratory at room temperature on a mixed birch-sunflower seed diet. Nymphs brought in from the field and reared grew faster than the averages given. 121 VOLUME XLIV TABLE 53 Stadia of Eremocoris ferns Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 9.8 6.0 5.6 6.4 7.4 7.9 46.0 (7-11) (5-10) (5-7) (5-8) (5-11) (5-11) (35-58) The nymphs with swollen abdomens feed very little and remain quiescent. Five newly moulted fifth instar nymphs of the northern species were allowed to feed on Tsuga seeds for twenty-four hours and then isolated with no food. By this time the abdomens were fully distended. After five to six days all moulted into normal sized adults. This clearly indicates that the feeding period can be restricted to a short period during the stadium. The adult lon- gevity varied from 28 to 112 days (mean 71). The larger extreme was an exceptional female which, however, laid no eggs for the last 42 days of her life span. The normal preoviposition period is 10.5 days (range 10-12). The precopulatory period could not be determined precisely; it is no longer than the preoviposition period. After being brought into the laboratory in March, 5 to 11 days (mean 8) days elapsed before oviposition occurs, although copulation in some cases had already occurred in the field. Adults collected in late February are not fertilized. The fecundity is high and varies from 148 to 291 eggs (mean 198) and is probably higher. Only one mating is necessary to fertilize the entire complement. Unmated sexually isolated females of both the austral and boreal forms lay from none to 11 eggs (mean 5.6). The eggs are laid singly at the rate of 4 to 6 a day. The eggs are typical drymine eggs rather stubby and cucumber shaped, with no spinules or any sticking mechanism. They are laid in loose litter, sand, cotton, both wet and dry with a distinct prefer- ence for dry substrates. Eremocoris setosus Blatchley Blatchley (1926) described this species from a manuscript of Barber’s as a distinct species. Barber (1928e) described setosus as a subspecies of the European E. plebejus Fallen distinguished by its even denser coating of long soft hairs. If setosus is a distinct subspecies of plebejus this would be a remarkable disjunct distribu- 122 ENTOMOLOGICA AMERICANA tion. Although Barber (1928e) recorded specimens of this south- eastern species from Tyngsboro, Massachusetts and Long Island, N.Y., I have been unable to locate the species is New England. There is also a record from Montreal, Canada (Moore 1950), but otherwise, setosus is restricted to the southeastern seaboard. Barber (1928e) found setosus abundant near Vienna, Virginia in sifting dead leaves in or about the woods. Blatchley (1926) took the species in Indiana from beneath logs on slopes of upland wooded pastures, and swept it from boneset and other Compositae. Gastrodes walleyi Usinger The genus Gastrodes includes 9 species, five in the Nearctic, three in the Palearctic, and one in the Oriental Region at Macao. Of the Palearctic species, one is restricted to Japan and northern China, and the other two are widespread Palearctic species. Four of the Nearctic species are restricted to western North America. The fifth and only eastern Nearctic species G. walleyi was described by Usinger (1938) from a single mutilated specimen collected by G. Beaulieu on July 1, 1914 at Ottawa, Ontario, Canada. This locality makes it probable that G. walleyi occurs in northern New England. It is evidently quite rare as it has not been collected again since the first collection and Kelton (in litt.) at the Canadian Entomology Research Institute, Ottawa, Canada has been so far unsuccessful in locating the species. The available records indicate that the species of Gastrodes feed in the trees on the seeds on conifers. The two common European species have been studied (Nageli 1933, Aitkens 1936, Pfaler 1936, Southwood and Leston 1959). G. abietum (Bergr.) is specific to Norway spruce ( Picea abies L). while G. grossipes is abundant on Scots pine ( Pinus sylvestris L.). G. conicola was collected in Cali- fornia on resinous mature cones of the digger pine ( Pinus sabiniana Dougl.) (Usinger 1933). The rarity of walleyi may be in part explicable by Usinger ’s note on G. conicola: “No specimens had ever been found during previous years of collecting nor, upon diligent search were any specimens later found to occur in the foliage or in old cones of previous seasons.’’ It is probable then, that G. walleyi will be found on a coniferous host, feeding on the seeds. I have searched without success in Maine and New Hampshire for it on the new and old cones of Pinus strobus L., Pinus rigida (Mill.), Picea rubens Sarg., Picea mariana (Mill.), Picea glauca (Moench.), Picea abies (L.), Abies 123 VOLUME XLIV balsamea (L.), Larix laricina (DuRoi and Tsuga canadensis (L.), but it probably occurs on one of these hosts. Pinus banksiana Lamb, is another possibility. TRIBE RHYPAROCHROMINI Peritrechus Fieber This large genus has a Holarctic distribution with the majority of the species in the Palearctic region. Of 26 species, 22 are from the western Paleactic, but apparently none are recorded from the eastern Palearctic — China, Japan, and Korea. This apparent faunal break in the eastern Palearctic is probably a collecting gap as a number of European species range quite far north, and four, P. angusticollis Sahib., P. convivus Stal, P. distinguendus Flor, and P. geniculatus Hahn, are recorded from “Siberia” (Slater, Cata- logue). Lindberg (1958) reported a colony of distinguendus on Newfoundland and considered it to have been introduced there by way of ballast from European ships. Of the four Nearctic species, two are western, one is widespread, and the other is limited to the eastern sea board. It is reasonable to consider the genus as a Palearctic one which has spread into the Nearctic region. Peritrechus fraternus Uhler Peritrechus fraternus is a widespread species recorded from Mexico and lower California north to British Columbia and Alberta and east to Quebec and New Hampshire, but in the eastern United States it is not recorded south of New Jersey (Slater, Catalogue) except in the mountains of North Carolina at Spruce Pine (Brimley 1938). I have collected the species in the same county, Mitchell, at 6,000 feet on Roam Mt., North Carolina. It appears then that the eastern and western populations have a quite different north- south extent. It would be important to ascertain if a species com- plex may be involved. Since the type material of P. fraternus was originally described by Uhler (1871) from Massachusetts, it is very probable that, whatever the species situation may be, the population in New England represents this species. 124 ENTOMOLOGICA AMERICANA Environment While it is ecologically restricted, Peritrechus fraternus is one of the more common species of rhyparochromines in southern New England. The great majority of the collections of this species are from wash litter communities. Since the wash litter is a concen- tration point for various seeds, it forms a favorable habitat. The species is found in strand wash at the high tide mark along the ocean shore at Rocky Neck and Noank, Connecticut and at Woods Hole and Brewster’s Point, Massachusetts; in litter along the shore of Lake Cayuga, New York, and Twin Lakes, Connecticut ; and in the flood wash of various streams. In the salt water beach wash it is found only at the highest strand levels, above the level dominated by the sand flea Orchestia, where the substrate moistness is mesic, about 5-6, and the salt content is low. At Brewster’s Point it is also found on the high beach in wind blown litter at the bases of seaside goldenrod ( Solidago sempervirens L.), beach rose ( Rosa rugosa Thunb.), beach plum ( Prunus maritima Marsh.), beach pea (Lathyrus japonicus Willd.), and poison ivy ( Rhus radicans L.). In similar inland litter aggregations, especially where large litter layers are deposited, this species attains very dense populations at some sites, up to 86 per square meter, although at others as low as less than 1 per square meter. Such lowr abun- dances are usually in old litter lines, the highest in fresh litter deposited during spring flooding. Open or semi-open areas are favored and heavily shaded strands contain very few Peritrechus. Nearly always, probably because of the nature of the site of litter deposition, the soil is sandy or gravelly. Repeatedly P eritrechus concentrates in a mesic zone of intermediate moisture (4—6) and temperature (70-90°F.) in the litter. At times such favorable aggregations of seeds attracts other rhyparochromines such as Perigenes constrictus , Heraeus plebejus, Pachybrachius basalis, and Scolopostethus thomsoni (A), but always in the strand litter bio- tope Peritrechus is dominant. It also occurs in the similar aggrega- tions of dry litter along some new roadsides. Sometimes it is found in low densities in new sandy often ruderal habitats, but here, it is always an infrequent member of this community which is usually dominated by rhyparochromines such as Ligyrocoris diffusus, Pachybrachius basalis , and Emblethis vicarius. Occa- sionally I found this species on small rises and hummocks in salt marshes along with Peritrechus paludemaris Barb, (see P. palude- maris for further discussion). On Roam Mountain, North Carolina, I collected a population at 6,000 feet on an open cool bald dominated by Rumex acetosella and fescue grass Festuca spp. While the area had been maintained in 125 VOLUME XLIV this condition by man’s activities, the habitat is so very different from the New England habitats that the question must be raised as to whether or not this mountain population is conspecific with the new England population. Perhaps it is a latitudinal variation in habitat selection. Since this mountain population is slightly larger and darker but otherwise very similar to the New England population it is unfortunate that I was unable to cross the two popu- lations because of diapause interference. At inland sites, Peritrechus overwinters in the same habitats in which it matures, but along the seashore there is a definite retreat nearby to drier, more protected sites away from the strand margin. Many of the following notes in the literature are too generalized to compare with my findings. Van Duzee (1894) collected it hiber- nating under dead leaves. He (1914) also collected what is sup- posedly this species on low flatland between the bay and the Pacific Ocean at San Diego, California. Smith (1910) collected it in a cranberry bog. Blatchley (1895, 1926) considered it scarce in Indiana and collected it under chips and dead leaves in fall, and under stones and other cover in winter. A habitat perhaps related to the wash litter is the drop seed habitat in which Walkden and Wilbur (1944) found Peritrechus is Kansas. Froeschner (1944) lists it as common in Missouri under rocks, logs, and in grass clumps. The strand and flood litter habitats, while their general presence is a permanent situation, the individual aggregations of litter, espe- cially on floodwash sites, are temporary biotopes. General Biology P. fraternus is entirely macropterous, and no brachypterism is known as is apparently true of the other species of the genus. It is not surprising that it is entirely macropterous, with such an obvious requirement to disperse to new habitats. The species may disperse, at least on occasion, at night for Hussey (1922) collected 30 at a lighted sheet at Devil’s Lake, North Dakota. Knowlton (1960) collected it in Utah at lights. Froeschner (1944) notes that in Missouri it may sometimes come in numbers to light. As is so frequent among lygaeids the nymphs and adults are quite differently colored. The adults are dark black and mottled brown and blend extremely well, when quiet, into the litter and sand. However, all nymphal instars are a rather bright pale red- dish color which may render them conspicuous to bird predators. When disturbed these insects move rapidly and agilely hide under some concealing object. They may '‘freeze” but do not feign death like Eremocoris and Stygnocoris. Their normal move- 126 ENTOMOLOGICA AMERICANA ments are fairly quick and active in contrast to Peritrechus palude- maris. These insects are prone to aggregate together under a bit of litter cover, and are reared much more readily in the laboratory if cover like crumpled methyl cellulose is provided. I have not discerned any known predators although it may clearly be subject to predation by the large numbers of predatory carabid and staphylinid beetles in the wash litter. Parasitism by Catkarosia sp. tachinid flies is very low ; one male specimen of a long series of adults and nymphs from Rocky Neck, Connecticut was found parasitized. When litter from its biotope is brought into the laboratory along with it, P. fraternus oviposits readily and is easily reared to maturity on the seeds contained in the litter. This is equally true if the other arthropods are either present or killed by baking. Among the profusion of seed species, seeds of Panicum spp. are dis- tinctly preferred; other seed species are probed at, but are not fed on except for some Solidago seeds. On sunflower seeds growth is excellent with little mortality up to the fourth and fifth instars when growth becomes quite slow and the mortality very great. While the water requirements are high, P. fraternus may survive for as long as 24 hours without water. When Panicum seeds are placed on a sheet of methyl cellulQse, they are subsequently dragged by the insects to concealed positions beneath the sheet. As in L. diffusus this species exhibits a definite defense reaction or display when one individual intrudes on another which is feeding. The defender first wags its antennae rapidly and alternatly, and endeavors to position its body between the intruder and the seed. If this fails to discourage the intruder, the defender either drags the seed away, or it ceases to feed, and, with antennae wagging and fore legs flailing, it charges the in- truder. The intruder may move away or persist, in which event the charge is repeated. In each observation the defender wins. It is remarkable that after such a scuffle, which may carry the insect two or four inches away from a seed, the exact same seed — among many — will be searched out and the exact same hole pre- viously drilled is found. Because of the intense diapause of this species and the essentially univoltine life cycle of much of the population, mating behavior could not be extensviely studied. The male becomes sexually excited when a female comes near, and recognizes her without touching her with his antennae. Recognition is shown by the male’s extending his motionless antennae towards the female. The male advances by suddenly leaping on the female, and positioning himself parallel to the female and tapping her head with his rapidly 127 VOLUME XLIV vibrating* antennae. In most observed encounters tlie female escaped the male. Under caged conditions there may frequently be 4—6 males around a female. On several occasions in the field simi- lar aggregations were found. Occasionally a male would leap upon a male. A male which had matured before the diapause point, is sexually excited by diapausing females of both P. fraternus and P. paludemaris. After advancing several times, however, the male lost interest in the diapausing females, while fertile females continue to elicit a reaction. Life History Peritrechus fraternus was observed to overwinter by Van Duzee (1894), Blatchley (1895, 1926) and Froeschner (1944). The observed phenology at Storrs is as in Table 54. It is significant that some early instars occur in the middle of July, for they represent a partial second generation. Ordinarily these nymphs might be thought of as progeny from an adult which lived exceptionally long. However, it was found that the insects which become adult in late June or early July (at least before July 9) become sexually reproductive and lay a small number of eggs until after about July 15 when such egg laying ceases. The adults which mature near this time go directly into reproductive diapause. The best hypothesis appears to be that some photoperiod effect initiates diapause. Most of a given population in New England does not undergo a partial second generation especially at inland sites. At a Rocky Neck salt marsh margin a considerable number did in 1960 (see paludemaris for further discussion). The phenology given in table 54 is pooled from the populations studied at Noank, Rocky Neck, and Mansfield, Connecticut. How- ever, one population progressed differently. At Storrs, Connecticut, on a grassy, south facing roadside bank strewn with debris, a popu- lation entirely of fifth instar nymphs was found on September 10, 1958. These became diapausing adults in mid- and late September. This may represent the partial second generation. The diapause condition is an intense one and is not broken by short cold exposures (one month), photoperiod conditions, or ele- vated temperatures (32°C.). The cultures do not survive longer cold exposures, nor does diapause development complete itself dur- ing the winter in laboratory warmth as in other species. Only the few cultures which survive in the laboratory until middle April become reproductive which is nearly the same time as the field popu- lations. It thus appears that diapause development will go to completion in this species over a long period of warmth. The occurrence of the partial second generation in early maturing adults 128 ENTOMOLOGICA AMERICANA suggests that the diapause state is initiated by shorter or shortening photoperiods, but no experimental proof is at hand. The stadia in table 55 are rather variable in later instars espe- cially those that feed on sunflower seeds. The very prolonged development of some nymphs is presumably abnormal and results from some limitation in the caged conditions. Nymphs brought in from the field and reared develop much more rapidly. If Panicum seeds are provided, the newly hatched nymphs develop much more normally and with a smaller mortality. The earlier instars, how- ever, develop equally well on sunflower seeds as Panicum seeds. TABLE 54 Phenology of Peritrechus fraternus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult Mar. 15- Apr. 27 100% June 1 8% 23% 15% 54% June 11 5% 9% ■ 38% 28% — 20% June 24 21% 32% 35% 12% — July 3 7% 13% 35% 45% July 11 21% 43% 15% 21% July 16 4% 22% 74% July 25 — 6% 29% 65% Aug. 5 2% 98% Aug. 10 100% on The average longevity of non-diapause adults is 52 days. The diapause adults in the field live through the summer and winter to spring and die in June. Those kept in warmth frequently live nearly that long (278 days) but most die before early spring without laying any eggs. With the few adults which mature before diapause conditions set in, the preoviposition period is about 4 days. Adults collected on April 15 are already reproductively active, mating, and contain ripe ova. Eggs are laid by spring adults a few days after being brought into the laboratory. The early spring females laid in the laboratory from 84-386 eggs (average 193). The partial second generation females laid 78-104 eggs (average 129 VOLUME XLIV 83). The rate per day is 6.1 and varies from 4-8.6 eggs. Most sexually isolated virgin females which overwintered laid no eggs except for one female which laid 79 at a low rate of 2 per day. The two virgin females which became adult at the time that the partial second generation females became reproductive, laid no eggs and later went directly into diapause. The eggs are laid singly into litter crevices, hollow stems, dry soil, and in tight cotton and methyl cellulose. The females spend a considerable time selecting oviposition sites. The elongate smooth eggs are sticky when laid and stick tight into crevices, to litter fragments, glass, and to methyl cellulose. The eggs often are com- pletely covered with grains of sand and litter fragments, which renders them difficult to see. TABLE 55 Stadia of Peritrechus f rater mis Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 7.2 (6-9) 5.3 (4-7) 5.9 (4-8) 6.6 7.7 12.2 (5-8) (5-11) (6-18) 43.5 (27-70) Peritrechus paludemaris Barber Barber (1914b) noted this species as being very closely related to fraternus , but many species of the genus Peritrechus are rather similar appearing, and actually paludemaris is quite different. Barber used the relation between the distance across the anterior angles of the pronotum and the width of the head to distinguish the species. In practice this character is difficult to use and resulted in some misidentifications. P. paludemaris is considerably larger, the membrane is mostly fuscous, and the antennae are much longer, especially the fourth antennal segment which is one and a half times as long as the third segment. P. paludemaris is found along the eastern sea board from Massa- chusetts to Florida (Barber 1914b). As there is a specimen in the Slater Collection from Biloxi, Mississippi, it is probable that this species is also found all along the Gulf Coast as well. The species is here recorded for the first time from Connecticut. Environment In the literature all records of this species are from salt marshes 130 ENTOMOLOGICA AMERICANA (Barber) 1914a, 1923, 1928d, Blatchley 1926) or seashores (Torre- Bueno 1931, 1946) of Massachusetts, New York, Maryland, and Florida. I collected this species in brackish and salt marshes at Rocky Neck State Park, Connecticut, and at Hammonasett State Park, Connecticut. The preferred biotope of this species is in the litter of the slightly raised hummocks in salt marshes. The marsh grass is composed largely of a single species, Spartina patens (Ait.) and some Distichlis spirata (L). on the hummocks. Other plants frequently on the hummocks are Hibiscus palustris L., Aster subulatus Michx., Pluchea purpurascens (Sw.), and in a brackish area, Spirea latifolia (Ait.). The ground moisture level of course is saturated (10), but the exposed litter is quite dry and grades off towards the ground. P. paludemaris is found at the narrow litter zone between these extremes. Temperatures are moderate, and measured at 78° F. P. paludemaris is never found on the mucky surface of the salt marsh flat, but on the hummocks which protrude slightly above the high tide level. In late autumn at Rocky Neck and at New Haven, Connecticut, several adults were found sheltered under Andropogon scoparius clumps a short distance (about 50 and 75 yards respectively) from the salt marsh. At the same time no paludemaris could be found in the marsh, despite careful search. This suggests that P. palude- maris may migrate away from the marshes to more protected sites in the autumn. The abundance of P. paludemaris is always relatively low com- pared to the numbers in which fraternus frequently occurs, and is quite discontinuous, some nearly identical hummocks with, others without insects. The greatest abundance is reached in mid- June when the population consists of early instars at abundance of 4-5 per square meter on certain hummocks. Usually the abundance is considerably less than 1 per square meter. This is the only lygaeid found on low hummocks far out in the salt marsh. In sites which are higher, drier and closer to the land margin of the marsh, P. fraternus often appears during July. During June in 1961 and 1962 at Rocky Neck, P. paludemaris was the only lygaeid on the salt marsh, and even the litter along the margin did not harbor P. fraternus which was abundant at other wash sites. However in July of 1960, 1961, and 1962 P. fraternus appeared on the drier hummocks and along the margins. As explained under P. fraternus , this apparently represents the partial second generation of P. fraternus. General Biology There probably is a seasonal shift from hibernation sites to the 131 VOLUME XLIV salt marshes, as indicated earlier. The precarious water level of the salt marsh must impose a hazard which may outweigh any advantage of brachyptery in the essentially permanent habitat of the salt marsh. P. paludemaris is similarly colored to fraternus although a shade lighter, but behaves quite differently. While fraternus moves rapidly in a scurrying fashion to take refuge under bits of debris, paludemaris moves much more slowly and deliberately, and hurries in short bursts of speed much like Myodocha and Antillocoris. It is much less active in taking refuge under objects. This difference in movements extends to the nymphs, which are quite different in appearance from the nymphs of fraternus. P. paludemaris nymphs are very densely pilose which gives them a grayish red color, and have a broad black band extending down the middle of the head and thorax. The pilosity may protect the nymphs from submer- gence at high tide. One tachinid parasite of the genus Catharosia was bred from this species. No predators are known. It has not been collected at lights. While this species feeds readily on sunflower seeds, its natural food are the seeds of Spartina patens (Ait.) and probably of other salt marsh plants. No seed defense or mating behavior was ob- served. Life History Just enough information is gathered to show that this species is univoltine, with an obligative diapause. The observed phenology is as in Table 56. Date May 25 June 10 June 28 July 11 July 26 Aug. 8 TABLE 56 Phenology of Peritrechus paludemaris Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult 100% 56% 44% 25% 50% 25% — - 25% 75% ^ 33% 67% 100% Oviposition clearly begins at least by late May. All adults are 132 ENTOMOLOGICA AMERICANA in a obligative diapause which is unaffected by rearing the late instars under short (12 hours) or long (16 hour) photoperiods. I could not break the diapause with either photoperiods, with elevated temperatures (32°C.) or a cold exposure of one month. In warmth the diapausing adults live until early spring. The available stadia based on a small sample are as in Table 57. TABLE 57 Stadia of Peritrechus paludemaris Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 6 7 8-9 10 11 Since no females were found in spring and diapause was not broken, no reproductive data is available. TRIBE MEGALONOTINI Megalonotus sabulicolus (Thomson) This introduced Palearctic species has been carried in previous American literature (including Part I of this paper) under the name M. chiragrus (Pabricius). There has been little agreement among European workers on the status of sabulicolus , and Slater and Sweet (1958) following several European authors, considered M. sabulicolus as one of the many color forms of the variable species M. chiragrus (c.f. Stichel 1925). Southwood (1963) and Roubal (1963), however, have presented evidence to show that M. chiragrus and M. sabulicolus are two dis- tinct species not merely color varieties. The two can be distin- guished by size, width of pronotum, length of antennae, paramere shape, and prevalence of brachyptery. M. chiragrus is larger with longer antennae and is usually brachypterous. As Southwood (ibid) rightly points out, the two forms are not subspecies since their ranges greatly overlap. Subspecies cannot be sympatric without intergrading, and Southwood found no or little intergrading between chiragrus and sabulicolus in England. The available literature (Slater, Catalogue) indicates that the 133 VOLUME XLIV two forms are sympatric over much of Europe, with records of sabulicolus concentrated around the Mediterranean. Hoberlandt (1955) in discussing the distribution of M. chiragrus in Turkey, designated the distribution of M. chiragrus as Euro-siberian and M. sabulicolus as Holomediterranean. He indicated that chiragrus has a more northern distribution and is found at higher altitudes in the south Palearctic than sabulicolus. It is noteworthy that the type locality of sabulicolus is in Sweden at the northern limit of the range of sabulicolus. According to Southwood (ibid), although there is some habitat overlap, in England the two species are ecologically distinct and sabulicolus is limited to sandy habitats in southern England. Since the differences are not exclusive, Southwood (ibid) raises the pos- sibility these may be forms ; environmental selection favoring a sabulicolus genotype in sandy habitats. A similar close relationship is shown among Scolopostethus species and Southwood reasons by analogy with Eyles’s (1963, in lift.) breeding work on Scolopos- tethus , that these forms of Megalonotus are good species. More- over, if these are ecological forms, more variability in response to ecological conditions would be expected than Southwood indicates for the English populations. This species pattern has a close parallel in Ischnodemus sabuleti (Fallen) and I. quadratus Fieber which ecologically displace one another (character displacement) over a broad zone of overlap where sabuleti is restricted to wet habitats, quadratus to dry habi- tats (Slater 1960). When considered in this light, the ecological distribution of the chiragrus complex should prove interesting since the area of sympatry is very large and many forms are described. The question that arises is the identity of the introduced Mega- lonotus in North America and how it relates to the chiragrus -sabuli- colus problem. The evidence for the introduction of Megalonotus into eastern North America on produce is given by Slater and Sweet (1958). Scudder (1961) considered ballast to be the prob- able mode of introduction of Megalonotus on the Pacific coast of North America. Scudder (ibid) noted that most of the British Columbia material is referable to M. chiragra sabulicola. Since 1958 I have found several additional populations of Mega- lonotus in Connecticut and one at Brewster, Cape Cod, Massachu- setts. On the basis of the coloration of the antennae and hind tibiae by which sabulicolus and chiragrus were originally recog- nized, both sabulicolus and chiragrus- like patterns occur in all the New England populations. But on the basis of the other characters given by Southwood (1963) all the New England populations clearly pertain to M. sabidicolus not chiragrus. However obvious 134 ENTOMOLOGICA AMERICANA and useful, the color variability of the antennae is so great, as is shown by the multiplicity of forms (Stichel 1925), that errors can easily result. The delimiting of the species in North America to sabulicolus further verifies that it was introduced into North America since sabulicolus is limited to the western Palearctic. The genus Mega- lonotus itself with 20 described species is concentrated in the Medi- terrean region of the west Palearctic, with only chiragrus itself having a Palearctic distribution. Since M. chiragrus is the most common species in England, and M. sabulicolus is known there from only a few localities, it is probable that the New England population came from continental Europe as suggested by the produce records (Slater and Sweet 1958). If M. sabulicolus is possibly a sandy area genotype or ecopheno- type of chiragrus one would expect the species to occur in New England and British Columbia in different forms in sandy or non- sandy habitats. All but one of the populations found in southern New England occur in sandy or gravelly habitats. Each of these populations closely fits Southwood’s description of sabulicolus and is completely macropterous. A population on Cape Cod near Brewster, Massachusetts occurs on a steep hillside covered with a low grass and partly shaded by pitch pines ( Pinus rigida Mill.). The population from this quite different habitat is definitely sabulicolus in morphological characters although 70% of the popu- lation has the third antennal segment dark as in chiragrus. Three of nine Cape Cod males examined also have a cryptic form of brachyptery in which the forewing membrane almost attains the apex of the abdomen, but the hind wings are short, reaching only tergum six. According to Southwood M. sabulicolus is usually macropterous while chiragrus is clearly brachypterous. It also may be significant that unlike Stygnocoris, Megalonotus was not found along the northern coast of New England from northern Massachusetts north to Maine despite repeated search for it among its host plant Centaurea. It is possible that the Holo- mediterranean adaptation of sabidicolus may limit it to southern New England. The New England populations of M. sabidicolus should be kept under observation to see if it extends its range and whether the M. chiragrus genotype will emerge. At present, the evidence from the New England population indicates that M. sabulicolus is retain- ing its European characteristics. Its release from competition with chiragrus and other Megalonotus species is another factor but it may be offset by competition with the native rhyparochromine fauna. 135 VOLUME XLIV Environment Several of the habitats of M. sabulicolus at Noank and Canaan, Connecticut were described by Slater and Sweet (1958). Subse- quently other sites were found as mentioned earlier. Except for the Cape Cod habitat described before, these new sites are sandy ruderal areas where the star thistle or bachelor’s buttons ( Centaurea spp.) grows. The association of sdbulicolus with Centaurea is very close and the insects are found in the litter about the base of these plants. These all constitute disturbed habitats often roadsides and with the possible exception of the Cape Cod colony, Megalonotus is not found in natural habitats. Interestingly, Centaurea itself is not only introduced from Europe but like Megalonotus also has its distributional center in the Mediterranean region (Fernald 1950). These habitats are all hot dry sites with the ground between the plants open to direct insolation. This is especially true of the site at Canaan in the cooler northwestern highlands. It was noted at Canaan (Slater and Sweet 1958) that the large population dropped off rapidly in numbers in late autumn. Re- peated observations over six years show that this results from a decimation of the population at the advent of cold weather. Only a few adults survive the winter to start the poprdation going the next spring. The adults overwinter in litter at the base of Cen- taurea. The European records given in Slater and Sweet (1958) are mostly from England and Finland and probably refer to M. chira- grus itself. Scudder (1961) notes that what is probably sabulicolus is abundant on boulevards in Vancouver, British Columbia. Ash- lock (in litt.) notes chiragrus as abundant in California and feeding on strawberries. The species is apparently much more abundant on the west coast than the east coast. These hot sandy sites are all temporary habitats which exist in one place for only a few years although such a habitat may be maintained longer by repeated disturbances along a roadside or on a shifting flood plain. The Cape Cod habitat forms an exception for it appears to be a long persisting habitat. General Biology The largely macropterous condition of sabulicolus correlates with its occurrence in temporary sandy habitats. The few cryptic submacropters from Cape Cod is an interesting development and should be kept under observation as mentioned earlier. If selected for, this population may eventually become brachypterous. The only available dispersal record is the collection of 117 136 ENTOMOLOGICA AMERICANA specimens of Megalonotus in a flight trap in 1931 in Oregon. The mottled brown and black pattern of the adults of this hairy medium sized (5 mm.) rhyparoehromine blends it effectively into the sandy background of its biotope but the black nymphs are much more conspicuous. This insect has a rather flattened shape and despite its average sized legs it runs rapidly to take cover under bits of litter when disturbed. Williams (1946) in California found Megalonotus along with Emblethis to be the prey of the astatine sphecid wasp Diplopectron. Several bugs were placed in a cell in the wasp nests which were built in sandy soil. No other parasites or predators are recorded for M. sabulicolus. The natural food plant seeds were those of Centaurea. The insect is never found on the composite heads but feeds on the fallen seeds. In the laboratory it will also feed and oviposit on Oenothera sp. and sunflower seeds both hulled and unhulled. Much better growth is had on a sunflower -Centaurea seed mixture. The insect actively drags or carries Centaurea and Oenothera seeds about by its beak to more sheltered sites for feeding. This was observed in the field as well as in the laboratory. In the mating behavior, the male responds to a receptive female by rapidly vibrating his antennae and advancing on the female with a hesitant jerky movement. The male mounts the female, aligns himself parallel and vibrates his antennae rapidly on her head. After effecting copulation the male swings into an end to end position. Copulation lasts for at least two hours. Life History According to Pfaler (1936) M. chiragrus in Finland overwinters as an adult and has two generations a year. Similarly, the New England population of sabulicolus is bivoltine and has an adult diapause. At Canaan, the few overwintered adults lay their eggs in May and early June. The phenology at Canaan is given in Table 58. At Storrs the phenology is advanced by about three weeks so that only late instars are present in early September before the advent of cold weather. In Canaan much of the second generation especially the nymphs are apparently killed by cold weather. The two generations overlap considerably, diapausing adults occurring in the field as early as August 10 and some first genera- tion adults lay eggs through August. Second generation adults go directly into an intense diapause state. Of twenty cultures only four individuals in one spontaneously broke diapause after 132 days under warm laboratory conditions. A brief 30 day exposure 137 VOLUME XLIV to 6°C. did not affect the diapause state, nor did long photoperiods in December. Adult longevity in the diapause state in laboratory warmth ranged from 125 to 185 days (mean 132 days). Such adults which broke diapause oviposited only a few eggs (17-26) and the resulting nymphs could not be reared past the second instar. No experimental work with photoperiodism was made on the diapause mechanism. The second generation diapause state occurs under warm conditions in field and laboratory. Reasoning from the condition in other bivoltine species, the diapause is probably facultative, based on photoperiod. Even when forced by early rearing the cultures did not produce second generation eggs until early July. The preoviposition period of summer adults maturing normally in July was only 5-7 days, but in a culture forced to maturity by June 16 it was 16-19 days. TABLE 58 Phenology of Megalonotus sabulicolus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 20 100% June 7 50% 30% 20% June 25 20% 30% 40% 7% 3% July 6 — 10% 15% 45% 30% July 28 10% 10% 5% 15% 60% Aug. 16 14% 14% 15% 10% 19% 28% Sept. 13 5% 5% 10% 15% 25% 40% Sept. 25 1% 19% 80% Oct. 15 5% 95% Nov. 1 100% The stadia given in Table 59 are derived from both first and second generations on a Centaur ea- sunflower seed mixture. While field collected nymphs can be reared on sunflower seeds, nymphs hatched from eggs in the laboratory could not be reared on sunflower seeds. The normal longevity in the laboratory ranged from 25 to 49 days. The fecundity in the laboratory is high, in eight females rang- ing from 129 to 215 eggs (mean 167). The eggs are laid at an average rate of 8.1 eggs a day (range 3-11 eggs). Four sexually 138 ENTOMOLOGICA AMERICANA isolated unmated females laid from none to 27 eggs (mean 18 eggs). The eggs of Megalonotus are remarkable because of their pecu- liar tack-like processes which attach the eggs to plant fuzz, ground litter, and sand. A cementing substance is also involved. The eggs are oviposited into available loose dry substrates. Wet sub- strates are avoided. TABLE 59 Stadia of Megalonotus sabulicolus Egg Instar 1 Instar 2 Instar 3 instar 4 Instar 5 Total 10.3 (9-11) 4.5 (3-7) 4.8 (3-8) 5.0 (4-6) 6.4 (4-10) 8.4 (5-13) 37.0 (27-45) Sphragisticus nebulosus (Fallen) The genus Sphragisticus is apparently monotypic. Ashlock (in litt.) tells me that simulans which Barber (1918c) described in Sphragisticus actually belongs in the tribe Gonianotini not Mega- lonotini and therefore cannot be a Sphragisticus. Sphragisticus nehulosus has a wide North Holarctic distribution. As stated in Part I, it is not clear whether Sphragisticus is an introduced species or one of the few Holarctic species. If it is a Holarctic species as thought by Horvath (1909) it is the only Nearctic representative of the tribe Megalonotini. Moreover, there are no relict populations of Sphragisticus in Mexico or Guatemala in contrast to other Holarctic lygaeid genera and to the relict popu- lations of Sphragisticus in Algeria and the Caucasus in the Pale- arctic. If it is an introduced species, it is an early introduction since Say recorded the species from North America in 1831. Its far northern distribution from Lappland to Eastern Siberia ( Jurinskij 1926) suggests that it could have made the transit from the Pale- arctic to the Nearctic. In the Nearctic it is only recorded as far north as British Columbia and Quebec, but I have seen a specimen from Fairbanks, Alaska (Univ. of Conn. Coll.). It is conceivable that Sphragisticus may have in late Pleistocene times spread natur- ally into the Nearctic which may account for the apparent lack of distinction between the Nearctic and Palearctic populations. A study of the population variability throughout the Holarctic may resolve this question. 139 VOLUME XLIV In the Nearctic Sphragisticus has a distinctly northern distribu- tion, and in the east is not recorded south of New Jersey and Missouri (Slater, Catalogue). While Hussey (1922) found it com- mon in Michigan, Froeschner (1944) considered it scarce in Mis- souri and Slater (1952) had no records of it from southern Illinois. Environment In New England Sphragisticus is largely restricted to disturbed habitats such as ruderal areas and along the edges of cultivated fields. It does not penetrate into natural habitats except in natur- ally disturbed areas, as a sandy flood plain which is in early succession. Penth (1952) in a thorough study of Ileteroptera populations in different associations at Mainz, Germany, found Sphragisticus similarly limited to disturbed ruderal habitats along roads and field edges. An arable habitat with considerable bare soil and litter does not support Sphragisticus populations as large as slightly latter succession stages where low often prostrate pioneer forbs have invaded the area. In later plant succession stages when the ground is filled in with taller forbs which shade the ground, Sphragisticus drops out. The plant associations of such favorable habitats typi- ically includes Angropyron repens, Stellaria media (L.), Cerastium vulgare (L.), Lepidium virginicum L., Capsella bursa-pastoris (L.), Polygonum spp., Silene spp., Specularia perfoliata (L.), Chenopo- dium album L., Rumex crispus L. and others. The soil of favorable areas varies from a light loam to sand, and none were found in clayey or wet soils. The litter layer is frequently very sparse except along field margins. In such open areas with low forbs the sun shines directly on the ground creating a dry hot microenvironment. In the most favorable habitats the abundance of the insect reaches 20-25 per square meter, but is more frequently around 5-10 per square meter. Sphragisticus apparently does not com- pete particularly well with the native rhyparochromine species such as Ligyrocoris diffusus , Pachybrachius basalis, Zeridoneus costalis and Emblethis vicarius and is nearly always the least abun- dant when the species occur together, except in the narrow succes- sion stage just described. The invasion, rise and disappearance of Sphragisticus was observed several times in fallow fields in Noank and Canaan, Con- necticut. It was rarely found for more than two years in a fallow field, except along edges maintained by cultivation. The literature on Sphragisticus is larger than usual because of its occurrence in cultivated fields, and several host plant records are 140 ENTOMOLOGICA AMERICANA given under the general biology which substantiates this habitat selection. Sphragisticus was found in large numbers in fields in autumn in Minnesota (Lugger 1900). It has been frequently re- corded from cultivated fields (Bruner 1891, Bruner and Barber 1894, Forbes 1900, 1905, Barber 1923). Blatchley (1926) said that it is quite common in Indiana in rubbish ; around sandy fields, espe- cially where melons have been cultivated. General Biology Sphragisticus is entirely macropterous which correlates with the temporary nature of the arable field habitats. There are no disper- sal records from New England, but Froeschner (1944) collected a specimen at lights in Missouri. The coloration of adult Sphragisticus is a mottled black and light brown which blends very well into the light, often sandy soils of its preferred habitats. The nymphs like the nymphs of Megalonotus and most Gonianotini are shiny black and appear much more con- spicuous than the adults. The insect is flattened with short legs and when disturbed it scurries rapidly to cover under bits of litter. It sometimes freezes after a brief run as does Emblethis vicarius. There are as yet no known predators or parasites, except for Williams’ (1946) record of it as one of the lygaeids utilized by the sphecid wasp Diplopectron to provision its larval cells. This insect feeds in the laboratory on the seeds of Oenothera sp., Rum ex spp., Chenopodium album, Verbascum thapsus, and a number of unidentified seeds from the ground litter. The insect feeds readily on sunflower seeds. Several notes in the literature indicate probable host plants. Bruner (1891) in Nebraska recorded it from sugar beets, grape vines and on the white pigweed ( Chenopodium album L.). Bruner and Barber (1894) named it the “ clouded weed bug” and said it was partial to Amaranthus, Chenopodium, purslane (Portidaca oleracea L.) and stink-grass ( Eragrostis megastachya (Koel.). Forbes: (1905) noted that Chenopodium album is its favorite food plant and it damages corn. Blatchley (1934) collected it under Amaranthus- in California. Larson and Hinman (1932) stated that it is injuri- ous in pea fields in Oregon. Hussey (1922) collected it on ragweed (. Ambrosia sp.) in Michigan. In the laboratory mating occurs infrequently because the females; repeatedly repulse the males. Copulation eventually does occur,, but it is difficult to observe the mating behavior to completion. In 21 trials involving different males and females at different times, the female did not open her ovipositor and dislodged the male. The male senses the female before actual contact and responds 141 VOLUME XLIV by turning his body and head toward her and holding his antennae stiffly erect at 60° angle from horizontal. When about a centimeter away the male suddenly leaps upon the female, who immediately attempts to dislodge the male. The male quickly orients himself parallel to the female and places the tips of his antennae on the female ’s head. The antennae which are held stiffly and slightly con- vergent apparently are not vibrated, but instead are moved slowly over a small radius on the female’s head. During this time the pygophore is extruded, rotated 180°, and pressed against the tip of the female ’s abdomen. In this position the parameres move like tiny jaws against the ovipositor. When sexually excited by a female the male will leap on other males and even on the female’s cast exuviae. Copulation lasts at least one and one-half hours in the usual end to end position. Life History Sphragisticus in Connecticut is bivoltine and the adults over- winter in a state of reproductive diapause. It was observed to over- winter as an adult (Scholtz 1847, Forbes 1905, Blatchley 1926) and lay eggs in spring (Pfaler 1936). The phenology given in Table 60 is for the seasonal cycle at Canaan, Connecticut on a hot sandy field which warms up earlier than most other sites. There is an overlap of old spring adults and new adults in late June, and in August there is an another overlap of ovipositing first generation adults with dia- pausing second generation adults. TABLE 60 Phenology of Sphragisticus nebulosus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 10 100% May 25 20% 80% June 2 15% 15% 10% 60% June 15 10% 15% 25% 30% 20% July 3 5% 95% July 31 10% 10% 5% 75% Aug. 20 5% 15% 45% 35% Sept. 15 5% 25% 70% Oct. 3 — 10% 90% Oct. 20 100% 142 ENTOMOLOGICA AMERICANA Second generation adults brought in the laboratory in mid- Au- gust remain in an intense state of reproductive diapause, and did not break diapause in midwinter. Adults collected in late October and November for the most part also remained in diapause but a few spontaneously broke diapause and laid eggs in December. Little work could be done on the diapause mechanism, but the available data fits the facultative diapause pattern found in other species in which short photoperiods bring about reproductive cessa- tion. Since Sphragisticus extends to the far north it is probably univoltine under those conditions. The available data is somewhat diverse, but suggests a pattern as in Pachybrachius basalis. The normal prereproductive period is about 5-9 days. One culture of new adults collected July 3 were ovipositing by July 9. In contrast several cultures reared to adults in the laboratory on June 23 did not become reproductive until mid- August. This may account for some of the late nymphs found in the field. On October 3 some very late fifth instar nymphs were col- lected with diapausing adults. The adults remained in diapause, but the 4 new adults reared from the nymphs began laying eggs after 17 days. The data is not sufficient for any further interpretation but indicates an interesting problem to resolve. The stadia given in Table 61 are based on laboratory rearing of field-collected early instars on sunflower seeds. Only four do seven measurements of each of the stadia are available. TABLE 61 Stadia of Sphragisticus nebulosus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 9.5 (9-10) 5.7 (5-7) 5.5 (4-7) 5.7 (4-7) 7.3 (5-9) 10.1 (9-13) 43.0 (36-53) The laboratory longevity of the adults of the first generation is long and varies from 42 to 70 days (mean 59 days). While this is probably longer than the field longevity, it accounts for the overlapping of generations in the field. The scant data available indicates that this species has a high reproductive potential. Two females laid 157 and 224 eggs. The eggs are laid at a rate of 6-7 a day. Six unmated females laid from none to 23 eggs (mean 18). The eggs are smooth and elliptical in shape with only a slight 143 VOLUME XLIV ventral curvature. This shape is generally convergent toward a lygaeoid type egg* (Putshkova 1956). The eggs are oviposited into the parenchyma of stems (Putshkova, ibid) and into litter, crevices and into tight cotton stoppers. Both wet and dry substrates are utilized although dry is preferred. TRIBE GONIANOTINI Emblethis vicarius Horvath The genus Emblethis includes a large number of Palearctic species, but apparently only one Nearctic species, E. vicarius . However, the Palearctic species are rather similar in appearance. Various specimens identified as vicarius from different Nearctic localities were run through Wagner’s (1958) key to the European Emblethis. These Nearctic specimens keyed out near different Palearctic species. It appears probable that E. vicarius is actually a species complex. The distribution of Emblethis is from the highlands of Guate- mala north to British Columbia and Quebec. The specimens from northeastern North America appear distinctly larger and darker. Since Horvath’s type specimens (a lectotype, Slater, in litt.) comes from eastern North America the form discussed here is E . vicarius. Environment Emblethis may be aptly called the “sandbug” for it is very characteristic of, and largely restricted to open sandy habitats (Uhler 1876, Hart 1907, Torre Bueno 1910, et al.). Walkden and Wilbur (1944) found it frequent on overgrazed pastures in Kansas. Lugger (1900) found it in large numbers in fields with Sphragisti- cus. It has been collected along sandy beaches in Long Island (Torre Bueno 1915) and in Florida (Barber 1914b). Hendrick- son (1930) recorded it from prairie associations of Stipa spartea- Andropogon scoparius and Andropogon scoparius-Boutelona curti- pendula found on dry steep hill sides and bluffs. Blatchley (1934) collected it under Amaranthus in southern California. The Guate- mala specimens were collected in the temperate zone at 7500 feet at Ostuncalco (Distant 1882). In New England Emblethis is found naturally along sandy beaches above the high tide line, usually about and under thin patches of litter. In inland areas similar but artificial areas are 144 ENTOMOLOGICA AMERICANA created in gravel pits, parking* lots, sandy fields and roadsides and populations of Emblethis are very frequent in such areas. As a characteristic species of dry sandy areas, Emblethis is the best example of the ability of lygaeids to survive high temperatures. Emblethis is frequently found out on the hot open sand or under the very thin litter. Measurements with a thermistor confirms its resistance to heat. Probes placed next to the insect in the field regularly measure from 43° to 50° C. On the very hot sand over 50° C. the insect does not remain still for long but moves from one small bit of litter to another. These small bits are equally hot but are poor heat conductors. Frequently it stands high on its hind legs to avoid the heated ground. But on sand of 45° C. it could “freeze” for 25-30 seconds. The nymphs are apparently less resistant to heat and under intense illumination keep to shaded cover under litter or around the bases of plants such as Andropogon clumps. In the evenings or on cloudy days the nymphs also run about on the open sand. Despite the dryness of their habitat (1), the insects will die in 24 hours without water. In the sandy habitat of Emblethis the plants which are often present are xerophytic pioneers such as Andropogon scoparius Michx., Eragrostis pectinacea (Michx.), Hypericum perf olium L., Rumex acetosella L., Potentilla spp., Trifolium arvense L., Carex sp. Sometimes, such as on beaches, the plant association is non- existent and the insect live in the dry litter with no green plants present. The abundance varies greatly from one site to another. It reaches 40-50 per square meter at a few places such as a sandpit at Storrs, Connecticut. More usually it is 10-15 per square meter. While beaches form long persisting habitats, the patches of seed bearing litter do not. In inland areas, sandpits and sandy fields are certainly temporary habitats. Although later succession may proceed rather slowly because of the xeric soil conditions, early succession proceeds rapidly. General Biology The completely macropterous condition of this species correlates with the temporary nature of its habitat. In New England it has not been collected at lights as in Missouri (Froeschner 1944). Torre-Bueno (1910, 1915) found Emblethis washed up in beach wash and he deduced from the wind conditions that it had been flying* over water. Emblethis has a remarkably effective procryptic coloration. The adults are mottled brown with dark punctures and rather broad and flattened in shape. There is considerable variation in 145 VOLUME XLIV color in the same population from pale brown to dark brown. The color variation has no relation to the age of non-teneral insects. The nymphs are similarly colored, but show less variation in color. Emblethis infrequently conceals itself under litter and is often found quiescent on the open sand. When disturbed, like other similarly colored animals such as the piping plover ( Choradius melodics) it moves in rapid bursts, suddenly freezes, and seems to literally vanish into the sand as the eye is carried past it. A few predators are known. Knowlton, Maddock and Wood (1946) found 79 specimens of Emblethis in the stomach of the lizard Scoloporus graciosus graciosus (Baird and Girard) in Utah. Williams (1946) found Emhlethis adults and nymphs among the lygaeids used by the astatine spliecid wasp Diploplectron to pro- vision its nests. Emblethis vicarius is parasitized by the catharosine tachinid Petia (= Procatharosia calva Coq.). In mixed associations of Em- blethis, Cnemodus, Pseudocnemodus and Pachybrachius, Catharosia itself was found only in the myodochines, Petia in Emblethis. Ash- lock informs me (in litt.) that he has taken Petia from Emhlethis in California. This suggests that there may be a definite host specific relationship. The parasite overwinters in the host Emble- this and emerges in late autumn and early winter from the hosts which are kept at room temperature. In the laboratory Emhlethis feeds on seeds of Andropogon scoparius, Bromus sp., Chenopodium album L., Solidago sp., Stel- laria media (L.), Bumex acetosella, R. obtusifolia L., Lechea villosa Ell., Oenothera sp., Hieracum sp. and other unidentifiable fallen seeds, but does not feed on Panicum sp. and Paspalum sp. Sun- flower seeds are fed on but the nymphs grow slowly. More rapid growth occurred on a mixture of fallen seeds from the field. The first instar nymphs showed a strong feeding reaction to Oenothera and Bumex obtusifolia. One adult pierced the bracts and the car- pels before reaching and penetrating the seed of Bumex obtusifolia. At the point of contact with each succeeding structure the salivary sheath cone was laid down. There are several host association records from the economic entomology literature: In Nebraska on sugar beets (Bruner 1891), as a general weed feeder and on Chenopodium album L., and roots of Eragrostris major Host (=E. megastachya Koel.) (Bruner and Barber 1894, Forbes 1900). No seed defense behavior was ob- served. The mating behavior is not known in detail since E. vicarius mates infrequently in the laboratory. An active male on contact with a receptive female vibrates his antennae rapidly. After 146 ENTOMOLOGICA AMERICANA climbing on the female he aligns himself parallel and taps his an- tennae rapidly on the female’s head. Copulation occurs in the end to end position. In one instance it lasted 6.5 hours. Life History In southern New England EmMethis vicarius is bivoltine and overwinters as an adult. Because of its open sandy habitat EmMe- this begins ovipositing early and the first instars are collected in early March as given in Table 62. The first and second generations overlap considerably in July. The entire first generation apparently becomes reproductive. Second generation adults, which appear as early as July 30 do not become reproductive but remain in an intense diapause state. Thus ovipositing first generation and diapausing second generation adults are found in the field simultaneously in early August. TABLE 62 Phenology of Emhlethis vicarius Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult Apr. 15 100% May 10 30% 70% May 22 25% 12% 8% 55% May 29 15% 35% 35% 10% 5% June 7 5% 20% 50% 15% 10% June 16 10% 30% 35% 15% 10% July 9 27% 15% 3% — 35% 20% July 30 26% 18% 23% 18% 5% 10% Aug. 13 5% 10% 10% 20% 25% 30% Aug. 25 5% 10% 40% 45% Sept. 7 5% 20% 75% Sept. 20 15% 85% Oct. 15 100% The diapause state is not usually broken at room temperatures. In an exceptional case, a male and a female which lived 224 and 227 days respectively in the laboratory became sexually reproduc- tive March 15 but died a few days later. A facultative diapause under photoperiod control is probable in which long photoperiods promote reproduction, short initiate diapause. Long photoperiods and warm conditions did not affect the diapause condition in 147 VOLUME XLIV December. The stadia given in Table 63 are mostly derived from rearing field collected nymphs on sunflower seeds and seeds from the natural habitat. A few can be reared entirely through their life cycle on mixed field seeds and sunflower seeds, but sunflower seeds alone are successful although first instars from the field are readily reared on sunflower seeds. The growth rate is probably slower than under hot field conditions. TABLE 63 Stadia of Emblethis vicarius Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 9.7 6.2 6.3 6.2 8.5 9.5 46.0 (9-10) (6-7) (5-7) (5-8) (6-10) (7-13) (36-52) Longevity of the non-diapause adults in the laboratory varies from 24 to 48 days (mean 31). Diapausing adults live a long time under laboratory room temperatures (134 to 233 days). However, the adults feed all winter long. The preoviposition period of first generation adults is about 10 days. Overwintered adults brought in from the field in March became reproductive in 15-18 days. The fecundity is high, and in six females it ranged from 150 to 218 eggs (mean, 181 eggs). Unmated and sexually isolated females laid no eggs. The eggs are laid singly at a rate of 6 to 11 per day (mean 8). Lack of a proper substrate greatly inhibits oviposition reducing it to only 15-35 eggs. Provided with sandy substrate, oviposition occurs copiously as shown in the fecundity figures just given. The eggs are cucumber shaped and beset with fine short setules. After considerable probing and testing with both antennae and ovipositor, the eggs are oviposited into the sand or under small pebbles and bits of litter on the sand. The female employs a more horizontal placement of the eggs than in myodochines like Ligyro- coris diffusus and rarely releases the ovipositor to the extent of Ligyrocoris. Partly because of the setules but more from a cement layer, the sand and pieces of litter densely cling to the eggs effec- tively concealing the eggs and making them hard to count as well. The eggs are laid in dry substrates, never in moist substrates. 148 ENTOMOLOGICA AMERICANA Trapezonotus arenarius (L.) Trapezonotus has a Holarctic distribution with nine known species in the Palearctic, and four in the Nearctic region. One species, arenarius , is presumably common to both regions. In both regions the genus has a large north-south distribution, extending south to North Africa and the Middle East in the Palearctic and into Panama in the Nearctic. T. arenarius (L.) has an exception- ally wide-spread distribution and is the only species which reaches eastern North America. It is found in the Palearctic south to the Madeira Islands, Morocco and Turkey, north to Lappland in Europe and east to Siberia, China and Kamchatka (Slater, Cata- logue). However, T. arenarius in the Nearctic region appears to have an entirely different southern limitation. It occurs only from British Columbia east through Manitoba to Quebec and is recorded in the United States only in the highlands of New Eng- land and northern New York. It is not recorded from the western United States and is the only species of Trapezonotus in Canada. The other species of Trapezonotus have a montane distribution in the southwestern United States, and one, T. caligmosus Dist., extends south from Arizona into the high mountains of Guatemala and Panama. T. vandykei Y.D. was collected at 10,000 feet in Colorado, and represents the apparent northern limits of the Nearc- tic group aside from T. arenarius. Blatchley (1926) believed T. arenarius to be an introduced species and Seidenstucker (1951) apparently agreed with the judgment. However, certain aspects of the biology and ecology of this species to be discussed suggest that T. arenarius is not an introduced species, and indeed rather raises the question of the supposed identity of the Nearctic and Palearctic populations. Certainly the European population is very similar to the North American, and the question revolves upon whether the North American “ arenarius ” is a vicariant species (Ross 1962) or a subspecies. In this respect, certainly populations from northeast Asia should be studied. The extremes overlap, but the New Eng- land arenarius appears smaller than the European T. arenarius : New England male, 4.36 (4.22-4.55) to European 4.5 (Seiden- stucker 1951). The European arenarius was described by Seidenstucker (1951) as being constantly macropterous except for very few brachypterous forms from the Alps. Yet the North American population is largely brachypterous. As Lindroth (1957) emphasized, intro- duced species are largely macropterous in their early stages of establishment and generally are restricted to coastal areas. In 149 VOLUME XLIV New England and New York, T. arenarius is not found along the coast but only in the highlands, and is quite scarce in southern New England. Lindberg (1958) did not find T. arenarius in Newfoundland, but did find that the European T. distinguendus (Flor) had established itself along the shore. Even species presently recognized in the arenarius complex (Seidenstucker 1951) have been questioned as being only ecological “forms” and the various differences in the claspers, etc., were considered the result of allometric growth in these forms (South- wood and Leston 1959). It is clear that this European complex requires a careful biological study before the status of American populations can be understood. Our species keys out to T. arena- rius in Seidenstucker 7s key. Along with the different distributions of the Palearctic and Nearctic populations, the habitat preferences are apparently differ- ent. In the extensive European literature T. arenarius is recorded from a large variety of habitats. Southwood and Leston (1959) described arenarius ( sensu lato) in England as inhabiting sand dunes, heaths, light sandy soils, dry woodlands, chalk soils, but as usually absent from damp soils. Seidenstucker (1951) gave a similar habitat range for arenarius ( sensu stricto) in Germany and noted a definite preference for humosere rather than heath and sandy soils, and often light woods. He considered this broad habitat range the explanation for its widespread distribution through cultivated regions. Krogerus (1932) found it to be a xerophilous species common on sandy littoral areas in Finland. The North American population is not found in sandy dune shore areas or in dry woods. Nor is there any relation to agricultural practice, in fact, quite the opposite. These differences seem to make it advisable to treat the two populations as distinct, and only briefly consider the literature on the European T. arenarius. Environment In general, Trapezonotus is a species of open upland habitats. It is found most abundantly in northern New England and northern New York. In southern New England it is found on several north facing drumlin slopes about Storrs, and in northwestern Connecti- cut on the summit balds of Canaan Mountain. Toward the north the species is found in a greater variety of habitats, from dry roadsides near Gorham, New Hampshire to the alpine meadows on Mount Washington at 5,300 feet. The exposure of these habitats is always largely open, often on a slope. The soil is nearly always poor and gravelly, and is over- drained. These dry sites (2—3) at lower elevations support a sparse 150 ENTOMOLOGICA AMERICANA short vegetation which varies from fescue (Festuca capillata Lam.) dominated areas of short grasses, to old bare areas with clumps of Andropogon scoparius with the xeric lichens ( Cladonia ) and the mosses ( Polytricum pilifera) in the wide interstices. On higher elevations Trapezonotus is found in the short (6 inches to 1 and one half foot) dense Vaccinium pennsylvanica scrub, especially along the bed rock outcrop areas which are dominated by scattered xerophytic oak Quercus ilicifolia. On Mt. Washington, New Hamp- shire, adults, and fourth and fifth instar nymphs are found above the three line at 5,300 feet in a Vaccinium patch in an alpine sedge- grass-meadow. This was, of course, much more moist (6-7) than the other habitats mentioned, and with a more uniform ground coverage. Brindley (1935) collected arenarius at 6,000 feet in the Alps. The biotope proper is the thin litter layer especially at the margins of bare or nearly bare areas. It is rare in completely barren sites or in dense ground-covering vegetation, and never where the herbs are over a foot or two in height. Thus, its distribu- tion is quite discontinuous at a given site, and the abundance is usually about 1-3 per square meter or less. At some outcrop margins in Vaccinium cover, it is found at 5-7 per square meter density, and at one favorable barren site at Storrs as high as 17 per square meter. In these habitats it is often found with the other seed bugs : Xestocoris nitens, Carpilis consimilis, Kolenetrus plenus, Ligyrocoris depictus, and Pseudocnemodus canadensis. In overwintering, Trapezonotus moves a short distance into marginal areas of deeper vegetation where I found them hibernat- ing in groups of two or three. The European Trapezonotus is recorded as hibernating in large colonies in moss ( Rieber and Puton 1876, Oliver 1904). General Biology As already mentioned the Nearctic population of T. arenarius is largely brachypterous, and the samples before me are only 11% macropterous. In this population there is also a sexual difference in wing polymorphism in T . arenarius with the condition in the female evidently more plastic. The relationship is summarized in Table 64. This low proportion of macropters correlates with the long lasting nature, often of a serclimax type, of the habitats of this species. Nevertheless several of the habitats especially several roadsides near Gorham, New Hampshire and at Camden Hills, Maine, are nearly ruderal in aspect and appear to be rather short lived sites. So it is apparent that the low proportion of long 151 VOLUME XLIV winged females must be adequate to colonize such an area. One such probable colonization was observed at a site studied at Storrs, Connecticut for five years. In 1958 the species here was absent or at least very rare, for this site was searched closely and carefully for another rather uncommon rhyparochromine in the Storrs area. In 1959 several females were found in May, and dur- ing the summer the population increased rapidly to 17 per square meter in some microhabitats. In the subsequent year, 1960, it had dropped to but 1-2 per square meter and despite very careful search it could not be found in 1961 and 1962. Over this time the plant community changed hardly at all. A very similar decline affected another population about four miles away, but the density here was much lower. TABLE 64 Brachyptery patterns in Trapezonotus arenarius Sex 1 2 3 4 Male 94% 6% Female 8% 41% 22% 29% “1” wing ■ attains tergum 6; “2,’ ’ wing to margin of tergum 7 ; ‘ ‘ 3, ” wing to middle of tergum 7 ; “4,” macropter. Both the shining black late instar nymphs and the brown and black patterned adults apparently have procryptic coloration. The amount of pigmentation varies considerably, but is not related to populations or age changes. In the laboratory these short legged insects strongly prefer samples of litter from the habitat and rarely venture on white or unsheltered backgrounds. In the litter the insects are very cryptic and quickly take refuge under bits of litter and in rolled leaves. Considerable disturbance of the litter is required to alarm them into movement, which, roach fashion, is quickly to the next crevice. When the first instars are put into a dish provided with only methyl cellulose, they are extremely active and crawl continuously around. But when litter is provided they actively seek “ refuge” in the litter and become very quiet. In fact when the litter is comprised of rolled Gaylussica or Vaccinium leaves, it is nearly impossible to determine their numerical presence in the field. No parasites were reared from Trapezonotus. The adults and 152 ENTOMOLOGICA AMERICANA all instars are actively preyed on by the nabid Pagasa fusca (Stein) which is fairly abundant in the same habitats as Trapezonotus. When some litter was brought into the laboratory with adults of Trapezonotus in early April it contained the eggs of the nabid as well. The tiny first instar nabids agilely climbed on top of adult Trapezonotus and pierced them through the back of the head with their stylets. The Trapezonotus quickly succumbed and the di- minutive nabid proceeded to feed. The nabids were reared up to the third instar in this fashion. Gronblom ( 1946 ) found that in Europe, the ‘ ‘ G-rabwespe ’ ’ Astata stigma Panz. provisioned its nest with T. arenarius. This sphecicl is of the same subfamily, the Astatinae, as are the Nearctic wasps which prey on Lygaeidae. In its feeding habits this species is evidently nearly omnivorous on seeds. When exposed to sunflower seeds it reacts very rapidly and feeds within a few seconds. During its feeding I have never observed the distal two labial segments to be bent or removed as in other lygaeids. The seeds are pierced very rapidly compared to other lygaeids. It feeds in the laboratory also on seeds of Fes- tuca sp., Solidago sp., Vaccinium pennsylvanicum, Gaylussica, and Betula spp. It does not react, however, to Panicum or Paspalum seeds. Trapezonotus is reared very readily from field nymphs or from the eggs through to the adult on sunflower seeds, but in some colonies the mortality is very high in the first instar. Cobben (1953) found Trapezonotus feeding on the seeds of Er odium circu- tarium L’Herit. No seed defense behavior was observed. The seeds are often dragged by the beak into litter refuges. T. arenarius, like all other known members of the genus known to me (11 of the 14 species) except the monotypic subgenus Gnopherus is sexually dimorphic : the females have entirely black antennae and legs while the males have the fore legs, basal two- thirds of the mid and hind femora and the basal antennal segment orange-brown (ferrugineous) . However, no functional significance of this dimorphism was observed. Courtship as such is very brief. When placed with a female the male randomly comes in contact with her. The first response of the male is to freeze, extend his antennae, and extrude his genital capsule. When the male touches the female again with his out- stretched antennae, he may suddenly leap upon her. He immedi- ately begins tapping her head with his very rapidly vibrating antennae, and pressing his revolved genital capsule against the females ovipositor area. If the female is receptive in the next moment the male slides off and goes into the reversed copulatory 153 VOLUME XLIV position. If not receptive the female decamps. The whole process may take only four seconds. Mating lasts from two and a half to five and one quarter hours. The male for the first hour or two appears semi-paralyzed and the female literally drags him about. Later the male becomes more active and walks backwards with the female. Copulation reoccurs frequently in the laboratory, and at least six copulations were observed with one female. However, only one mating is required to fertilize a female ’s entire complement of eggs. No male reacts to another male, nor did the sexually active males react to diapause females present in the field at the same time. The single very long lived female mentioned earlier also failed to elicit any response from the males which came out of diapause in October. Life History T . arenarius has a univoltine life cycle with an obligative adult diapause. The phenology observed at Storrs in 1960-1961 is as in Table 65. TABLE 65 Phenology of Trapezonotus arenarius Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 13 100% June 7 9% 9% 18% 9% 55% June 12 10% 20% 30% 10% 10% 20% June 23 5% 28% 43% 21% 3% July 1 5% 5% 10% 25% 35% 20% July 8 7% 9% 22% 53% 9% July 17 2% 8% 30% 60% July 29 2% 10% 88% Aug. 15 100% on In northern New England, late instars are found considerably later, and at Gorham, New Hampshire and Laconia, New Hamp- shire on August 12 and 15 respectively, fifth instars still compose 33% and 25% of the populations found. The new adults are in a state of obligative diapause. They feed very little in late summer and may go without food for several 154 ENTOMOLOGICA AMERICANA weeks or water for several days. They remain in this strong diapause condition until late autumn or winter when some colonies complete diapause, mate, lay a few eggs, and die. It appears from the data that a sudden high tem- perature elevation is the best explanation for the diapause break for photoperiod conditions caused no change, and all colonies of a given year either break or remain in diapause. The change correlates with the temperature records, which show that in 1957 and 1959 the rearing room became briefly overheated, exceeding 105 °F. and diapause terminated shortly after this event. In the other years the temperature remained at least between 65 °F. and 85°F. extremes. Leonard (in lift.) similarly has found that high temperature releases Blissus spp. from diapause in late autumn. The stadia of nymphs reared in the laboratory are as in Table 66. This appears to parallel the field development and is perhaps somewhat slower. The last two stadia are considerably longer, which agrees with the late field occurrence of last instar nymphs. The longevity of the overwintered adults was quite long, and the generation overlap was complete, with some ovipositing females present along with new diapause adults. TABLE 66 Stadia of Trapezonotus arenarius Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 11.6 6.3 6.4 6.8 9.4 11.1 50.0 (10-13) (5-8) (5-9) (6-9) (8-14) (9-15) (46-59) Overwintered adults live in the laboratory usually until between July 1 and July 30, but an unusual female lived 236 days after being brought into the laboratory on May 28. It ceased to oviposite about July 9, but remained alive like normal males until January 1. This represents a total longevity of a year and a half. Most interesting is the nature of the cessation in oviposition in this female. When left in warmth the adult lives either until diapause is broken or dies in midwinter. None survived past January 27. It is clear, then, from this longevity data that the southward spread of this species (or population) may be limited by a requirement for a long cold diapause period. As stated earlier this species has a long oviposition period. Colonies collected on April 19 and 23, however, while copulation oc- 155 VOLUME XLIV curred at this time, did not begin ovipositing until May 4. Colonies collected on May 15 and May 21 were ovipositing and the females contained fully developed eggs. The oviposition period is long, ap- proximately 50-55 days, but probably less in the field due to natural mortality. In perhaps inverse correlation the rate of oviposition in the laboratory is low and averages 2.8 eggs per day (2-4 a day), so that the total fecundity is 58-141 eggs (average 97), which is fairly similar to other rhyparochromines with shorter oviposition periods. For this reason the rapid population increase in 1960 is most inter- esting for it may imply a high survival rate in that particular popu- lation. In those colonies which broke diapause, only 16-25 eggs (average 19) are laid, at a rate of less than 1.2 eggs a day. The nymphs of these eggs appear unusually weak and die off rapidly without de- veloping in direct contrast to the spring colonies. Three virgin females which broke diapause in October laid in all, 17 eggs (aver- age 5.4). When given a choice, oviposition occurs into dry substrates, but T. arenarius will lay its eggs on wet cotton if no other is provided. When ovipositing this species does not prefer deep contact with the substrate, but lays its eggs singly near the margins, and lays eggs under stones, twigs, seeds, into leaf crevices, hollow stems, and into loose dry litter. The short thick eggs are beset with hairs which cling to objects and debris and further conceal the eggs. Delochilocoris umbrosus (Dist.) This species has had a variegated generic history, and is listed under a variety of names. It was originally described by Distant (1893) in the preoccupied genus Dorachosa and Bergroth (1893) substituted Delochilocoris for Dorachosa. Horvath (1908) how- ever, made Delochilocoris a synonym of Aphanus Lap. Most of the records are listed under this name. Uhler, unfortunately, mistook this species for Microtoma carbonaria Rossi (Barber 1918b). Ac- cording to Britton (1938), the 1920 record of Rhyparochromus plenus from Connecticut refers to this species. It was found that Rhyparochromus should replace Aphanus (China 1943), Ashlock (1960) resurrected Delochilocoris and returned umbrosus to it, but it has recently been discerned that umbrosus must be placed in a new genus distinct from the type species, illuminatus Dist. (Slater, Ash- lock, and Sweet in prep.). Thus the literature references are con- fusing. Both of these species belong to the series of gonionotine genera 156 ENTOMOLOGICA AMERICANA endemic to the Nearctic. “D.” umbrosus has a wide distribution and is recorded from southern New England and Ontario, west to Colorado and California, and south through Florida and Mexico to Guatemala and Panama, the latter three being the type localities. In Central America it was collected by Champion in the temperate highlands at 8,000 feet in the Quiche Mts. at Ostuncalco, Guatemala, and between 4,000-6,000 feet on the Volcan de Chiriqui in Panama (Distant 1893). It is interesting that while illuminatus attains only the extreme warm temperature tier of southern states, it is also not recorded south of Guatemala. Blatchley (1926) states that while umbrosus is frequently found in southern Indiana, it was not found in the northern counties. Both Barber (1923) and Froeschner (1944) commented on the gen- eral scarcity of this species. I have found it only in southern New England and also to be one of the more infrequent species. For this reason the following discussion is relatively limited. Environment Its relative scarcity is apparently not due to a restriction to an uncommon habitat for umbrosus is found in a diversity of habitats. Nevertheless these habitats are generally characterized as fully ex- posed open subclimax areas, which are not particularly old sites. The ground level biotope is usually dry and moisture level ranges from 2-4, and litter layer is rather sparse. Both adults and nymphs are found in the following habitats : temporary new fields dominated by pioneer weedy plants; on a sandy flood plain in Oenothera and Verbascum thapsus L. litter; in dry pastures on mullein leaves (V erbascum thapsus) and at the edges of rock out- crops; and in mullein leaves and grass (Agropyron repens (L.) at a sandy ruderal site. One ovipositing female was collected at 1,800 feet on exposed rock ledges on Canaan Mountain. In all these sites the species occurs in low abundances of 1-2 per square meter, at most. At one site in Mansfield Center, Connecticut on July 12, 1957, it was found in the greatest abundance (6-7 per square meter) and in all nymphal stages. This was a flood wash area at the base of a sandy scree from a sand pit. The gravelly ground supported a low ruderal vegetation, and was thinly covered with light dry flood debris. Also present on the ground in this habitat were the lygaeids Emblethis vicarius, Ligyrocoris diffusus, Peritrechus fraternus, and a few Megalonotus sabulicolus. At each of the above sites, umbrosus was associated with different lygaeid species, which reflects the ecological diversity of the habitat selection of Delochilocoris. The various habitat notes in the literature agree with the ob- served findings and appear to be in open areas. Blatchley (1926) 157 VOLUME XLIV collected it in spring* and summer in Indiana by sweeping low herb- age, more commonly in sandy localities. He found it hibernating beneath logs, chunks and leaves of mullein (1895). Vestal (1913) collected it in Illinois under a board at the edge of a field. Forbes (1905) found it in corn fields apparently on husks and hibernating under bark. Froeschner (1944) found it “hibernating in grass clumps, or under rocks, logs or mullein leaves.” Walkden and Wilbur (1944) collected the species in native hay habitats of brome grass and blue grass in Kansas. Gillette and Baker (1895) col- lected it under stones in Colorado and Rainwater (1941) recovered it from Spanish moss in Louisiana and South Carolina. Nearly all of the D. umbrosus habitats are apparently of recent origin and temporary nature. At not one site, even the most favor- able one, was the species found in a subsequent year. In two weedy field sites which were carefully collected earlier, the occurrence of the species apparently represented a new migration into the site. The “native hay” habitats in Kansas (Walkden and Wilbur 1944). however, would be a climax prairie association rather than a new subclimax. General Biology Such utilization of newer subclimax communities makes under- standable the completely macropterous condition of this species. Several references may refer to its ability to disperse. Glick (1939) collected it 200 feet in the air in Louisiana. Caudell (1902) found it on mountain snow in Colorado. It has not been collected at lights. When disturbed in the field, this black, rapid-moving insect runs but a short distance and takes cover in the nearest crevice. This protective behavior is reflected in the flattened shape of the insect. Perhaps the black coloration may enhance the shadow effect in such crevices. At any rate, it is frequently found in light background habitats and both the adults and the equally black nymphs are very conspicuous when disturbed, and the contrast would be apparent to both color blind and color perceiving predators. No parasites or predators are yet known. However Billings and Glen (1911) found it infected with a Sporotrichum fungus. In the laboratory it fed readily on sunflower seeds and could be completely reared from egg to adult. It also feeds on seeds of the mullein, V erbascum thapsus, grasses, Agropyron repens , Pani- cum sp., and a number of unidentified composite seeds collected in wash litter. No seed defense behavior was elicited, perhaps because in the laboratory only small populations were available. Seeds are, how- ever, moved to more protective sites. 158 ENTOMOLOGICA AMERICANA Mating behavior was observed only in the male courtship be- havior. The male or female consistently decamps before copulation is completed. In contrast to other species the female apparently does not show an avoidance reaction, and it is the male which usu- ally decamps. Since the available material is rather old by middle June, it may have been too late for normal copulation. Four similar trials were made, and the following sequence was observed re- peatedly. At any point the male may decamp. The sequence is always repeated. If the sexes come close to each other during random movementsy the male may perceive the female without actually touching her and extend his antennae toward her. Or, the male may directly touch the female by accident. The female may become passive or more often turn and face the male, and touch antennae. The male fre- quently decamps or becomes passive at this point. If not, the male becomes sexually stimulated and vibrates his antennae rapidly with the distal three segments held horizontal and at right angles to the scape. The male moves upon the female with slight shaking move- ments reminescent of Ligyrocoris and protrudes and revolves the pygophore. The response of the female is to become very passive, but as the male climbs upon the female from the side, she may place her hind leg across the male’s head. Once on the female the male’s antennae are held parallel and tapped on the female’s head. The pygophore, with the claspers working, is moved back and forth in the area of the ovipositor. In no case did the female release the ovipositor and allow copulation. Frequently the females were ob- served to touch antennae and quiver them briefly, and become quiet. Life History D. umbrosus overwinters as an adult (Blatchley 1895, 1926, Froeschner 1944). It has not been definitely ascertained whether there are two generations or a very long single generation. The data pattern resembles Peritrechus fraternus, and it is very prob- able that two generations are involved. The observed phenology at Storrs is as in Table 67. In the laboratory, the overwintered adults live until late June and early July, and July 2 is the latest date of oviposition. It seems that very few adults can survive that long in the field. Along with this, the late occurrence of early instars in mid-July, and fifth in- stars on September 8 strongly suggests that two generations are involved, especially as oviposition occurs in May and fifth instars are found as early as June 17. If so, the pattern is very similar to Peritrechus for in the labora- tory cultures, the new adults, which are not obtained until late 159 VOLUME XLIV July, are in reproductive diapause. Also, adults reared from nymphs collected on July 14 in the field were in diapause. If so the diapause may be similarly governed by a photoperiod stimulus. The diapause is a strong one, and is not broken by cold ex- posures. The diapausing adults survive as late as mid-winter and die without coming out of diapause. The average longevity of such diapausing adults was 148 days, the range (73-184 days). The stadia in days are as given in Table 68. A few first instars lived 27 days and then died, despite feeding. TABLE 67 Phenology of Delochilocoris umbrosus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult May 31 100% June 17 33% 33% 34% July 10 50% 50% July 14 6% 18% 12% 18% 18% 28% Aug. 4 10% 15% 25% 50% Sept. 8 37% 63% Oct. 2 100% TABLE 68 Stadia of Delochilocoris umbrosus Egg Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Total 9i> T2 8A (L3 84) 104 54.3 (9-10) (8-10) (7-10) (6-7) (6-10) (7-14) (46-62) Oviposition of fertile eggs occurs from May 9 to June 2. The average fecundity is 123 eggs (93-152 eggs). The eggs are laid at a rate of 3.5 eggs a day, but one female over a week laid eggs at 5.9 a day. Oviposition ceases when seeds are removed. Sexually iso- lated females do not lay any eggs. Oviposition is always on a dry site, and into some sort of loose substrate. Loose dry soil is preferred over methyl cellulose, and there is a strong tendency to lay the eggs under objects like twigs, pebbles and seeds. Even into methyl cellulose and litter the eggs 160 ENTOMOLOGICA AMERICANA are oviposited in a relatively horizontal plane rather than vertically as in Ligyrocoris. The short eggs are beset with small spines and cling to litter, obscuring the eggs. In this respect it is interesting that after oviposition, the female usually moves the ovipositor back and forth which further covers the eggs with minute bits of litter. Malezonotus fuscosus Barber The Nearctic genus Malezonotus is predominantly western in distribution, with five of the seven species restricted to the area west of the Great Basin — none are as yet known from the Rocky Moun- tain ranges. Although Ashlock (1958) mentioned only one species, M . august at us , as being found outside a mountainous area, the re- maining two are low-land species. M. rufipes has a wide distribu- tion from southeastern Arizona through the southern United States to Virginia. M. fuscosus, itself, has the most restricted distribution and was known only from New Jersey, and Long Island (Ashlock 1958, in lift.). Malezonotus fuscosus is here recorded for the first time from Connecticut. This apparently represents the first collection of the species since Barber (1918c) collected the type series in 1916 in Lakehurst, New Jersey. A few were collected earlier: Barber (1914a) states that the records of Malezonotus fuscosus (as rufipes ) listed in The Insects of New Jersey (Smith 1910) are based on two specimens which he collected at Lakehurst. The sole other speci- men was collected in 1915 at Fire Island Beach on Long Island, New York by Torre-Bueno (Barber 1918c). M. f uscosus was collected in east central Connecticut at a single station near Mansfield Center, in an area which is largely dominated by an outlier pine barren community. This New Jersey-Long Island-southeastern New England distribution is similar to many plant species which are endemic to the coastal pine barren formation (Nichols 1913a, Transeau 1913, Fernald 1925, 1933, Conard 1935 and Braun 1955). Environment This single station is a south-exposed slope of a small hill, the top of which is covered with a stand of Pinus rigida. The over- drained dry soil of this slope for the most part supports an Andropogon community except along an especially dry margin be- tween the andropogonetum and a fescue field. This marginal ridge area supports a very sparse plant cover of small thin grasses Festuca capillata L. and Eragrostis pilosa (L.) ( ?), lichens (Cladonia) , and a number of small low forbs, Sericocarpus aster oides (L.), Hier- 161 VOLUME XLIV acium scabrum Michx., Solidago odora Ait., Potentilla canadensis L., Antennaria capillatum, and some Bubrus villosus L. This small rim-like area, but 10 x 40 meters in extent, is the habitat of M. fuscosus. Very few specimens could be found outside this area in the marginal Andropogon association. While the habitat appears to be a moderately old one from the presence of a heavy growth of lichens, it may be in part maintained from Andropogon ecesis because the late summer mowing of the fes- cue field also covers this barren spot. This mowing, however, hardly affects the sparse low vegetation of the site. The poor soil and litter is very dry (2) and the sun-exposed ground temperatures reach at least 135° F. The soil, while sandy, is rather dark which may represent charcoal from an old burn. The abundance of Malezonotus is low and varies from 0.1 per square meter in spring to 2-4 per square meter in mid-summer. The population is highly discontinuous and consists of small groups scattered here and there in the area. It is the only rhyparochromine in this dry area despite a rich myodochine fauna in neighboring biotopes. Both of Barber’s collections (1914a, 1918c) of Malezonotus at Lakehurst, New Jersey were under huckleberry ( Gaylussacia buccata L.) but none could be found under nearby Gaylussacia cover at the Connecticut sta- tion. The collection under huckleberry may indicate at least an open site on the pine barrens. Other species of Malezonotus apparently occur in open habitats also: M. rufipes (Stal) (as sodalicus (Uhl.) ) was collected in num- bers, hibernating under Andropogon (Froeschner 1944) ; and M. sodalicus has been noted as common in strawberry fields in British Columbia (Parshley 1919) ; and M. angustatus was collected under dried cattle droppings in a pasture (Downes 1924). General Biology M. fuscosus appears to be most closely related to M. rufipes (Ashlock 1958) so it is interesting that nearly all fuscosus speci- mens are brachypterous and rufipes is entirely macropterous. All reared specimens are brachypterous. The only known macropter of fuscosus is the specimen collected in beach wash up on Long Island by Torre-Bueno. Presumably this may indicate dispersal over or near the ocean. As the site described earlier is essentially a tempor- ary one, although apparently a long persisting stage, this low pro- duction of macropters must be sufficient to maintain this rare spe- cies. However, its rareness may stem in part from a low vagility coupled with a particular habitat selection. When disturbed, M. fuscosus is an extremely rapid moving in- sect especially in relation to its small body and medium length legs. 162 ENTOMOLOGICA AMERICANA Like Trapezonotus it hides closely under small chips, etc., and will not move until directly disturbed. Its dark fuscous color blends in well with the soil of its biotope. The late instars are shining black. This insect is very nervous and active in its normal movements, and constantly vibrates its antennae. Ashlock noted (1958) that M. angustatus was difficult to observe because of its great activity. Malezonotus fuscosus feeds readily on a variety of small seeds : Antennaria capillata, Car ex sp., S ' ericocar pus asteroides, Hieracium sp., Eragrostris sp., Festuca capillata L., Solidago odor a Ait., Po- tentilla canadensis L., Rumex acetosella L., Panicum sp., Aquilegia canadensis (L.), and sunflower seeds. The insects oviposit readily on the seed diet and could be reared very readily from field-collected second instars. However very slow development occurred in nymphs hatched in the laboratory and only a few attained the third instar. I could not promote growth with different seeds, nor with green material from the habitat. Ashlock (1958, in lift.) records a similar difficulty in attempting to rear Malezonotus angustatus V.D. The seeds are often dragged by the beak to more protected sites for feeding. No seed defense behavior was observed which may be related to the low laboratory culture abundances. As in Trapezonotus, the sexes are dimorphic in coloration, the males having distinctly orange fore femora, the female dark legs. The significance is not evident in the observed mating behavior. The male does not respond until after he comes in contact with the female. The sexual response of the male is to vibrate his antennae very rapidly, holding with the terminal three segments horizontal and at right angles to the body and the scape. Next the male ad- vances on the female with slow somewhat jerky movements. If the female remains passive the male climbs upon her and proceeds to vibrate his antennae close to her head and places the revolved genital capsule over the apex of the ovipositor. This is similar to the ac- tions of Delochilocoris but the movements are much more nervous and hesitant. Mating is frequently repeated. Ashlock (1958) reports mating frequent in M. angustatus and that they mate in an obtuse angle. M. fuscosus may or may not rest in an obtuse angle. When the pair is moving the male is clearly directly behind the female. Life History M. fuscosus has a bivoltine life cycle with a facultative but strong diapause, and overwinters as an adult. The available phenology is as in Table 69. Barber’s collections were both in early spring, March and April, and the insects presum- ably were in hibernation. The adults collected in June actively oviposit as do the first 163 VOLUME XLIV generation adults in August. However, the second generation uniformly enters into a diapause which I could not break with a cold exposure. The diapause is not related to feeding as the fall adults feed readily and become quite fat, so that the abdominal con- junctiva are exposed. Dissection, however, showed immature ovaries. A most interesting aspect of the diapause is that there is very little mortality in warmth and the adults, although in re- productive diapause, are active all winter in the laboratory, and a number live until early summer. Diapause was broken in an aliquot in the spring by placing the insects in a long photoperiod (15 hours) room. While a relation between diapause and photoperiodicity is suggested by the evidence, further experimental work is needed. Diapause was not broken until after 47 days of long day illumina- tion. TABLE 69 Phenology of Malezonotus fuscosus Date Instar 1 Instar 2 Instar 3 Instar 4 Instar 5 Adult June 10 July 9 Aug. 11 5% Sept. 20 100% 20% 60% 20% 10% 20% 40% 25% __ 100% Only the egg development period of 15 days is available. The very long stadia of laboratory-hatched nymphs are abnormal and not included. Some remained as first instars for as long as 28 days. The field evidence indicates a total development period of approx- imately 40 days. The longevity of two adults which overwintered in warmth was remarkably long; female 323 days; male, 453 days. The over- wintered adults, however, from the field die in late June and early July. The fecundities are 58, 68, and 102 eggs which are laid at a rate of 3-4 per day. Deprivation of seed food causes oviposition to cease. One virgin female laid no eggs. The eggs are laid in the laboratory in seed heads of Antennaria and Solidago, in grass culms, in heavy plant pilosity and in other litter crevices. It lays eggs sparingly in methyl cellulose, and pre- ferred plant debris crevices and fuzz. 164 ENTOMOLOGICA AMERICANA Literature Cited Adam, C. C. 1915. An ecological study of prairie and forest in- vertebrates. Bull. nat. Hist. Surv. 11 : 31-280. Aitkins, A. E. 1936. The “ Spruce Cone Bug” Gastrodes abietis Linn. (Hemiptera : Heteroptera : Lygaeidae) . Ent. mon. Mag. 72 : 139-49. Allee, W. CM A. E. Emerson, O. Park, T. Park, and K. P. Schmidt. 1949. Principles of Animal Ecology. W. 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(original not seen) 193 INDEX To Volumes 43, 44 Pages to volume 44 and plant names in italics abdominalis, Ligyrocoris 8 abies, Picea 123 abietis, Eremocoris 119 abietum, Gastrodes 123 Acer 89 acetosella, Rumex 105, 107, 33, 38, 125, 145, 146, 163 Achillea 94, 84 Acompus 74, 56 acris, Ranunculus 84 acuminatus, Aster 118 aeneoventris, Alophorella 58, 22 affinis, Scolopostethus 32, 57, 75, 103, 105 Agropyron 62, 33, 36 alba, Betula 119 alba, Quercus 33 albimaculus, Cnemodus 30, 33 albocinctus, Pacliybracliius 12, 19, 25, 34, 36, 46, 47, 50, 54, 58, 59, 62, 66, 71, 74, 76, 84, 87, 121, 122, 15, 21, 24, 80, 111 album, Chenopodium 3, 4, 8, 17, 140, 142, 146 Alfalfa 93 Alisma 121 Allothrombium 57 alnifolia, Cletlia 121 Alydidae 61 Amaranthus 141, 144 Ambrosia 10, 141 americanus, Plinthisus 12, 15, 19, 24, 35, 37, 39, 44, 46, 48, 50, 54, 66 Anaphalis 89 anatrepsis 93 Andropogon 25, 35, 36, 49, 62, 89, 100, 105, 3, 12, 31, 33, 42, 43, 145, 161, 162 andropogonetuih 12, 161 angustatus, Malezonotus 161, 163 angusticollis, Eremocoris 116 angusticollis, Peritrechus 124 angustifolium, Vaccinium 113, 26, 47 annua, Poa 36 annulicollis, Sisamnes 111 anmilicornis, Myodocha 91, 93 Anolis 56 Anthocoridae 56, 60 Antillocorini 11, 13, 15, 16, 74, 62 Antillocoris 15, 32, 35, 38, 42, 52, 56, 62, 68, 69, 76, 62, 64, 66, 72, 79, 132 Aphanus 156 arenarius, Trapezonotus 13, 20, 25, 37, 42, 45, 47, 51, 71, 74, 78, 85, 87, 113, 26, 55, 112, 117, 149, 156 art emisii folia, Ambrosia 92, 104, 8 Artliemis 92 arvense, Tri folium 145 Aspergillus 9 Astatinae 57 Aster 62, 94, 119, 76, 79, 96, 97, 102, 108 asteroides, Sericocarpus 161, 163 atlanticus, Scolopostethus 13, 19, 24, 42, 45, 47, 50, 62, 68, 74, 84, 86, 121, 76, 96, 103, 107, 111, 118, 119 atra, Cinochira 57, 113 atrata, Microtoma 12 atropicta, Ozophora 79 baccata, Gaylussacia 47 bacteria 27, 63 balsamea, Abies 60, 124 banksiana, Pinus 124 barberi, Carpilis 46 basalis, Pachybrachius 12, 19, 24, 25, 46, 47, 50, 58, 60, 63, 65, 66, 70, 71, 74, 76, 84, 85, 94, 15, 20, 23, 25, 68, 90, 125, 140, 143 bassiana, Beauveris 26 Betula 62, 48, 63, 89, 97, 114, 118, 119, 153 bicolor, Solidago 96 bifiora, Impatiens 99 bilobatus, Pachybrachius 11, 12, 19, 54, 56, 58, 65, 70, 15, 16, 18, 21, 24, 26 birch 102, 104, 112, 117, 121 Blissus 155 borealis Eremocoris 117 bouringiana, Cattleya 71 Brassica 36 Braconidae 59 breviusculus, Blissus 53 Bromus, 17, 146 brunneus, Drymus 74, 75, 95, 99 ENTOMOLOGICA AMERICANA buccata , Gaylussacia 162 bullatus, Geocoris 42 bursa-pastoris, Capsella 140 caesia, Solidago 118, 119 caliginosus, Trapezonotus 149 Calluna 84 calva, Petia 146 canadensis, Aquilegia 94, 17, 27, 163 canadensis, Erigeron 8 canadensis, Potentilla 94, 104. 107, 42, 48, 162, 163 canadensis, Pseudocnemodus 12, 20, 25, 42, 43, 46, 50, 54, 56, 58, 63, 66, 71, 74, 76, 87, 26, 28, 29, 30, 42, 151 canadensis, Sambucus 99 canadensis, Solidago 84 canadensis, Tsuga 119, 60, 62, 64, 89, 104, 117, 124 capillata, Festuca 114, 47, 71, 72, 151, 161, 163 capillatum, Antennaria 162, 163 carbonaria, Microtoma 156 Carex 26, 62, 113, 121, 122, 103, 112, 114, 116, 145, 163 caricis, Ligyrocoris 12, 19, 25, 35, 37, 45, 46, 49, 50, 54, 62, 71, 74, 76, 84, 87, 103, 109, 114, 120, 123, 21, 111 carolinianum, Rosa 121 carota, Daucus 104, 107, 3, 84 Carpilis 15, 41, 49, 54, 35, 46, 51, 53, 55, 68 catawbiense, Rhododendron, 56, 66, 141 Catharosia 57, 59, 90, 106, 115, 4, 9, 13, 17, 22, 27, 33, 38, 69, 119, 127, 132, 146 Catharosini 59 Centaurea 62, 85, 115, 3, 135, 138 centipede 56 Chalcidae 57 Chauliops 41 Chenopodium 141 Chilacis 90 chiragrus Megalonotus 12, 18, 24, 29, 46, 47, 51, 57, 66, 71, 74, 78, 133, 137 cimicoides, Ilyocoris 32 cincticornis, Heraeus 97 circutarium, Er odium 153 Cladonia 105, 114, 26, 47, 55, 151, 161 clavigera, Sisamnes 12, 20, 25, 29, 38, 40, 45, 47, 50, 71, 74 Cleradini 71, 38, 41, 43 Cnemodus 15, 39, 53, 59, 62, 69, 89, 5, 12, 31, 34, 146 coatimundi 53 collinus, Xestocoris 71 compactus, Plinthisus 59 composites 56, 62, 88, 100, 104, 107 compressa, Danthonia 47 conicola, Gastrodes 123 consimilis, Carpilis 12, 19, 20, 25, 29, 37, 45, 46, 50, 71, 74, 76, 87, 113, 26, 42, 46, 49, 51, 52, 55, 151 constrictus, Perigenes 12, 20, 24, 44, 46, 50, 58, 71, 74, 77, 8, 9, 11, 125 contractus, Sisamnes 65, 41, 43 contractus, Taphropeltus 32 convivus, Peritrechus 124 cordifolius, Aster 79 Coreidae 61, 69 Coreoidea 63 Corixidae 32, 38 corymbosum, Vaccinium 48, 76, 79, 108, 118 costalis, Zeridoneus 12, 19, 24, 26, 46, 56, 58, 66, 71, 74, 77, 2, 7, 8 crassus, Drymus 13, 19, 24, 35, 37, 38, 45, 46, 50, 74, 77, 84, 86, 95, 96, 101, 103 crispus, Rumex 94, 140 cristabellus, Anolis 26 Cryphula, 15, 47, 49, 52, 56, 62, 68, 76, 88, 67, 70, 72, 81 curtipendula, Boutelona 3, 144 curvipes, Pacliybrachius 16 cyanus, Centaurea 29 Cydnus 90 Danthonia 62, 72 decoratus, Scolopostethus 63, 75, 103, 113 Delochilocoris 15, 68, 70, 78, 156, 157, 163 dentatum, Viburnum 76, 108, 118 depicturata, Ozophora 78 depictus, Ligyrocoris 12, 19, 25, 37, 45, 46, 49, 50, 54, 58, 62, 71, 74, 77, 84, 87, 88, 102, 113, 115, 118, 119, 121, 124, 5, 14, 151 dichotoma, Aristida 113, 26 Dieuches 56 195 VOLUME XLIV diffidens, Scolopostetlius 13, 24, 34, 42, 45, 47, 50, 57, 62, 65, 71, 74, 84, 86, 60, 61, 64, 96, 103, 110, 112, 117, 119 diffuSus, Ligyrocoris 12, 20, 26, 44, 46, 49, 50, 54, 58, 59, 62, 65, 71, 74, 77, 84, 102, 110, 112, 114, 118, 122, 124, 2, 3, 5, 7, 8, 10, 42, 93, 100, 112, 125, 127, 140, 148, 157 Dimorphopterus 66 diphylla, Mitella 96 Diploplectron 57, 137 , 141, 146 Diptera 30 discretus, Antillocoris 62, 66 discors, Cymus 87 dispositus, Perigenes 8 distinguendus, Peritrechus 124 distinguendus, Trapezonotus 18, 150 Dorachosa 156 dorsalis, Oedancala 87 Drymini 11, 13, 15, 16, 26, 31, 36, 47, 74, 75, 95 Drymus 14, 16, 27, 44, 47, 48, 54, 56, 62, 65, 68, 77, 78, 84, 86, 57, 58, 85, 95, 97, 99, 100, 102 Dryndella 57 Dysdercus 33, 61, 66, 67 dysenterica, Puticaria 84 elatior, Festuca 48 Elymus 119 Emblethis 14, 15, 31, 53, 54, 58, 62, 67, 71, 78, 144, 147 Entomophtliera 90, 113 equestris, Lygaeus 53 Eragrostris 163 Eremocoris 14, 36, 52, 54, 55, 57, 59, 62, 65, 68, 69, 76, 86, 61, 96 Erica 84, 90, 116, 118, 119, 126 Erigeron 94, 105, Erythroneura 80 Exptocliiomera 15, 35, 54 fasciatus, Oncopeltus 60 ferus, Eremocoris 13, 19, 24, 25, 31, 34, 42, 46, 47, 50, 52, 53, 56, 63, 67, 71, 74, 75, 86, 60, 62, 76, ~96, 108, 116, 122 ferus, Nabis 94, 106 ferruginea, Carpilis 20, 46 fescue 73, 74 Festuca 62, 88, 100, 105, 114, 119, 3, 4, 17, 50, 68, 72, 74, 125, 153 flava, Sarracenia 93 Formica 55, 32 formiciforme, Xenydrium 62 fracticollis, Pachybracliius 12, 26, 15 Frag aria 92, 94 fraternus, Peritrechus 13, 20, 25, 43, 45, 47, 50, 54, 56, 58, 71, 74, 76, 84, 85, 16, 42, 124, 132, 157, 159 fuligineus, Stygnocoris 83, 90, 92 fungi 27 furcatus, Andropogon 104 furze 84 Fusarium 64 fusca, Pagasa 56, 106, 115 ,153 fuscipes, Neotoma 78 fuscosus, Malezonotus 13, 19, 25, 45, 47, 51, 68, 71, 74, 76, 15 161, 164 gale, Myrica 118, 119 Galium 92, 99, 100 Gastrodes 14, 18, 73, 123 Gaylussacia 27, 119, 152, 153, 162 geniculatus, Peritrechus 124 Gelastocoris 66, 67 Geocorini 61 Geocoris 52, 62, 38 Gerridae 32 Gerris 64 giraffa, Myodocha 91 glabra, Rhus 76 glauca, Picea 123 Gnaphalium 114, 115 Gnopherus 153 Gonianotini 11, 13, 15, 16, 26, 36, 48, 52, 54, 70, 74, 139, 141, 144 Gonianotus 68 gracilis, Sphaerobius 12 graciosus graciosus, Sceloporus 146 grandentata, Fagus 89 grandis, Scolopostethus 103 greeni, J uncus 47 grossipes, Gastrodes 57, 123 guttatus, Heraeus 99 Hedeoma 27 Heinsius 41 Hemiptera 60 hemlock 61, 63, 96, 117, 121 Ilenestaris 52 ENTOMOLOGICA AMERICANA Heraeus 15, 54, 85, 97, 98, 100, 16, 90 Heterogaster 75 Heteroptera 30, 31, 53, 57, 63, 75, 76, 140 Hieracum 146, 163 hirtipes, Cnemodus 32 Homoptera 60, 61 huckleberry 162 Hypericum 62, 91, 92, 94, 121, 4, 17, 42, 43 ■ilici folia, Quercus 47 , 151 illuminatus, Delochilocoris 156, 157 inflatus, Cnemodus 30, 31 insignis, Sphaerobius 12, 20, 25, 27, 46, 50, 54, 55, 58, 66, 71, 74, 77, 12, 14 intybus, Cichorium 3, 4 Ischnodemus 66 jacobeae, Nithecus 8, 31 japonicus, Lathy rus 125 jays 51 Jussiacea 22 Icalmii, Bromus 47 klugii, Anolis 26 knulli, Zeridoneus 2 Kolenetrus 20, 54, 68, 55, 57 Laconia 112, 113, 115 laevis, Bidens 99 laricina, Larix 124 Lasius 52 lateriflorus, Aster 118, 79 latifolia, Spirea 48, 55, 91, 97, 131 latimarginatus, Ligyrocoris 65, 2 latus, Drymus 95 Ledum 90 lenta, Betula 79, 118 Lepachys 107 Lepidium 17 Lepidoptera 30 Leptoglossus 60 Lethaeini 11, 13, 15, 16, 74, 67 lethierryi, Scolopostethus 75 leucanthemum, Chrysanthemum 92, 104, 106, 3, 4, 10 Ligyrocoris 10, 15, 48, 54, 66, 69, 84, 85, 87, 101, 109, 115, 117, 119, 122, 123, £, 4, 6, 11, 14, 21, 148, 159, 161 limbatus, Dolichonabis 67 limbatus, Geocoris 56, 42 Lithobius 56, 85, 113 litigiosus, Ligyrocoris 2 lizards 56 lundi, Peritrechus 32 lupulina, Carex 4 luridus, Cymus 87 luridus, Pachybraehius 12, 26, 15 lustrans, Catharosia 58 lutea, Betula 117 Lygaeinae 73 Lygaeus 52 Lygus 22 maculatum, Geranium 84 maculatus, Callosobruchus 32 major, Eragrostis 146 major, Nabis 85, 113 Malcinae 69 Malezonotus 15, 49, 68, 70, 78, 88, 89, 15, 161, 162 margaritacea, Anaphalis 89, 91 maritima, Prunus 125 mavortius, Cnemodus 12, 19, 25, 27, 42, 45, 47, 50, 54, 56, 58, 71, 74, 76, 30, 31, 34, 35, 42, 67 media, Stellaria 140, 146 Medicago 107 Megalonotini 10, 11, 13, 16, 26, 36, 48, 53, 54, 85 Megalonotus 16, 18, 62, 68, 84, 89, 134, 137, 139, 141 megastachya, Eragrostis 141, 146 melanocarpa, Aronia 47 melodus, Choradius 146 Mentha 113, 114, 133, 139 Microvelia. 36 millefolium, Achillea 92, 84, 86 minor, Nabis 85 minutus, Antillocoris 13, 19, 24, 31, 34, 35, 37, 42, 45, 47, 50, 52, 71, 74, 86, 62, 66, 76, 108 Miridae 2, 4, 5, 30, 31, 58, 60, 73, 89 miriformis, Myrmus 72 mirmicoides, Nabis 85, 113 miruginodis, Myrmica 91 Mizaldus 62, 64, 66, 67 Monarda 100, 107 muhlenbergii, Panicum 4 muhlenbergii, Paspalum 94, 72 multifarius, Cnemodus 30 197 VOLUME XLIV mycetomes 63 Myodocha 15, 32, 54, 62, 91, 94, 97, 98, 100, 101, 16, 19, 64, 79, 90, 132 Myodochini 10, 11, 12, 14, 16, 26, 27, 29, 31, 36, 44, 48, 49, 53, 55, 58, 67, 70, 74, 76, 87, 88, 91, 100, 54 Myrica 34, 76, 119 Nabidae 56, 60 nana, Betula 119 nana, Exptochiomera 11, 12, 19, 53 nebulosus, Sphragisticus 12, 13, 20, 24, 25, 29, 46, 47, 51, 57, 66, 74, 76, 139, 142, 143 Neotoma 59 nettles 112, 115 nevadensis, Amphispiza 56 Nitaparavala 41 nitens, Xestocoris 13, 19, 25, 29, 45, 47, 50, 52, 71, 74, 87, 89, 113, 26, 47, 55, 68, 71, 73, 74, 81 nodosa, Ptochiomera 12, 19, 25, 26, 45, 47, 50, 56, 58, 74, 35, 36, 39, 40, 43 Notonectidae 32, 38 novaeangliae, Aster 91 Nysius 52, 63 obscurus, Ligyroeoris 2 obtusa, Festuca 118, 119 oh tusi folium, Rumex 3, 27, 36, 38, 146 occidentalis, Cephalanthus 121 occidentalis, Thuja 118 occultus, Pachybrachius 15 odontogaster, Gerris 38, 39 odora, Solidago 162, 163 Oenothera 105, 17, 33, 114, 137, 141, 146, 157 officinalis, Saponaria 84 officinalis, Veronica 48 oleracea, Portulaca 141 Oncidium 71 Oncopeltus 60, 61, 66, 67 Orchestia 125 Orthoptera 33 Ozophora 15, 36, 54, 62, 86, 87, 75, 76, 78, 79, 118 Ozophorini 11, 13, 15, 16, 26, 31, 44, 53, 67, 75 Pachybrachius 15, 54, 59, 65, 84, 85, 87, 109, 5, 15, 17, 18, 14 pacificus, Heraeus 97, 43 pallescens, Ozophora 79 paludemaris, Peritrechus 13, 19, 25, 34, 45, 47, 50, 54, 58, 59, 62, 125, 127, 128, 130, 133 palustris, Hibiscus 131 Pamera 15 Panicum 62, 105, 115, 3, 13, 16, 17, 20, 33, 47, 48, 50, 67, 69, 79, 127, 129, 146, 153, 158, 163 papyrifera, Betula 48, 60, 62, 64, 102, 104, 117 Paspalum 62, 94, 4, 13, 17, 146, 153 patens, Spartina 131, 132 patula, Atriplex 114 patula, Solidago 99 pectinacea, Eragrostis 145 pedestris, Stygnocoris 12, 13, 18, 20, 24, 26, 44, '46, 50, 54, 63, 74, 84, 56, 83, 85, 88, 94 Penicillium 9 Pennsylvania, Casnonia 91 pennsylvanica, Phymata 106 pennsylvanicus, Camponotus 32 pennsylvanium, V accinium 55, 151, 153 pennsylvanica, Myrica 66, 76, 118 Pentatomidae 52, 61, 63, 69, 71 Pentatomoidea 60 Pentatomomorpha 57, 61, 71 Perellus 61 perfoliata, Specularia 140 perfoliatum, Eupatorium 99 perfolium, Hypericum 145 Perigenes 15, 48, 54, 68, 69, 85, 4, 5, 8, 10 peregrina, Comptonia 55 Peritrechus 14, 15, 28, 31, 62, 68, 75, 84, 42, 124, 126, 130, 159 personatus, Reduvius 43 Petia 57, 58, 59, 90, 146 Phymata 106 Picea 62, 86, 89, 104, 118, 119 picipes, Melanolestes 56, 94, 106 picturata, Ozophora 13, 19, 24, 34, 46, 47, 50, 53, 63, 71, 74, 77, 86, 67, 75, 76, 78, 82', 108 pictus, Scolopostethus 105 Piesma 71 pilicornis, Drymus 95 piliferium, Poly trichum 47, 151 pilipes, Drymus 95 198 ENTOMOLOGICA AMERICANA pilosa, Eragrostis 161 pilosulus, Antillocoris 13, 19, 35, 45, 50, 71, 62, 66 pilosulus, Scolopostethus 75 pine 50, 55, 112 pini, Rliyparochromus 78 Plantago 3, 38 plebejus, Eremocoris 122 plebejus, Heraeus 12, 19, 24, 45, 47, 50, 57, 59, 66, 70, 74, 76, 97, 101, 125 plenus, Kolenetrus 12, 20, 25, 35, 44, 46, 50, 54, 74, 88, 54, 56, 58, 151 plenus, Rliyparochromus 156 Plinthisinl 11, 12, 15, 26, 36, 59 Plinthisus 14, 38, 41, 44, 56, 62, 68, 76, 86, 59, 61 Plociomera 35 Poa 17 podagricus, Eremocoris 75 Podisus 61 Poecilomyces 90 Polygonum 105, 107, 140 Polytrichum 26 Ponerine 62 populifolia, Betula 25, 35, 62, 92, 115, 119, 27, 47, 50, 55, 56, 61, 62, 64, 79, 91, 104, 117, 118 Potentilla 4, 7, 43, 84, 145 pratensis, Poa 42 prinus, Quercus 47 Procatharosia 57, 58, 90, 146 Prytanes 35, 46 Pseudocnemodus 15, 48, 62, 88, 5, 26, 28, 146 pseudograndis, Scolopostethus 105 Pseudopamera 1 02 Ptochiomera 15, 26, 47, 56, 62, 35, 38, 40, 50 pinnila, Potentilla 36 punctata, Crataegus 68 punctiloibula, Dennstaedita 76, 108 purpurascens, Pluchea 131 purpurea, Sarracenia 93, 66 purpurem, Eupatorium 99 purslane 141 pusilla Allophora 90 Pyrrhocoridae 61 Pyrrhocoris 66, 67 Pythium 64 quadratus, Ischnodemus 134 quinque folia, Anemone 96 quinquemaculata, Ozophora 79 quadristriatus, Sphaerobius 11, 12, 19, 12, 15 Quercus 48, 55 racemosa, Smilacina 96 radicans, Rhus 125 Raglius 75 recta, Potentilla 104, 107, 3, 4 Reduviidae 4, 60, 91 repens, Agropyron 104, 3, 8, 9, 36, 38, 140, 157, 158 reperta, Ozophora 75 Rhodnius 38, 39 Rhopalidae 61 Rhyparochromini 124 Rliyparochromus 75, 156 rigida, Pinus 17, 48, 123, 135, 161 robustus, Cymus 87 roseipennis, Nabis 56 rotundifolia, Drosera 121 rubens, Picea 89, 104, 123 rubra, Festuca 26, 27, 47, 48, 67, 71, 72 rubrum, Acer 86, 107, 33, 76, 108, 118 Rudbeckia 62, 107, 115, 4, 10, 114 rufa, Formica 13, 85, 91, 113 rufipes, Acompus 26 rufipes, Malezonotus 161, 162 rugosa, Rosa 125 ruida, Ectatoma 62 Rumex 62, 92, 107, 17, 114, 141 rusticus, Stygnocoris 12, 13, 18, 20, 24, 26, 35, 44, 46, 50, 56, 74, 84, 83, 91, 94 ryei, Drymus 95 sabuleti, Ischnodemus 63, 75 sabulicolus, Megalonotus 133, 139, 157 Sagittaria 121 Salix 90 sanguinalis, Digitaria 36 Sarracenia 21 saxatilis, Spilostethus 53 scabrinodis, Myrmica 91 scabrum, Heiracium 162 Scirpus 62, 87, 121, 21, 22, 114 Scolopostethus 14, 52, 54, 55, 68, 75, 76, 84, 86, 62, 103, 111, 113, 116, 118, 119, 134 199 VOLUME XLIV scoparius, Andropogon 23, 99, 3, 12, 13, 17, 26, 31, 41, 43, 48, 50, 55, 67, 71, 72, 131, 144, 146, 151 sedge 118 sempervirens, Solidago 125 serotina, Prunus 76 serotina, Rudheckia 104, 106, 107 serphyllifolia, Veronica 48 serripes, Myodocha 12, 19, 24, 42, 45, 47, 50, 56, 57, 66, 71, 74, 91, 95, 97 servillei, Pachybrachius 21 setosus, Eremocoris 11, 13, 19, 122, 123 sigillatus, Gryllodes 56 Silene 104, 84, 140 similis, Perigenes 8, 9 simulans, Sphragisticus 139 Sisamnes 15, 35, 48, 54, 56, 62, 76, 35, 39, 43, 50 Slaterellus 41 Smilax 76 sodalicus, Melezonotus 162 Solidago 92, 99, 104, 107, 119, 4, 27, 84, 86, 91, 97, 127, 146, 164 sordidus, Elasmolomus 56, 64 Sparganium 121 spartea, Stipa 144 Spartina 62 Sphaerobius 15, 48, 49, 55, 62, 78, 89, 12, 14 Sphagnum 104, 118, 121, 13, 15, 63, 112 Sphragisticus 14, 16, 31, 62, 68, 73, 76, 85, 89, 139, 144 spicata, Danthonia 113, 48, 71 spinulosa, Dryopteris 118 spirata, Distichlis 131 Sporotrichum 158 squamosa, Solidago 47 St achy s 4 Stellaria 105 Stenocephalidae 61 stigma, Astata 153 Stilbocoris 67 Stipa 3, 12 stratuius, Anolis 26 strawberry 62, 136 stricta, Carex 25, 84, 87, 121, 21, 24, 111 strictae, Caricetum 121 strohus, Pinus 47, 48, 71, 89, 123 subulatus, Aster 131 Suffenus 54 sugarbeets 146 sulphurea, Arpliia 81 sunflower 17, 38, 43, 50, 56, 73, 74, 79, 102, 114, 121, 132, 138, 141, 148 Stygnocorini 10, 11, 12, 13, 36, 74, 75, 83 Stygnocoris 16, 18, 29, 44, 48, 54, 56, 62, 66, 68, 71, 76, 77, 84, 57, 58, 83, 84, 90, 120, 126, 135 Sylvadrymus 95 sylvaticus, Drymus 32, 95 sylvestris, Ligyrocoris 12, 15, 20, 24, ' 26, 37, 44, 46, 50, 54, 62, 74, 77, 78, 84, 102, 103, 109, 113, 117, 120, 123 sylvestris, Drymus 75 sylvestris, Pinus 123 Tanacetum 62, 84, 86, 90, 113 Tapinoma 52, 64, 72 Taraxacum 94 Tenebrionidae 27 thapsus, Verhascum 105, 141, 157, 158 thomsoni, Scolopostethus 13, 20, 24, 25, 32, 37, 42, 46, 47, 50, 52, 57, 62, 71, 74, 84, 87, 121, 103, 108 111, 116, 119, 125 Thrombidium 57, 105 Thuja 119 Thymus 84 Thyreocoris 16 Tingidae 60 Tingoidea 4 Toads 51 tormentosum, Spirea 89, 91 toxicodendron, Rhus 76 Trapezonotus 14, 15, 56, 62, 68, 70, 88, 54, 140, 150, 153, 163 tremidoides, Populus 47, 48 Trichophora 57, 61, 63 Trifolium 107 trimaculata, Cryphula 13, 19, 20, 25, 42, 45, 47, 50, 52, 58, 63, 71, 74, 88, 67, 68, 70, 71, 81 trinotata, Ozopliora 75, 79 tripunctatus, Lygaeospilus 26, -47 tropicus, Scolopostethus 103 Tsuga 62, 86, 60, 61, 114, 119, 122 typhina, Rhus 76 Tyroglyphus 9 200 ENTOMOLOGICA AMERICANA uhleri, Xestocoris 71 uliginosus, Geocoris 56, 38 umbrosus, Delochilocoris 13, 20, 24, 25, 51, 71, 74, 77, 85, 156, 158, 160 unus, Drymus 13, 19, 24, 26, 34, 44, 46, 50, 56, 70, 71, 74, 77, 84, 86, 62, 95, 103, 108, 118 Urtica 112 Yaccinium 24, 25, 35, 62, 86, 88, 114, 115, 27, 62, 72, 76, 89, 96, 108, 109, 117, 119, 151, 152 vacillans, Yaccinium 47 Veronica 62, 88, 50, 53 Viburnum 24, 62, 86, 62, 108, 109, 118 vicarius, Emblethis 13, 20, 25, 27, 46, 47, 51, 57, 59, 71, 74, 77, 42, 125, 140, 141, 144, 146, 148, 157 villosus, Bubus 162 vinctus, Pachybrachius 56, 25 virginiana, Frag aria 107, 89 virginicum, Lepidium 36, 140 vulgare, Tanacetum 104, 106, 84,. 85 89, 91, 140 vulgatum, Cerastium 84, 86 walleyi, Gastrodes 11, 13, 18, 123Y xanthoptera, Arphia 81 Xestocoris 15, 47, 49, 52, 56, 62, 68 69, 76, 87, 88, 64, 68, 71, 73 Zeridoneus 15, 48, 54, 59, 62, 108, 109 2, 3, 5, 6, 8, 14, 90, 140 201 PRINTED BY BUSINESS PRESS, INCORPORATED , <>v " Os,