vg&aay j? %. MfS ,'« lii&^m SST 7/M' Jv r X) * ■' A '^^SiSe" ' AV t\ %' ’’ •£i:'* [j, ^;tka&ei%, //^V-,f/ / \5l & S Al liiir^MtsS] 'X?# vvr. ■iS3^;.. C/ ll^s/:- ,rW ^$S§Nn XIctSa^IX ' Vft X #r®#? r% Xi fe X #} . ■'W&JS$f# Xc ,<‘isg2?l ,b Ua'AffipT , X WMW X ' V) r' s&R >-i\ '"X !!££[$■ ^ &**» " VllV/ .g^iriciag^pfgg j; gw ■ |S%' v*T i,fi ;]a;|?\ ,.1, 0! \?.V. I a- fill', ’’“■■J-j 3.-;" Vn Eh., rX l*r IT V? ^’Xji f A s II jj II fj[ ;i ; iff 1: #*• \\ Wi?CC -Hill iianJui v w&m vjl. 7 J..; \ ,4^ /<0m\ • Vy /M #Y,P N> MA luA a \ k«Ki,( ',»*4Nf.|\ ®l k ljfcf$[> V ,« fej iv 4,. X+^'i4siS| muj r vfpi mi® V ’Wm 4$ "M‘K ' i'“M wp/if iWRw ~’*P X” . v> '■'**' -C.asr ;. Vi % i* \y! ’i; ~—r- ■ il 5 a SiX*' ' '~’i' i i § of J 0^m lgip-4Aj{S (^3’^' „„ Vj^ J\ * ffy Skm -A^ USfcIStS' ^?XJX4-Sl ^ ’’wr^ri ' 6' '■^M- ■a,' u.. •«. p*» 'J'j 'A' tMw vr xa vf'W- y /M$ff 4 A> ^^^.Bfr^aSS^ &* ~V *V Ot i ( * g | X-i'-_ — ^■fX] '18 > g |J ~f, i'"'Xc#| a \a#fXffy # %, --cig^ ,,x 0 l/'n, P? .t4 ^/.w sV’" /A 4^^4j(pXi ^ 6' ., « ■i. • 11aissBi“ii3i:ia:is,l, *i, '*• y:">’ y;i<'X'V)i;1, |j>te V *** y;^;-":l|i|imnllii!i^',:!llji::i||i!; a 4, wr^M A \Q X^-i-n^k Journal of the New York ENTOMOLOGICAL SOCIETY Devoted to Entomology in General VOLUME LXXIV Published by the Society New York, N. Y. ALLEN PRESS, INC. Lawrence, Kansas INDEX OF AUTHORS ALEXANDER, CHARLES P. Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XII 66 ALEXANDER, CHARLES P. Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XIII 180 BAHADUR, J. and B. B. L. SRIVASTAVA The Nerves of the Thoracic Segments of the Larva of Prodenia litura (Lepidoptera: Noctuidae) 168 BENNETT, FREDERICK D. Notes on the Biology of Stelis ( Odontostelis ) biline olata (Spinola), a Parasite of Euglossa cordata (Linnaeus) (Hymenoptera: Apoidea: Megachilidae) 72 BROWN, F. MARTIN David Bruce (1833-1903) and Other Entomological Collectors in Colorado 126 dos PASSOS, CYRIL F. Pieris narina oleracera (Harris) in New Jersey (Lepidoptera: Pieridae) 222 FREDRICKSON, RICHARD W. An Apparent Association of Mites (Acarina) with the Rock Barnacle Balanus 101 GUPTA, A. P. Further Studies on the Internal Anatomy of the Meloidae (Coleoptera) . II. The Digestive and Reproductive Systems of the S. A. Blister Beetle, Picnoseus nitidipennis Fairmaire and Germain (Coleoptera: Meloidae) 72 HUNG, AKEY C. F. and WILLIAM L. BROWN, Jr. Structure of Gastric Apex as a Subfamily Character of the Formicinae (Hymenoptera: Formicidae) 198 IVIE, WILTON Two North American Spiders (Araneae: Linyphiidae) 224 KLOTS, ALEXANDER B. Melanism in Connecticut Panthea fur cilia (Packard) (Lep- idoptera: Noctuidae) 95 KLOTS, ALEXANDER B. Life History Notes on Lagoa laceyi (Barnes and McDun- nough) (Lepidoptera: Maegalpygidae) 140 KLOTS, ALEXANDER B. The Larva of Amblyscirtes samoset (Scudder) (Lepidop- tera: Hesperiidae) 185 LUDWIG, DANIEL and MARGARET R. GALLAGHER Vitamin Synthesis by the Symbionts in the Fat Body of the Cockroach, Periplaneta americana (L.) 134 MANISCHEWITZ, JACK R. Studies on Parasitic Mites of New Jersey 189 O’BRIEN, JAMES F. Origin and Structural Function of the Basal Cells of the Larval Midgut in the Mosquito, Aedes aegypti Linnaeus 59 ROZEN, JEROME G., Jr. Taxonomic Descriptions of the Immature Stages of the Parasitic Bee Stelis ( Odontostelis ) biline olata (Spinola) (Hymenoptera: Apoidea: Megachilidae) 84 ROZEN, JEROME G., Jr. and BARBARA L. ROZEN Mature Larvae of the Old World Bee Genus Panurgus (Hymenoptera: Apoidea) 92 iii TREAT, ASHER E. A New Blattisocius (Acarina: Mesostigmata) from Noctuid Moths 143 VASVARY, LOUIS M. Musculature and Nervous System of the Thorax, of the Sound Mechanism, and of a Typical Pregenital Abdominal Segment of the Male of the Annual Cicada, Tibicen chloromera (Walker) (Homoptera: Cicadidae) 2 VOGEL, BEATRICE R. Spiders from Powdermill Nature Reserve 55 WOOLLEY, TYLER A. and HAROLD G. HIGGINS Xenillidae, a New Family of Oribatid Mites (Acari: Cryptostigmata) 201 NOTES HOPF, ALICE L. Help for Ailing Caterpillars? 111 dos PASSOS, CYRIL F. The Discovery of Additional Journals of Frank E. Watson .... 188 BOOK REVIEWS KLOTS, ELSIE B. Monarch Butterflies by Alice L. Hopf 64 and Fireflies in Nature and the Laboratory by Lynn and Gray Poole 64 FREDRICKSON, RICHARD W. The Tarantula by William J. Baerg 109 ARNETT, ROSS H., Jr. The Beetles of the Pacific Northwest by Melville H. Hatch __ 109 MILLER, DAVID C. Wandering Through Winter by Edwin W. Teale 110 QUEDNAU, F. W. The Callaphidini of Canada by W. R. Richards 228 BROWN, F. M. A History of Entomology by O. E. Essig 229 VASVARY, L. M. Plant Galls and Gall Makers by E. P. Felt 230 RECENT PUBLICATIONS 58, 116, 164 PROCEEDINGS of the NEW YORK ENTOMOLOGICAL SOCIETY 117, 160 BYLAWS of the NEW YORK ENTOMOLOGICAL SOCIETY 103 MEMBERSHIP of the NEW YORK ENTOMOLOGICAL SOCIETY 112 NECROLOGY 122 IV V gf fjT 10 0 7 3 'j'fr 6 ^ 4 y ^ Vol. LXXIY MARCH 1966 No. 1 Devoted to Entomology in General W i-'v; -v s) y'^ K: A A' :c , FA /y - /\ ! i / H ■ I* The ■fX V , V| \y PX y M bAA New York Entomological Society m / ^'A/HA Organized June 29, 1892 — Incorporated February 25, 1893 ■ • 7) ^ 'trW A Reincorporated February 17, 1943 '//AA #:£ Vz:am /v^v^ ,\ y Jm _ ' A' ' ' XvX'AA y y Fib TaaI Xj?" - y , ■ •/ : V 1 A ’■ ''V ''' ( x ' ,\( -1 ' y ■' 1 \ ' ^ ■ ■ The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St., & Central Park W., New York 24, N. Y. A-v/ X7 Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. 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Schmitt 3 ' i ’ 0 fe- i ■ jXA 1 2 Mn Robert Buckbee j •’ i a Mailed March 3 b 1966 < rb Ab> „7,',y jb:vy( , .#-9 >: ; a,, \\ ; y.y.7, The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press Inc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas. ' A bbA ■ -/A iN': ■■ A) /H ;v $ y m f; yX ( xA-/.. i XX if: xx r ay ;ava / 7 A! ;yy . r. /; : /-X A i! i'v \ ... ' 1 ' ' . - K't- 1 . : \ . v>\,_ A j ■ - y% • 1 j , : ! . 1 ■■ 1 ) / -A i / • As A • /. ^ ' < .i Journal of the New York Entomological Society Volume LXXIV March 31, 1966 No. 1 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen Columbia University College of Pharmacy 115 West 68th Street, New York, N. Y. 10023 Associate Editor James Forbes Fordham University, New York, N.Y. 10458 Publication Committee Dr. Pedro Wygodzinsky Dr. Asher Treat Dr. David Miller CONTENTS Musculature and Nervous System of the Thorax, of the Sound Mechanism, and of a Typical Pregenital Abdominal Segment of the Male of the Annual Cicada, Tibicen ehloromera (Walker) (Homoptera: Cicadidae) Louis M. Vasvary 2 Spiders from Powdermill Nature Reserve Beatrice R. Vogel 55 Origin and Structural Function of the Basal Cells of the Larval Midgut in the Mosquito, Aedes aegypti Linnaeus James F. O’Brien 59 Recent Publications 58 Book Reviews 64 2 New York Entomological Society [Vol. LXXIV Musculature and Nervous System of the Thorax, of the Sound Mechanism, anti of a Typical Pregenital Abdominal Segment of the Male of the Annual Cicada, Tibicen chloromera ( W alker ) ( Homoptera : Cicadidae ) 1 Louis M. Vasvary Rutgers — The State University, New Brunswick, N. J. Abstract: The musculature and innervation of the thorax, sound mechanism, and the fourth abdominal segment of the male annual cicada, Tibicen chloromera (Walker) are described. The ventral nerve cord consists of a subesophageal ganglion, prothoracic ganglion, and a thoracic-abdominal ganglionic mass. There are no ganglia present in any of the abdominal segments. The prothoracic ganglion supplies innervation to some of the muscles of the cervical area and the muscles of the prothorax. The thoracic-abdominal ganglionic mass provides in- nervation to the posterior tergo-sternal muscles of the prothorax, the muscles of the pro- thorax, the muscles of the mesothorax, metathorax, and all of the abdominal segments. The abdominal segments are innervated by lateral nerve branches which arise from a pair of nerves that originate from the posterior portion of the thoracic-abdominal ganglionic mass located in the mesothorax. No median nerves are visible between the subesophageal ganglion, prothoracic ganglion, and the thoracic-abdominal ganglionic mass. The median nerves are probably included within the interganglionic connectives. The members of the family Cicadidae are among the largest insects classified in the order Homoptera. Their periodic occurrences in large numbers and the shrill “song” produced by the males have probably aroused the curiosity of man since the beginning of time. Despite their large size and the interest they have received by virtue of their sound-producing apparatus, cicadas have been some- what neglected by morphologists. This study was undertaken as a contribution to our knowledge of the musculature and innervation of the thorax, of the sound mechanism, and of a typical pregenital abdominal segment of the male of the annual cicada, Tibicen chloromera (Walker). A study of the nerve patterns in insects may be approached with at least two different objectives in mind. From a physiological or histological standpoint, a knowledge of nerve and muscle arrangements is a necessary prerequisite for pre- cise investigations. From a morphological standpoint, a knowledge of the hexapod nervous system is essential in establishing nerve and muscle homologies and thereby provide additional information on the course of phylogenetic devel- opment. This paper is an attempt in the latter direction with the full understand- ing that detailed investigations of many more forms are necessary in order to establish the course of phylogenetic development. 1 Paper of the Jour. Series, N. J. Agric. Expt. Station, Rutgers-The State University of New Jersey, Dept. Ent. and Econ. Zool. March, 1966] Vasvary: Morphology of Annual Cicada 3 13 Figure 1A Fig. 1A. Lateral view of the subesophageal ganglion of the male annual cicada Tibicen chloromera (Walker) . The concept of an underlying homology of segmental musculature has pro- vided important evidence on the evolution of the insect thorax and appendages. This concept is based on the assumption that at sometime in the past history of the Hexapoda, the abdominal somites, as leg-bearing segments, had essentially the same structure as the primitive thoracic and gnathal segments. If we assume that the innervation pattern as well as the musculature was homologous in each ancestral segment then the nerve configuration manifested in insects today is a variation of the ancestral pattern. Moreover, since the inherent purpose of the nervous system is to transmit nerve impulses, selective pressure on the nerve pattern would be less than on the structures innervated (Schmitt, 1959). This assumption should not be interpreted to imply that the nervous systems of insects have remained static in the course of phylogenetic development, but rather that through investigations of the segmental innervation patterns of insects and by establishing criteria of homology of nerves through the utilization of primitive muscle groups and nerve junctions, a knowledge of the course of the phylogenetic development of the nervous system should be possible. Unfortunately, only a very few comparative morphological investigations have been presented in the literature concerning the establishment of nerve homologies in insects. The writer hopes that this paper will be a significant addi- tion to the existing studies and serve to cultivate further interest regarding the concept of a basic plan of segmental innervation. 4 New York Entomological Society IVol. LXXIV Fig. IB. Dorsal view of ventral muscles that cover the prothoracic ganglion and anterior portion of the thoracic-abdominal ganglionic mass of the male annual cicada Tibicen chloromera (Walker). OBJECTIVES 1. Determine the musculature of the thorax of the male of the annual cicada, Tibicen chloromera (Walker) and compare this musculature to that of Hnechys sanguinea var. philaemata as described by Maki (1938) and to that of Cicada (= Tibicen ) plebeia as described by Berlese (1909). 2. Describe the ventral nerve cord of Tibicen and compare its configuration to the ventral nerve cords previously described in the family Cicadidae. 3. Determine and describe the cervicothoracic nervous system of the male of Tibicen chloromera (Walker) and, if feasible, to establish criteria of homology. 4. Determine the musculature of the first abdominal segment of Tibicen which contains the sound mechanism and compare this musculature to that described by Maki (1938) for Huechys and to that of Cicada (= Tibicen) plebeia as described by Berlese (1909). 5. Determine the innervation of the first abdominal segment of the male of T ibicen chloromera ( W alker ) . 6. Determine the musculature of a typical pregenital abdominal segment of March, 1966] Vasvary: Morphology of Annual Cicada 5 Table 1. Ventral muscles covering the prothoracic and thoracic-abdominal ganglia of Tibicen chloromera (Walker). Muscle number Origin Insertion 1 Pleural arm of prothorax Zygomatic with muscles 2 and 3 over the prothoracic ganglion and the anterior portion of thoracic-abdominal ganglionic mass. 2 Anterior margin of episternum ventral to tergo-pleural 40 Zygomatic with muscles 1 and 3 over the prothoracic ganglion and the anterior portion of thoracic- abdominal ganglionic mass. 3 Anterior mesofurcal arm Zygomatic with muscles 1 and 2 over the prothoracic ganglion and the anterior portion of thoracic- abdominal ganglionic mass. Tibicen and compare this musculature to that described by Maki (1938) for Huechys. 7. Determine the innervation of a typical pregenital abdominal segment of Tibicen and, if feasible, to establish criteria of homology. REVIEW OF LITERATURE The literature will be reviewed under five major headings corresponding to their order of presentation in this paper. 1 . THE VENTRAL NERVE CORD Comparatively little is known concerning the general nerve configuration in the families of the order Homoptera. The principal writers reporting on the ventral nerve cord of cicadas are: Binet (1894), Dufour ( 1833), Hilton (1939), and Myers (1928). It may be stated that within the family Cicadidae a high degree of specialization has taken place as far as the nervous system is con- cerned (Myers, 1928). The chief evidence of this specialization is the fact that all abdominal ganglia have become consolidated within the large thoracic-ab- dominal ganglionic mass located in the mesothorax. Binet (1894) described the subintestinal nervous system of Cicada orni. By microscopic sections of the thoracic-abdominal ganglionic mass, Binet was able to distinguish the abdominal ganglia by the absence of crural lobes correlated with the absence of legs in corresponding segments (Myers, 1928). Dufour (1833), in an earlier publication, described the ventral nerve cord in Cicada orni as having a cephalic ganglion and two thoracic ganglia. The thoracic ganglia are nearly fused, forming one oblong body which is covered dorsally by a mass of muscles which occupy the lower wall of the thorax. Dufour states that the anterior thoracic ganglion gives rise to four pairs of principal nerves, while the posterior ganglion gives rise to six pairs of nerves. The nerve cords 6 New York Entomological Society LVol. LXXIV Fig. 2. First stage dissection showing muscles of the cervix, thorax, and first abdominal segment and the ventral nerve cord in longitudinal section in the male annual cicada Tibicen chloromera (Walker). March, 1966] Vasvary: Morphology of Annual Cicada 7 which innervate the abdominal segments are adherent at their origin but separate before finally dividing in the abdominal cavity. Dufour did not describe the subesophageal ganglion which, according to Myers (1928), may have been mistaken for the brain. Hilton (1939) described the central nervous system for both the immature and adult stages of a cicada. Unfortunately, no mention is made of the species studied. There were two ganglia in both the immature and mature cicada other than the superesophageal ganglion. Hilton further stated that there are many large, long nerves issuing from the caudal portion of the large thoracic-abdominal ganglion. Myers (1928) described the brain and ventral nerve cord of Melampsalta sericea. The round subesophageal ganglion is connected to the first ganglionic mass by a pair of long, stout, well-separated interganglionic connectives. The first ganglionic mass lies largely in the prothorax. Two short, very stout inter- ganglionic connectives join the first ganglionic mass to the second thoracic mass which lies wholly within the mesothorax. Myers states that the second thoracic mass is much longer than broad and displays signs of a two fold origin. However, from the standpoint of gross anatomy, the abdominal ganglia cannot be dis- tinguished. Nerves that innervate typical abdominal segments superficially ap- pear to arise as a single cord as they leave the second thoracic mass. Later the single cord splits into two nerves as it enters the abdomen. 2. THORACIC MUSCULATURE The thoracic musculature of two species of cicadas have been described by Berlese (1909) and Maki (1938). Berlese (1909) described in some detail the thoracic musculature of Cicada ( — Tibicen ) plebeia . Muscles are identified in figures by Roman or Arabic numerals while descriptions of muscle origins and insertions are included in the text. An attempt is made to homologize the thoracic musculature of several species of insects. Unfortunately, with respect to Cicada (= Tibicen) plebeia , it appears many of the muscles that originate on the furcal and pleural arms and attach to the coxae and trochantine are omitted. Maki (1938) presents a very detailed description of the thoracic muscles of Huechys sanguinea var. philaemata. Muscles are identified by their position and function; however, in tables and figures, Arabic numerals are utilized for muscle numbers. Muscle origins and insertions are described in the text. In his study of Hemiptera, Maki presents the thoracic musculature of Eurostus validus , Sigara substriata , Cicadella ferruginea , Macrohomotoma gladiatum , and Huechys sanguinea var. philaemata . Maki includes in his tables the musculature of Nezara viridula by Malouf (1933), Cicada plebeia by Berlese (1909), and Psylla mali by Weber (1929). Snodgrass ( 1927 and 1935), illustrates a portion of the thoracic musculature of Tibicina (= Magicicada) septendecim as an example of indirect wing muscles. 8 New York Entomological Society I Vol. LXXIV Table 2. Prothoracic musculature of Tibicen chloromera (Walker). Muscle Muscle 0rigin number (or attachment) Insertion (or attachment) Dorsal muscles Median dorsal 4 Posterior edge of head First phragma Median dorsal 5 Dorsolaterally on middle of tergum First phragma Lateral dorsal 6 Dorsolaterally on middle of tergum Anterolateral region of first phragma Lateral dorsal 7 Dorsolaterally on middle of tergum Anterior edge of first phragma Anterior dorsal 8 Posterior edge of head Dorsolateral midportion of tergum Ventral muscles Internal ventral 9 Posterior tentorial arm Sternal apophyses External ventral 10 Posterior end of cervical sclerite Pleural arm of prothorax Tergo-sternal muscles Anterior intersegmental 11 Posterior edge of head Ventrolateral cervical sclerite Anterior intersegmental 12 Posterior edge of head Ventrolateral cervical sclerite Anterior intersegmental 13 Anterior dorsolateral region of tergum Posterior tentorial arm Anterior intersegmental 14 Dorsolateral region of tergum Posterior tentorial arm Anterior intersegmental 15 Anterior dorsolateral region of tergum Ventrolateral cervical sclerite Anterior intersegmental 16 Dorsolateral region of tergum Base of tentorium Posterior tergo-sternal 17 Anterolateral portion of mesotergum Pleural arm of prothorax Tergo-pleural muscles Anterior tergo-pleural 18 Dorsolateral portion of pos- terior edge of head Base of prothoracic pleural arm Anterior tergo-pleural 19 Dorsolateral portion of pos- terior edge of head Base of prothoracic pleural arm Ordinary tergo-pleural 20 Middle of lateral region of tergum Pleural arm of prothorax Coxal muscles Tergal promotor 21 Dorsolateral region of tergum Anterior rim of coxa Tergal promotor 22 Lateral region of tergum Apodeme of trochantin Sternal promotor 23 Prof urea Anterior basal rim of coxa Tergal remotor 24 Middle dorsolateral region of tergum Remotor apodeme of coxa Tergal remotor 25 Oblique ridge at middle of lateral region of tergum Remotor apodeme of coxa Tergal remotor 26 Tergum external to 25 Posterior basal rim of coxa Tergal remotor 27 Lateral region of tergum beneath 26 Posterior basal rim of coxa Sternal remotor 28 Profurca | j 1 Posterior basal rim of coxa Tergal abductor 29 Midportion of dorsolateral region of tergum Apodeme anterolateral basal rim of coxa Pleural abductor 30 Pleural arm of prothorax Anterior basal rim of coxa Pleural abductor 31 Pleural arm of prothorax Anterior basal rim of coxa Trochanteral muscles Tergal depressor 32 Midlateral region of tergum Depressor apodeme of trochanter Pleural depressor 33 Pleural arm of prothorax Depressor apodeme of trochanter March, 1966] Vasvary: Morphology of Annual Cicada 9 The muscles illustrated in the mesothorax are the longitudinal dorsal, oblique dorsal, anterior tergo-sternal, and posterior tergo-sternal. The metathoracic de- pressor muscles of the trochanter and the coxal part of the depressor muscle of the trochanter are also included. The above muscles are homologous to those of Cicada (= Tibicen ) plebeia (Berlese, 1909), Huechys sanguinea var. philaemata (Maki, 1938), and Tibicen chloromera with respect to their origins and insertions. 3. THE CERVICOTHORACIC NERVOUS SYSTEM Detailed descriptions of the thoracic nervous system have not appeared in the literature for any member of the family Cicadidae nor for any insect in the order Homoptera. Moreover, the literature contains only a relatively few studies regarding the thoracic nervous systems of insects. One reason for this lack of information is due to the time-consuming nature and patience necessary for such research. Therefore, the majority of nerve studies have been restricted to anatomical facts and descriptions of nerve cord configurations. The principal writers who have contributed detailed information on thoracic nervous systems of insects are: Holste (1910) on Dytiscus marginalis , Johansson (1957) on Oncopeltus fasciatus , Maki (1936) on Chauliodes formosanus, Marquardt (1939) on Carausius morosus , Matsuda (1956) on Agulla adnixa and Blattella germanica , Niiesch (1957) on Telea polyphemus , Pipa and Cook (1959) on Periplaneta americana , Schmitt (1959) on Dissosteira Carolina , and Wittig (1955) on Perla abdominalis. Schmitt (1962 ) states that an additional reason for the lag of nerve topography studies in insects is due to the difficulty in relating the findings on one group to those on another group. Furthermore, Maki (1936) and Pipa and Cook (1959) state that there exists a remarkable degree of variability in nerve distribution patterns of different individuals of the same insect species. However, Pipa and Cook (1959) also state that the existence of a fundamental plan in the peripheral distribution of thoracic nerves in widely separated insects is evident. Wittig (1955) describes the innervation pattern in the thorax of the larva and adult of Perla abdominalis . She presents a comparison of the innervation fields of the thoracic nerves of Perla abdominalis with those of Chauliodes formosanus as reported by Maki (1936), Carausius morosus as reported by Marquardt (1939), and Dytiscus marginalis as reported by Holste (1910) and establishes the existence of nerve homologies in these widely separated insects. Pipa and Cook (1959) state that the pattern of nerve distribution in Peri- planeta americana essentially agrees with that found in other insects which have been investigated. A similar indication in Periplaneta americana was made by Nijenhuis and Dresden (1955). Schmitt (1959) describes the cervicothoracic nervous system of Dissosteira Carolina and presents several areas of nerve homology with respect to Chauliodes trx 10 New York Entomological Society [Vol. LXXIV Fig. 3. Second stage dissection showing muscles of the cervix, thorax, and first abdominal segment in longitudinal section in the male annual cicada Tibicen chloromera (Walker). March, 1966] Vasvary: Morphology of Annual Cicada 11 formosanus as reported by Maki (1936). However, he states that the ptero- thoracic dorsal nerves pass beneath the ventral longitudinal muscles and differ in this respect from the prothoracic dorsal nerves of Dissosteira and all the thoracic dorsal nerves of Chauliodes. Schmitt compared the nerves of the pro- thoracic muscles of Dissosteira with those of the pterothorax and concluded that there is evidence of a loss of anterior prothoracic musculature as a result of the evolution of the cervix. The cephalic muscles of the cervical sclerites and the ventral lateral neck muscles are derived from this anterior prothoracic muscula- ture. Schmitt also includes a comparative study of the anterior ganglionic con- nectives of the dorsal nerves of Dissosteira , Periplaneta , and Orchelimum and indicates that the anterior ganglionic connectives of the dorsal nerves may have a wider distribution than in Orthoptera but are not recognizable because of juxtaposition with the connectives of the ventral nerve cord. Schmitt describes the median nerves and the innervation of the spiracular muscles in Dissosteira and mentions that the transverse nerves, dorsal nerves, and the innervation of the spiracular muscles of Chauliodes as described by Maki (1936) present a pattern identical with that in Dissosteira. There appears to be no essential dif- ferences in the innervation pattern of the thoracic spiracles as compared with the innervation pattern of the abdominal spiracles in both Chauliodes and Dissosteira. Schmitt concludes that the nerves to the thoracic spiracles agree sufficiently with the nerve pattern of the abdominal spiracles to indicate that the thoracic spiracles may be homologous with the abdominal spiracles. Schmitt (1962), in a later paper, despite unfortunate differences in nomen- clature applied by different workers, presents additional information establishing the presence of nerve homologies in several insects. Schmitt utilizes the dorsal longitudinal muscles as a starting point since these muscles are homologous both in the thorax and abdomen of insects. Usually, from a descriptive standpoint it is quite simple to identify the dorsal nerves to these muscles. Schmitt arranges in tabular form the names and designations used by various authors for the nerves to the thoracic dorsal longitudinal muscles, designations of the anterior ganglionic connectives, designations of the subesophageal nerves to the protergal muscles, and a comparison of thoracic nerve designations used by various authors with those utilized by Maki for Chauliodes. The wing nerves, median and transverse nerves, innervation of the ventral muscles and spinosternal musculature, and a discussion of the prothoracic nervous system in various insects is also presented. 4. THE MUSCULATURE AND INNERVATION OF THE SOUND MECHANISM The majority of investigations appearing in the literature concerning the sound mechanism of cicadas describes the construction of the sound apparatus and the mechanics of sound production. Myers (1928) presents a summary of the studies pertaining to the sound-producing apparatus as well as including his 12 New York Entomological Society [Vol. LXXIV Table 3. Mesothoracic musculature of Tibicen chloromera (Walker). Muscle Muscle Origin Insertion number (or attachment) (or attachment) Dorsal muscles Median dorsal 34 Anterior median portion of tergum Median area of second phragma Lateral dorsal 35 Middle of dorsolateral por- tion of tergum Lateral portion of second phragma Ventral muscles Longitudinal ventral 36 Profurcal arm xMiterior mesofurcal arm Tergo-Sternal muscles Anterior tergo-sternal 37 Anterior portion of dorso- lateral region of tergum Ventrolateral sternal region Posterior tergo-sternal 38 Ventral portion of second phragma Posterior mesofurcal arm Tergo-Pleural muscles Tergo-pleural 39 Anterolateral margin of tergum Anterior margin of episternum Tergo-pleural 40 Lateral margin of tergum Anterior margin of episternum Tergo-pleural 41 Lateral margin of tergum Mesothoracic pleural arm Tergo-pleural 42 Lateral margin of tergum Prothoracic pleural arm Tergo-pleural 43 Lateral margin of tergum Wing process Tergo-pleural 44 Lateral margin of tergum Base of mesothoracic pleural arm Pleural-axillary 45 Episternum Third axillary sclerite Pleural-axillary 46 Episternum Third axillary sclerite Pleuro-subalar 47 Posterior margin of epimeron Subalar sclerite Sterno-Pleural muscles Sterno-basalar 48 Anterodorsal portion of epi- sternum Ventrolateral sternal region Furco-entopleural 49 Furcal arm of mesothorax Pleural arm of mesothorax Coxal muscles Tergal promotor 50 Anterolateral region of tergum Trochantin Trochantino-basalar 51 Laterodorsal margin of epi- sternum Trochantin Trochantino-basalar 52 Anterolateral margin of epi- sternum Trochantin Sternal promotor 53 Base of mesofurcal arm Anterior basal rim of coxa Tergal remotor 54 Anterolateral region of tergum Posterior basal rim of coxa Tergal remotor 55 Posterior dorsolateral region of tergum By a tendon to posterior basal rim of coxa Coxo-subalar 56 Posterior basal rim of coxa Subalar sclerite Sternal remotor 57 Posterior mesofurcal arm Posterior basal rim of coxa Sternal remotor 58 Mesofurcal arm Posterior basal rim of coxa Sternal adductor 59 Mesofurca Mesal basal edge of coxa Coxo-basalar 60 Dorsal margin of episternum Anterolateral basal rim of coxa Troehanteral muscles Tergal depressor 61 Anterolateral portion of tergum Depressor apodeme of trochanter Trochantero-basalar 62 Dorsal margin of episternum Depressor apodeme of trochanter Sternal depressor 63 Mesofurcal arm Depressor apodeme of trochanter Muscles of the spiracle Occlusor 64 Subspiracularum Ventral portion of atrial chamber March, 19661 Vasvary: Morphology of Annual Cicada 13 own findings based on M damp salt a sericea and M damp salt a muta , two species of cicadas found in New Zealand. Complete studies regarding the musculature of the first abdominal segment which contains the sound-producing apparatus have been described for Cicada (— Tibicen) plebeia by Berlese (1909) and for Huechys sanguinea var. philae- mata by Maki (1938). Berlese utilizes both Roman and Arabic numerals for muscle identification in his figures while descriptions of muscle attachments are included in the text. Berlese (1909) shows the structure of the sound mechanism in his figures 879 to 882. Berlese considers the sclerotized V-shaped structure, yAd2 in his figure 880, as the furca of the second abdominal sternite. However, Carlet (1876), Vogel (1923) and Myers (1928) who have given this structure the most attention, ascribe it to the first abdominal segment. Maki (1938) shows the musculature of the sound mechanism in his figure 24 and utilizes Arabic numerals for muscle numbers. Maki presents in tabular form the muscles of the first six abdominal segments with their muscle numbers. Descriptions of the muscle attachments are not included in the text. A complete presentation of the innervation pattern of the first abdominal seg- ment of cicadas has not appeared in the literature. However, the auditory or tymbal nerves which innervate the large tymbal muscles have been mentioned by various writers since Binet (1894). Swinton (1880) traced the auditory nerve from the thoracic ganglionic mass, presumably in the mesothorax, to the abdomen and around the tymbal muscle. The auditory nerve then forms a ganglion which enters a groove. According to Vogel (1923) the auditory nerve arises in the ventral nerve strands and rises, running parallel with the body wall, in a sclerotized groove and passes dorsally to the sense organ, where its fibers run into the base of each sense cell. Myers (1928), in poorly preserved material, found a distinct nerve emerging on each side of the last thoracic-ganglionic mass and running parallel to a sclerotized ridge leading up to the auditory capsule. Myers (1928) states that it is very improbable that the auditory nerve should arise from the abdominal strands, as Vogel ( 1923) states. Investigations utilizing electric stimulation of the auditory or tymbal nerve and the sympathetic nerve have appeared in the literature. Pringle (1954) concluded that the frequency of tymbal movements resulting from the contrac- tions of the tymbal muscle exceeds the rhythm of tymbal nerve stimulation. Pringle also reported that an isolated tymbal muscle does not give multiplied rhythmic reactions when stimulated but functions the same as a common skeletal muscle. Hagiwara and Watanabe (1956) found that at a certain intensity and frequency of nerve stimulation, repetitive potentials up to ten or more resulted from each stimulus in the tymbal muscle, tymbal nerve, and motor neuron. Voskresenskaya and Svidersky (1960) investigated the electrical activity of the tymbal muscle, the tymbal nerve, and the sympathetic nerve during and 14 New York Entomological Society [Vol. LXXIV Fig. 4. Third stage dissection showing muscles of cervix, thorax, and first abdominal seg- ment in longitudinal section in the male annual cicada Tibicen chloromera (Walker). March, 1966] Vasvary: Morphology or Annual Cicada 15 after electric stimulation and concluded that the sympathetic nervous system is essential to normal sound production in cicadas. 5. THE MUSCULATURE AND INNERVATION OF THE FOURTH ABDOMINAL SEGMENT The musculature of the pregenital abdominal segments of male cicadas have been described by Maki (1938) for Huechys sanguined var. philaemata and by Berlese (1909) for Cicada (= Tibicen) plebeia. Maki in his figure 24 shows the musculature of the first three abdominal segments and utilizes Arabic numerals for muscle numbers. Maki presents the muscles of the first six abdominal seg- ments and their muscle numbers in a table on page 168 where he compares the musculature of Erostus validus , Sigara sub striata , Huechys sanguined var. philaemata, Cicadella ferruginea, and Macrohomotoma gladiatum. Maki does not describe the muscle attachments for Huechys in his text; however, they are clearly shown in his figure 24. Berlese (1909) describes the musculature of the first three abdominal segments in Cicada (= Tibicen) plebeia and utilizes both Roman and Arabic numerals for muscle identification. Descriptions of the muscle attachments are included in the text. No studies dealing with the innervation of a pregenital abdominal segment of a male cicada have been found in the literature. Moreover, the literature con- tains only a few studies on the abdominal nervous system of insects. In recent years some interest has been shown regarding the establishment of basic segmental nerve pattern within the Hexapoda. Schmitt ( 1954) describes the nervous system of the pregenital abdominal segments of Dissosteira Carolina , Acheta assimilis, Periplaneta americana , and Diapheromera femorata. Schmitt utilizes various points of nerve homology or “landmarks” in presenting the in- nervation pattern of the above insects. The innervation of the ventral diaphragm in Dissosteira is also described. Libby (1959) describes the musculature and innervation of the second and third abdominal segments of the cecropia larva and concludes that the dorsal, ventral, and transverse nerve roots arising from each segmental ganglion of the cecropia larva seem homologous with those de- scribed by Schmitt (1954) for the pregenital segments of certain Orthoptera. Libby concludes, by utilizing the points of nerve homology set forth by Schmitt, that the homogeneity of the innervation pattern in such widely separated orders as Orthoptera and Lepidoptera lend further support to the concept of a basic segmental nerve pattern within the Hexapoda. Libby (1961) describes the mus- culature and innervation in the fourth abdominal segment of the adult male cecropia moth Hyalophora cecropia and compares his finding with the pregenital abdominal segments of Chauliodes formosanus , as described by Maki (1936), Acheta assimilis , as described by Schmitt (1954), and the larva of Hyalophora , as described by Libby ( 1959). Schmitt (1963) describes the abdominal nervous system in the nymph of Pteronarcys proteus and the adult of Pteronarcys calif ornica and concludes that 16 New York Entomological Society [Vol. LXXIV "7 > u Fig. 5. Fourth stage dissection showing muscles of the cervix, thorax, and first abdominal segment in longitudinal section in the male annual cicada Tibicen chloromera (Walker). March, 19661 Vasvary: Morphology or Annual Cicada 17 the ganglia of segments 3 and 4 have coalesced and only the first three segments contain both dorsal and ventral nerves. The transverse nerves of segments 4, 5, and 6 arise from the ganglia of the immediately following segments. No occlusor or dilator muscles of the spiracles could be found in the two above-men- tioned species of Pteronarcys. Schmitt also describes the muscles and nerves of the genital segments. Schmitt (1964) describes the nerve pattern of the pregenital abdominal seg- ments of N eoconocephaiis exiliscanorus and Cento philus gracilipes gracilipes, two Orthoptera classified in the family Tettigoniidae. The segmental nerve patterns of these two insects were comparable and conformed to the patterns described in the Acrididae, the Gryllidae, and the Blattidae, as described by Schmitt (1954), and in Carausius (Phasmidae) as described by Marquardt (1939). Similarities in the nerve patterns to Hyalophora cecropia as described by Libby (1959 and 1961) and by Beckel (1958) and in some degree to the Plecoptera and the Megaloptera were noted. No innervation to the alary muscles could be found in N eoconocephalus or Ceutophilus. Schmitt (1965) presents a comparative study on the transverse nerves of the pregenital abdominal segments of insects. By comparing the segmental innervation patterns of Peri planet a, N eoconocephalus, Hyalophora, Chauliodes , Pteronarycs, Acroneurai, Apis, and Tibicen, Schmitt concludes that, in those insects which apparently lack median and transverse nerves, these nerves are in- corporated in the longitudinal connectives and lateral segmental nerves. MATERIALS AND METHODS Insect Material Used in the This Study. — The male of the annual cicada, Tibicen chloromera (Walker), was selected for this study in order to provide information concerning the musculature and nervous system of the thorax, sound mechanism, and a typical pregenital abdominal segment. The annual cicada’s large size and ready availability make them especially attractive subjects for such investigation. N omenclature. — Nomenclature used in this study involve primarily the muscula- ture and nervous system. Various methods of nomenclature have been devised for each of these organ systems. Nomenclature of the musculature is based on the general outline set forth by Maki (1938) in his work on Huechys sanguinea var. philaemata. Muscles are named according to their position, attachment, or function and are assigned Arabic numerals which serve as muscle numbers in figures. Effective nerve nomenclature requires not only that it describe the nerves in question, but also that it can be applied or adapted to as many nervous systems as possible in order to demonstrate nerve homologies. However, before a stan- dard terminology can be devised, it is essential to have a relatively thorough knowledge of the musculature and nervous systems of manj^ different insect 18 New York Entomological Society [Vol. LXXIV Table 4. Metathoracic Musculature of Tibicen chloromera (Walker) | Muscle Origin Number (or attachment) Insertion (or attachment) Dorsal muscles Median dorsal 65 Dorsal portion of second phragma Dorsal portion of third phragma Ventral muscles Longitudinal ventral 66 Posterior mesothoracic furcal arm Metafurcal arm Tergo-Sternal muscles Anterior tergo-sternal 67 Anterior dorsolateral region of tergum Ventrolateral sternal region Posterior tergo-sternal 68 Anterolateral edge of first abdominal tergum Metafurcal arm Tergo-Pleural muscles Tergo-pleural 69 Lateral portion of tergum Pleural arm of metathorax Tergo-pleural 70 Lateral portion of tergum Dorsal border of episternum Pleuro-axillary 71 Pleural ridge Third axillary sclerite Pleuro-axillary 72 Pleural ridge Third axillary sclerite Sterno-Pleural muscles Sterno-pleural 73 Mesofurcal arm Anterior end of metathoracic episternum Furco-entopleural 74 Metafurcal arm Pleural arm of metathorax Coxal muscles Tergal promotor 75 Anterior dorsolateral region of tergum Trochantin Pleural promotor 76 Anterior portion of epi- sternum Anterior basal rim of coxa Sternal promotor 77 Metafurcal arm Anterior basal rim of coxa Tergal remotor 78 Mid dorsolateral region of tergum Posterior basal rim of coxa Tergal remotor 79 Posterior dorsolateral region of tergum Posterior basal rim of coxa by a tendon Coxo-subalar 80 Lateral basal rim of coxa Subalare Sternal remotor 81 Metafurcal arm Posterior basal rim of coxa Sternal remotor 82 Metafurcal arm Posterior basal rim of coxa Pleural abductor 83 Anterior region of epi- sternum lateral to 75 Anterolateral basal rim of coxa Trochanteral muscles Tergal depressor 84 Second phragma Depressor apodeme of trochanter Tergal depressor 85 Anterior portion of dorso- lateral region of tergum Depressor apodeme of trochanter Pleural depressor 86 Episternum Depressor apodeme of trochanter Sternal depressor 87 Metafurcal arm Depressor apodeme of trochanter Muscles of the spiracle Occlusor 88 Ridge between mesothorax and metathorax Ventral end of spiracle species. Several systematic methods of nerve terminology have been devised and each have their advantages and disadvantages. The method of nerve designation utilized in this paper is similar to that used by Whittig (1955) in her work on Perla abdominalis Burm. Ganglia, except for the subesophageal ganglion, are assigned Roman numerals. Nerve roots arising from each ganglion are designated by the Roman numeral of the ganglion followed March, 1966] Vasvary: Morphology op Annual Cicada 19 by the letter N and an Arabic numeral. Lower case letters following Arabic numerals are used to identify nerve branches. Prime (') and double prime (") designations are utilized where it appears necessary for better understanding of nerve branch description. Methods of Illustration. — Illustrations in this paper representing nerves and muscles are of two types. One type, the semiperspective illustration, is an at- tempt to represent as clearly as possible the various stages of dissection. Each stage is illustrated separately and in series beginning with the median muscle groups and progressing to the body wall. In illustrations that combine two consecutive stages of dissection, the lower half of the figure represents the earlier stage. The second type of illustration used in this study are diagrams indicating the spatial relationships of nerves. The right side of the insect is illustrated and viewed in a laterad aspect. Muscle innervations are designated by Arabic nu- merals which represent muscle numbers. Where two nerves cross, the laterad nerve is interrupted. Nerves which terminate in the integument are indicated by a short line drawn across the nerve. An explanation of abbreviated designations may be found under “Abbrevia- tions used in the Figures” at the conclusion of this paper. RESULTS AND DISCUSSION 1. THE VENTRAL NERVE CORD General: The ventral nerve cord of insects is the postcephalic portion of the nervous system which lies beneath the alimentary canal and extends posteriorly through the thorax and abdomen. This portion of the central nervous system contains the subesophageal ganglion, thoracic ganglia, and abdominal ganglia arranged metamerically and joined by paired longitudinal connectives. However, modifications of the above generalized ventral nerve cord exists in a number of insect orders and is evidenced by the reduction in number or complete absence of ganglia in abdominal segments. Snodgrass (1935) states that there is a tendency for the ganglia of the ventral nerve cord to migrate anteriorly and unite with each other. This process is referred to as condensation. The forward migration and fusion of ganglia results in the shortening and external disap- pearance of connectives and commissures. A dorsal view of the ventral nerve cord in the male cicada, Tibicen chloromera (Walker) is illustrated in Fig. 9 and consists of a subesophageal ganglion, pro- thoracic ganglion, and a thoracic-abdominal ganglionic mass. There are no ganglia in any of the abdominal segments. All abdominal segments are innervated by nerves originating from the posterior portion of the thoracic-abdominal ganglionic mass located in the mesothorax. The subesophageal ganglion is the anterior ganglion of the ventral nerve cord. 20 New York Entomological Society [Vol. LXXIV Table 5. Comparison of prothoracic musculature of Tibicen chloromera , Huechys sanguined var. philaemata (Maki, 1938), and Cicada (— Tibicen) plebeia (Berlese, 1909). Muscle groups Tibicen chloromera Huechys sanguined var. philaemata (Maki, 1938) Cicada (= Tibicen) plebeia (Berlese, 1909) Dorsal muscles Median dorsal 4 1 140 Median dorsal 5 2 CIX Lateral dorsal 6 3 110 Lateral dorsal 7 — CXII Anterior dorsal 8 4 CXXXVI Anterior dorsal - - cxxxv Ventral muscles Internal ventral 9 5 136 External ventral 10 6 CXXXI Tergo-Sternal muscles Anterior intersegmental 11 7 147 Anterior internal tergo-sternal 12 — - Anterior internal tergo-sternal 13 8 CXXXV Anterior internal tergo-sternal 14 — — Anterior internal tergo-sternal 15 9 CXXXVa Anterior internal tergo-sternal — 10 144 Anterior internal tergo-sternal 16 11 — Posterior tergo-sternal 17 12 112 Tergo-Pleural muscles Anterior tergo-pleural 18 13 - Anterior tergo-pleural 19 - - Ordinary tergo-pleural 20 14 - Coxal muscles Tergal promotor 21 15 113 Tergal promotor 22 16 - Sternal promotor 23 17 - Tergal remotor 24 18 116 Tergal remotor 25 19 - Tergal remotor 26 20 - Tergal remotor 27 21 — Sternal remotor 28 — — Tergal abductor 29 22 - Pleural abductor 30 23 — Pleural abductor 31 24 - Troehanteral muscles Tergal depressor 32 25 115 Pleural depressor 33 26 — In Tibicen chloromera eight pairs of nerves arise from the ganglion and innervate the salivary glands and lateral salivary gland ducts, muscles associated with' the feeding apparatus, and some muscles of the cervical area. The prothoracic ganglion and the anterior portion of the thoracic-abdominal ganglionic mass are covered dorsally by ventral muscles (Fig. IB). Dufour ( 1833) mentions similar ventral muscles in Cicada orni. An invagination of the first abdominal sternite serves as a muscle attachment for the large tympanal muscles. A sternal canal is located within this invagina- tion. Two pairs of nerves, IIN8 and IIN9, pass through the sternal canal. March, 19661 Vasvary: Morphology of Annual Cicada 21 Fig. 6. Fifth stage dissection showing muscles of the cervix, thorax, and first abdominal segment in longitudinal section in the male annual cicada Tibicen chloromera (Walker). 22 New York Entomological Society [ Vol. LXXIV One pair of nerves, IIN8, innervates the posterior muscles of the first abdominal segment while the other pair of nerves, IIN9, innervates the remaining abdominal segments. No median nerve is visible between the subesophageal ganglion, prothoracic ganglion, and thoracic-abdominal ganglionic mass. However, the median nerve is probably included within the interganglionic connectives. Spiracular muscles in the thoracic segments are innervated by nerves which arise from the dorsolateral portion of the prothoracic ganglion and thoracic- abdominal ganglionic mass. Spiracular muscles in pregenital abdominal seg- ments are innervated by a nerve branch from the dorsal nerve. The ventral nerve cord of the male Tibicen chloromera (Walker) is not re- stricted to a definitive positional relationship in the thorax by spinae or muscles that attach to these structures. Schmitt (1959) described an opposite situation in the thorax of Dissosteira , where possible future evolution of the ventral nerve cord towards condensation will require drastic skeletal and muscle system changes. Subesophageal Ganglion: The anterior portion of the subesophageal gan- glion is covered by the tentorial bridge (TB, Fig. 9). A pair of short, stout cir- cumesophageal connectives link the subesophageal ganglion to the brain. A lateral view of the subesophageal ganglion is shown in Fig. 1A. Eight pairs of nerves arise from the ganglion, five pairs of nerves from the lateroventral surface, and three pairs from the ventral area. The first pair of nerves, SN1, arise from the anterior medioventral surface of the ganglion in close association with the ventral portions of the circumesopha- geal connectives. SN1 nerves divide into labral nerves (LmNv) and nerves which innervate the protractor muscles of the mandibular bristles (pmdb). The second pair of nerves, SN2, are mandibular nerves and arise from the anterior lateroventral surface of the ganglion. The SN2 nerve divides soon after leaving the ganglion into a dorsal branch that innervates the retractor muscle of the mandibular bristle (rmdb) and a ventral branch that innervates the protractor muscles of the mandibular bristles. The third pair of nerves, SN3, are maxillary nerves and arise from the latero- ventral surface of the ganglion. The SN3 nerve bifurcates into anterior and posterior nerve branches. The anterior branches innervate the internal (rmxbi) and the external (rmxb2) retractor muscles of the maxillary bristles. Posterior nerve branches innervate both internal and external retractor muscles of the maxillary bristles, protractor muscles of the maxillary bristles (lpmxb and 2pmxb), and provide nerve branches which enter the base of the maxillary bristles, mxb (Fig. 1A). The fourth, SN4, and fifth, SN5, pairs of nerves arise from the mediolateral and posterolateral areas, respectively, of the subesophageal ganglion and co- March, 1966] Vasvary: Morphology of Annual Cicada 23 g o *-4-> CJ C3 C 3 4-> • bu g o in O -C 13 p a Ui c £ ? -si rt 03 c_) -*-> ..-c m u rG ■*-> c7) o M c3 3 C C <3 OJ 13 a OJ -C alesce to form the first cervical nerve SN4 + SN5. The SN4 + SN5 nerves ex- tend dorsally and innervate the anterior internal tergo-sternal muscle 13. The sixth pair of nerves, SN6, arises from the posterior lateroventral surface of the ganglion and innervates the salivary glands (S1G1) and the anterior in- ternal tergo-sternal muscle 16. The latter nerves are the second cervical nerves. A nerve branch from SN6, and SN6a, proceeds in a posterior direction and coalesces with the SN9 + INI nerve. The seventh pair of nerves, SN7, are the labial nerves and arise from the posterior medioventral surface of the ganglion ventrad to SN6. Nerve SN7 pro- 24 New York Entomological Society [Vol. LXXIV vides nerve branches to the dilator muscles of the salivary syringe (dlSyr) and the lateral muscles of the sclerotized rod (mr) before entering the labium. Johansson ( 1957) shows a similar innervation pattern for the labial nerve in the milkweed bug One o pelt us fasciatus (Dallas). The labial nerves innervate the dilator muscles of the salivary syringe before entering the base of the labium. The eighth pair of nerves, SN8, arises from the posterior medioventral sur- faces of the ganglion in close association with the interganglionic connectives and innervates the salivary ducts (SID). A pair of large nerves, SN9, arises laterally from each interganglionic connec- tive and coalesce with the IN 1 nerves which arise from the anterior surface of the prothoracic ganglion. INI nerves can easily be separated from the intergangli- onic connectives to their origin on the prothoracic ganglion. A pair of long, sturdy, well-separated interganglionic connectives link the subesophageal ganglion to the prothoracic ganglion. The interganglionic con- nectives pass laterally around the muscles of the sclerotized rod. The sclerotized rod is an extension of the labium and appears to have a stronger association with the prothorax than with the head since it hangs freely from the cervical mem- brane. Prothoracic Ganglion: The prothoracic ganglion lies wholly within the pro- thorax and is situated between the sternal apophyses. Three pairs of ventral muscles (1, 2, and 3) cover the entire ganglion dorsally (Fig. IB). Four pairs of nerves arise from the anterior portion of the prothoracic ganglion (Fig. 9). The anterior pair of nerves (INI) proceed anteriorly and join nerves SN9 which branch from the interganglionic connectives. Nerves IN2, IN3, and IN4 pass under the internal ventral longitudinal muscles (9) and innervate muscles of the cervix, prothorax, and prothoracic leg muscles. (A detailed de- scription of the innervation pattern is presented under the section entitled “The Cervicothoracic Nervous System.”) A pair of large nerves, IIN1, issues from the interganglionic connectives be- tween the prothoracic ganglion and thoracic-abdominal ganglionic mass (Fig. 9). The IIN1 nerves pass over the IIN2 nerves originating from the thoracic- abdominal ganglionic mass and then pass under the longitudinal ventral muscles 36 of the mesothorax. Nerve IINl innervates the longitudinal ventral muscles 36, median dorsal muscles 34, and lateral dorsal muscles 35 of the mesothorax. A pair of fine, short nerves, IN5, arise on each side of the middorsal portion of the prothoracic ganglion and innervates the ventral muscles 1 which cover the ganglion. Two pairs of fine nerves, IN6 and IN 7, arise from the middorsal area of the prothoracic ganglion and coalesce with nerve INS arising from the dorsal surface of the interganglionic connective between the prothoracic ganglion and thoracic- abdominal ganglionic mass. March, 1966 1 Vasvary: Morphology of Annual Cicada 25 A pair of very short, stout interganglionic connectives links the prothoracic ganglion to the large thoracic-abdominal ganglionic mass. Thoracic— Abdominal Ganglionic Mass: The thoracic-abdominal gangli- onic mass is the terminal ganglion of the ventral nerve cord and is located above the basisternum of the mesothorax. With the exception of the XIN2a nerves which innervate the posterior tergo-sternal muscles 17 of the prothorax, nerves originating from the thoracic-abdominal ganglionic mass innervate muscles of the mesothorax, metathorax, sound mechanism, and abdominal segments. Eight pairs of lateral nerve roots arise from the thoracic-abdominal ganglionic mass: one pair anteriorly, IIN2; two pairs laterally, TIN3 and IIN4; and five pairs posteriorly, IIN5, 1 1X6. IIN7, IIN8, and TIN9 (Fig. 9). Nerves IIN2, IIN3, and IIN4 pass under the longitudinal ventral muscles 36 while the remain- ing nerve roots extend posteriorly and pass over the mesofurca. IIN2 is the anterior wing nerve while IIN3 and IIN4 innervate muscles in the mesothorax. Nerves IIN5 and IIN6 pass under the posterior arms of the mesofurca and inner- vate muscles of the metathoracic segment with the exception of nerve branch IIN6a' which innervates the posterior tergo-pleural muscle 38 of the mesothorax. The 1 1 N 5 nerve is the dorsal nerve since it innervates the dorsal muscles 65. Ventral muscles are innervated by a nerve branch from IIN10 + IIN11. Nerve IIN6 provides a nerve branch IIN6a which is the posterior wing nerve. Nerves IIN7 supply innervation to the muscles located in the anterior portion of the first abdominal segment and the membrane forming the large abdominal air chamber. Nerves IIN8 provide nerve branches IIN8a to the large tympanal muscles before passing through the sternal canal to innervate the muscles located in the posterior portion of the first abdominal segment. The IIN9 nerves pass through the sternal canal and innervate muscles of the remaining pregenital abdominal segments by providing a lateral nerve branch to each consecutive segment. Two pairs of fine nerves (IIN10 and IIN11) arise dorsolaterally from the thoracic-abdominal ganglionic mass (Fig. 9). The IIN11 nerve divides soon after leaving the ganglion and provides a fine nerve branch IINlla which coalesces with nerve IIN5. Nerves IIN10 and XIN11 are connected by a fine nerve designated as XIN10 + IIN11. It appears that both the IIN10 and IIN11 nerves are responsible for innervation of the occlusor muscle (88) of the meta- thoracic spiracle and ventral muscle 3. Discussion: Unfortunately, only the gross anatomy of the central nervous sys- tem of cicadas has been described in the literature. Therefore, comparisons of ventral nerve cords in order to establish areas of homology are limited to their general configuration. Hilton (1939) in his Figure 190-1 presents an unlabeled drawing of the cen- tral nervous system of an unnamed adult cicada showing the brain and two gan- glia of the ventral nerve cord. If it is assumed that the anterior ganglion is the 26 New York Entomological Society rVoL. LXXIV Table 6. Comparison of mesothoracic musculature of Tibicen chloromera, Huechys san- guined var. philaemata (Maki, 1938), and Cicada {— Tibicen ) plebeia (Berlese, 1909). Muscle groups Tibicen chloromera Huechys sanguined var. philaemata (Maki, 1938) Cicada (— Tibicen ) plebeia (Berlese, 1909) Dorsal muscles Median dorsal 34 27 70 Median dorsal — — 69 Lateral dorsal 35 28 71 Ventral muscles Longitudinal ventral 36 29 105 + 106 Spino-furcal ventrals - - 104 Tergo-Sternal muscles Anterior tergo-sternal 37 30 LXXVIII Posterior tergo-sternal 38 31 73 Tergo-PIeural muscles Tergo-pleural 39 32 XCI Tergo -pleural 40 33 86 Tergo-pleural 41 34 - Tergo-pleural 42 - - Tergo-pleural 43 - - Tergo-pleural 44 - - Pleuro-axillary 45 35 XC1II Pleuro-axillary 46 36 XCII Pleuro-subalar 47 37 - Sterno-Pleural muscles Sterno-basalar 48 38 91 Furco-entopleural 49 39 100 Coxal muscles Tergal promotor 50 40 74? Trochantino-basalar 51 - 79 + SO Trochantino-basalar 52 - - Sternal promotor 53 41 - Tergal remotor 54 42 LXXXII Tergal remotor 55 43 75 Coxo-subalar 56 44 84 Sternal remotor 57 45 - Sternal remotor 58 - - Sternal adductor 59 - - Pleural abductor - 46 - Coxo-basalar 60 47 82 Trocha literal muscles Tergal depressor 61 48 76 Trochantero-basalar 62 49 81 Sternal depressor 63 50 - Muscles of the spiracle Occlusor 64 51 — subesophageal ganglion, then the remaining ganglionic mass contains all of the thoracic and abdominal ganglia. Dufour (1833), describing the ventral nerve cord in Cicada orni , states that the central nervous system consists of a cephalic ganglion and two thoracic ganglia. No mention is made of the subesophageal ganglion, which, according to Myers (1928), Dufour may have confused with the brain. Dufour does March, 1966] Vasvary: Morphology of Annual Cicada 27 mention that the cephalic ganglion is produced by a fusion of two hemispheroid lobes and the cleft which separates the two lobes is only superficial. Dufour continues by describing the thoracic ganglia as not being separate and distinct but nearly fused into one. However, with difficulty, a light demarcation of an anterior ganglion can be observed. Myers (1928) states that the ventral nerve cord in Melampsalta sericea consists of a subesophageal ganglion, prothoracic ganglion, and thoracic-ab- dominal ganglionic mass, each separated by visible interganglionic connectives. Berlese (1909), in his Figure 697, presents a diagram of the brain and the subesophageal ganglion of Cicada (= Tibicen) plebeia , and shows that the subesophageal ganglion is separated from the brain by a pair of stout circum- esophageal connectives. The remainder of the ventral nerve cord is not described. Snodgrass (1935), in his Figure 237, presents a longitudinal section of Tibicina (— Magicicada) septendecim showing two thoracic ganglia, one in the prothorax and the other in the mesothorax. The subesophageal ganglion is not illustrated. If the above investigations are correct, then there appears to be some diversity in the family Cicadidae regarding the number of ganglia in the ventral nerve cord. Cicada orni is the most specialized with a central nervous system composed of a cephalic ganglion and two very closely associated thoracic ganglia while in Melampsalta sericea and Tibicen chloromera there is a subesophageal ganglion and two separate thoracic ganglia. There also appears to be a diversity in the number of principal lateral nerve roots arising from the thoracic ganglia. Dufour (1833) mentions that the an- terior thoracic ganglion in Cicada orni gives rise to four pairs of principal nerves while six pairs of nerves issue from the posterior thoracic ganglion. Hilton (1939), in his Figure 190-1, of the central nervous systems of an unnamed species of cicada, shows three principal nerves arising from the anterior lobe of the thoracic-abdominal ganglionic mass while the posterior lobe possesses three pairs of lateral nerves and a single caudal nerve. Tibicen chloromera has three pairs of principal lateral nerve roots (not count- ing the INI nerve which adheres to the interganglionic connective) arising from the prothoracic ganglion. One nerve, IIN1, appears to arise from the intergan- glionic connective between the prothoracic ganglion and thoracic-abdominal ganglionic mass, and eight principal pairs of nerves arise from the thoracic-ab- dominal ganglionic mass. 2. THORACIC MUSCULATURE General: The thoracic musculature of the male cicada, Tibicen chloromera (Walker) is illustrated in Figs. IB to 8. Figs. 2 to 8 represent stage dissections which proceed from the interior muscle groups to the exterior muscle groups on the body wall. Arabic numerals are utilized for muscle numbers. Thoracic 28 New York Entomological Society | V ol. LXXIV Table 7. Comparison of metathoracic musculature of Tibicen chloromera, Huechys san- guined var. philaemata (Maki, 1938), and Cicada {— Tibicen) plebeia (Berlese, 1909). Muscle groups Tibicen chloromera Huechys sanguined var. philaemata (Maki, 1938) Cicada (= Tibicen) plebeia (Berlese, 1909) Dorsal muscles Median dorsal 65 52 37 Ventral muscles Longitudinal ventral 66 53 68 Tergo-Sterual muscles Anterior tergo-sternal 67 54 XXXVI Posterior tergo-sternal 68 55 XXXVII Tergo-Pleural muscles Tergo-pleural 69 56 - Tergo-pleural 70 - - Pleuro-axillarv 71 57 56 Pleuro-axillary 72 58 - Sterno-Pleural muscles Sterno-pleural 73 59 - Furco-entopleural 74 60 65 Coxal muscles Tergal promotor 75 61 42? Pleural promotor 76 62 48 + 49 Sternal promotor 77 63 - Tergal remotor 78 64 44 Tergal remotor 79 65 43 Coxo-subalar SO 66 XLIX Sternal remotor 81 67 61 Sternal remotor 82 — — Pleural abductor 83 68 - Trochauteral muscles Tergal depressor 84 69 XLV Tergal depressor 85 70 46 Pleural depressor 86 71 - Sternal depressor 87 72 - Muscles of the spiracle Occlusor 88 73 — muscles are listed with their muscle numbers, origins, and insertions in Tables 1 to 4. A comparison of the thoracic musculature of Tibicen chloromera , Huechys sanguinea var. philaemata described by Maki ( 1938) and Cicada (= Tibicen) plebeia described by Berlese (1909) is presented in Tables 5 to 7. Ventral Muscles Which Cover the Thoracic Ganglia: The prothoracic ganglion and the anterior portion of the thoracic-abdominal ganglionic mass are covered dorsally by three pairs of muscles (Fig. IB). The muscle numbers, origins, and insertions of the three muscles groups are described in Table 1. Ventral muscles 1,2, and 3 are mutually joined by zygomatic connections. Mus- cles 1 and 3 are quite sturdy, while muscle 2 is broad at its zygomatic junction and compressed dorsoventrally. Muscle 3 is joined laterally to ventral longi- tudinal muscle 36 for a portion of its length. Prothoracic Musculature: The prothoracic muscles in Tibicen chloromera are fundamentally homologous to Huechys sanguined var. philaemata (Table 5). The lateral dorsal 7, anterior internal tergo-sternals 12 and 14, anterior tergo- pleural 19, and sternal remotor 28 muscles in Tibicen chloromera were not re- ported in Huechys sanguinea var. philaemata . The anterior internal tergo-sternal muscle 10 reported by Maki (1938) and muscle 144 by Berlese (1909) are absent in Tibicen chloromera. This muscle arises on the tergum and attaches to the ventrolateral cervical sclerite. However, the anterior tergo-sternal muscles 12 in Tibicen chloromera , which has its origin 30 New York Entomological Society [Vol. LXXIV on the posterior end of the head and attaches to the ventrolateral cervical sclerite, is probably the homologue. Berlese (1909) did not report the presence of tergo-pleural, sternal promotor, sternal remotor, tergal abductor, pleural abductor or pleural depressor muscles in the prothorax of Cicada { — Tibicen) plebeia. The anterior internal tergo- sternal 14, anterior tergo-pleural 19, and sternal remotor 28 muscles in Tibicen chloromera do not have counterparts in the other two species of cicadas. Mesothoracic Musculature: The mesothorax is the largest division of the thorax and necessarily so, since it contains the very large dorsal longitudinal (34) and oblique dorsal (35) muscles. The dorsal longitudinal muscles serve as depressors of the wings while the oblique dorsal muscles are probably wing ele- vators (Snodgrass, 1927 and 1935). The tergo-pleurals 42, 43, and 44, trochantino-basalar 52, sternal remotor 58, and sternal adductor 59 muscles in Tibicen chloromera have not been reported in Huechys sanguinea var. philaemata by Maki ( 1938) or in Cicada (= Tibicen) plebeia by Berlese (1909). The trochantino-basalar muscle 51 in Tibicen chloromera is present in Cicada {—Tibicen) plebeia (79 + 80) but not in Huechys sanguinea var. philaemata. The pleural abductor of the coxa, Maki’s muscle number 46 in Huechys sanguinea var. philaemata , was not described by Berlese (1909) in Cicada {—Tibicen) plebeia nor is it present in Tibicen chloromera. Berlese (1909) includes median dorsal 69 and spino-furcal ventral 104 muscles in Cicada { — Tibicen) plebeia. Both of the above muscles are not present in the two other species of cicadas (Table 6). Berlese (1909) did not report the presence of pleural-subalar, sternal promotor, sternal remotor, sternal adductor, sternal depressor, or spiracular muscles. Metathoracic Musculature: The metathorax is extremely short, especially dorsally, where the entire notum is reduced to a narrow band behind the scutel- lum of the mesonotum. The metathoracic musculature in Tibicen chloromera is homologous to that of Huechys sanguinea var. philaemata , with the exception of the tergo-pleural muscle 70 and the sternal remotor muscle 82. Berlese (1909) did not report the presence of tergo-pleural, sternal-pleural, sternal promotor, pleural abductor, pleural depressor, sternal depressor, and spiracular muscles. However, all of the above mentioned muscles were reported by Maki (1938) in Huechys san- guinea var. philaemata and are present in Tibicen chloromera. 3. THE CERVICOTHORACIC NERVOUS SYSTEM General: A dorsal view of the ventral nerve cord in the male of Tibicen chloro- mera (Walker) is shown in Fig. 9. A general description of the thoracic nervous system is presented under the section entitled “The Ventral Nerve Cord.” March, 1966] Vasvary: Morphology of Annual Cicada 31 Fig. 9. Dorsal view of the ventral nerve cord of the male annual cicada Tibicen chloromera (Walker) from the head to the first abdominal segment. 32 New York Entomological Society [Vol. LXXIV The Cervix and the Prothorax: The narrowed membranous region between the head and prothorax of insects is called the cervix or neck and is pre- sumably derived from portions of both the labial and prothoracic segments. Muscles contained within the cervical region are believed to have evolved from both the labial and prothoracic segments. Therefore, the concept that the muscles of a segment are innervated from the ganglion of that segment suggests that each muscle of the cervix can be assigned either to the labial or prothoracic segment by determining the segment of innervation. Three pairs of nerves from the subesophageal ganglion innervate muscles located in the cervical region: SN4 + SN5, SN6 (Fig. 9), and SN7 (Fig. 1A). Nerve SN4 + SN5 innervates the anterior intersegmental muscle 13 and nerve SN6 innervates the anterior intersegmental muscle 16. The SN7 nerve innervates the muscles of the sclerotized rod, mr, and muscles within the labium, mlb (Fig. 1A). The sclerotized rod is an extension of the labium and hangs freely from the cervical membrane. A nerve branch from SN6, designated as SN6a in figs. 1A and 9, may be associated with the innervation of the anterior intersegmental muscle 15 and possibly other muscles in the cervicoprothoracic area. A precise determination could not be made since the SN6a nerve joins with a nerve formed by the coalescence of nerves SN9 and INI. The resulting nerve, INI + SN9 + SN6a, then coalesces with the IN2 nerve to form nerve IN2 + INI +SN9 +SN6a which innervates muscles associated with the cervical sclerites, dorsal muscles, and muscles located in the anterior portion of the prothorax. The SN9 nerves issue from the interganglionic connectives between the subesophageal ganglion and prothoracic ganglion. Nerve branch SN9a inner- vates the internal ventral muscle 9 and external ventral muscle 10 before joining the IIN10 + IIN11 nerve originating from the thoracic-abdominal ganglionic mass (Fig. 9). Nerve SN9 then coalesces with nerve INI and later receives the SN6a nerve before joining nerve IN2 originating from the prothoracic ganglion. The INI nerves arise from the anterior portions of the prothoracic ganglion adjacent to the interganglionic connectives. Nerves INI proceed anteriorly in close association with the interganglionic connectives before coalescing with the SN9 nerves. The IN2 nerves issue from the anterolateral area of the prothoracic ganglion and pass under the internal ventral muscles 9. The first nerve branch, IIN2a, provides two sensory nerve branches to the integument, then passes around the tergal promotor muscle of the coxa 21 and over the anterior basal rim of the prothoracic coxa into the leg. After IN2 coalesces with nerve INI + SN9 + SN6a to produce nerve IN2 + INI + SN9 + SN6a, a nerve branch is formed which combines with nerve branches from nerve IIN10 + IIN11 to innervate the anterior intersegmental muscle 15. Nerve IN2 + INI + SN9 + SN6a then March, 1966] Vasvary: Morphology of Annual Cicada 33 Fig. 10. Dorsal view of the ventral nerve cord of the male annual cicada, Tibicen chloro- mera (Walker), from nerve root IIN3 of the thoracic-abdominal ganglionic mass to the second abdominal segment and showing the innervation pattern of the first abdominal segment. ramifies into three nerve branches. One nerve branch proceeds anteriorly and innervates the anterior intersegmental muscles 11 and 12, the ordinary tergo- pleural muscle 20, and the anterior tergo-pleural muscles 18 and 19. The lateral nerve branch bifurcates into a ventral branch which innervates the pleural abductors 30 and 31 and a dorsal branch which innervates the tergal abductor 34 New York Entomological Society I Vol. LXXIV muscle 29 and the tergal promotor muscles 21 and 22. The remaining nerve branch proceeds dorsally and innervates the anterior intersegmental muscle 14, anterior dorsal muscle 8, median dorsal muscle 5, lateral dorsal muscles 6 and 7, and median dorsal muscle 4. The IN3 nerves arise from the anterolateral portion of the prothoracic gan- glion, pass under the anterior edge of the prothoracic pleural apophysis, and innervate the sternal promotor 23, pleural depressor 33, and sternal remotor 28 muscles. Nerve IN4 arises from the anterolateral portion of the prothoracic ganglion posterior to IN3 and proceeds in a lateral direction passing under the prothoracic pleural apophysis. IN4 provides a nerve branch into the leg before dividing into nerve branches IN4a and IN4b. IN4a is a sensory nerve and provides nerve branches to the posterolateral protergal area. Nerve IN4a innervates tergal remotor muscles 24, 25, 26, and 27 and the tergal depressor muscle 32. Nerve IN5 is very short and issues from the mediodorsal area of the pro- thoracic ganglion and innervates ventral muscle 1. Nerves IN6 and IN 7 arise from the mediodorsal portion of the ganglion pos- terior to nerve IN5 and coalesce with nerve IN8 which arises from the dorsal area of the interganglionic connective between the prothoracic ganglion and thoracic-abdominal ganglionic mass (Fig. 9). It appears that nerves IN6, IN7, and INS are responsible for the innervation of ventral muscle 2 and the occlusor muscle 64 of the mesothorax. The posterior tergo-sternal muscle 17 of the prothorax is innervated by nerve branch IIN2a which arises from the anterolateral area of the thoracic- abdominal ganglionic mass. Nerve IIN2a passes under nerve IINl and proceeds lateral to the longitudinal ventral muscle 36 and along the posterior edge of the prothoracic pleural arm to the posterior tergo-sternal muscle 17. Mesothorax: The mesothorax is the largest thoracic segment in the male of the annual cicada, Tibicen chloromera (Walker). Innervation of the meso- thoracic segment, with the exception of the posterior tergo-sternal muscle 38 and the occlusor muscle 64, is achieved by five pairs of nerves: IINl, IIN2, IIN3, IIN4, and IIN10 + IIN11. The large IINl nerves arise from the short interganglionic connectives between the prothoracic ganglion and thoracic-abdominal ganglionic mass GII (Fig. 9). Nerve IINl passes over nerve IIN2 and provides nerve branch IINl a to the longitudinal ventral muscle 36. Nerve IINla bifurcates into two nerve branches. One nerve branch enters muscle 36 along its mesal surface while the remaining nerve branch enters the lateral surface of the muscle 36. The IIN2 nerve passes laterad to the longi- tudinal ventral muscle 36 and ramifies into six nerve branches along the ventral edge of the median dorsal muscle 34. The large median dorsal muscle 34 is innervated by three nerve branches while the lateral dorsal muscle 35 is in- March, 1966] Vasvary: Morphology of Annual Cicada 35 Fig. 11. Posterolateral view of the nerves and muscles of the right side of the fourth abdominal segment of the male of Tibicen chloromera (Walker). First stage of dissection. 36 New York Entomological Society [Vol. LXXIV nervated by a single nerve branch which passes between the median dorsal muscle 35 and the anterior tergo-sternal muscle 37. The remaining nerve branches (IINlb, IINlc, and IINld) are sensory nerves and terminate in the integument of the mesotergum (Fig. 9). Nerve branch IINlb passes obliquely between the anterior tergo-sternal muscle 37 and the median dorsal muscle 34 and continues nresad to the furco-entopleural muscle 49 and tergal remotor muscle 54. Nerve IINlb provides a nerve branch to the trachea between muscle 54 and 55 before terminating in the integument in the posterior portion of the mesotergum. IINlc passes mesad to median dorsal muscle 34 and divides into two nerve branches. One nerve branch continues dorsally and terminates in the middorsal region of the mesotergum while the other nerve branch passes between the latter nerve and muscle 34 and terminates in the integument in the anterodorsal region of the mesotergum. Nerve IINld passes obliquely over the anterior tergo-sternal muscle 37 then proceeds along the posterior edge of muscle 37 and terminates in the integument of the middorsal region of the mesotergum. Nerve IIN2 arises from the anterolateral surface of the thoracic-abdominal ganglionic mass. Two nerve branches, IIN2a and IIN2b, issue from the base of nerve IIN2. The IIN2 nerve passes under the ventral longitudinal muscle 36, and provides a nerve branch IIN2c to nerve IIN3 (Fig. 9). Nerve IIN2 pro- ceeds anterodorsally along the posterior edge of the profurcal arm and then passes laterad to the anterior tergo-sternal muscle 37. The IIN2 nerve divides into three nerve branches, IIN2d, IIN2e, and IIN2f, prior to entering the forewing. Nerve branch IIN2d enters the integument below the tegula of the mesothorax. Nerve branch IIN2e terminates in the integument in the region of the third axillary sclerites and nerve branch IIN2f enters the mesothoracic tegula. Nerve IIN2 is the anterior wing nerve (AWN, Fig. 9) and enters the base of the mesothoracic wing. Nerve branch IIN2a innervates the posterior tergo-sternal muscle 17 of the prothorax. Nerve branch IIN2b passes under the ventral longitudinal muscle 36, proceeds around the posterior edge of the anterior tergo-sternal muscle 37, and innervates the furco-entopleural muscle 49 and the tergal remotor muscle 54, before terminating in the integument along the lateral edge of the mesotergum. Nerve IIN3 is a large nerve and arises from the lateral surface of the thoracic- abdominal ganglionic mass. The first nerve branch passes under the nerve IIN4 and bifurcates into two nerve branches. One nerve branch enters the integument while the remaining nerve branch passes over the anterior basal aim of the mesothoracic coxa and enters the leg. Nerve branch IIN3a innervates the sternal promotor of the coxa 53, sternal remotor of the coxa 58, sternal adductor of the coxa 59, tergal depressor 61, trochantero-basalar 62, and the tergal promotor of the coxa 50. Nerve IIN3 is then connected to IIN2 by way of nerve branch IIN2c. The next nerve branch, IIN3b, innervates the sterno-basalar muscle 48, March, 1966] Vasvary: Morphology of Annual Cicada 37 the trochantero-basalar muscle 62, the trochantino-basalar muscles 51 and 52, and the coxo-basalar muscle 60. Nerve IIN3 then ramifies into three nerve branches, one innervating the anterior tergo-sternal muscle 37, another which innervates the tergal promotor muscle 50, and a nerve branch designated as IIN3c (Fig. 9). Nerve branch IIN3c innervates the anterior tergo-sternal muscle 37 and the tergo-pleural muscles 39, 40, 42, and 43. Nerve IIN4 issues from the lateral surface of the thoracic-abdominal gangli- onic mass posterior to IIN3 and proceeds posteriorly and ramifies into three nerve branches (Fig. 9). One nerve branch innervates the sternal depressor muscle of the coxa 63 before passing over the posterior basal rim of the coxa and into the mesothoracic leg. Nerve branch IIN4a innervates the furco-ento- pleural muscle 49 and the sternal remotor muscle 57. Nerve IIN4 passes around the posterior edge of the sternal remotor muscle 57 and provides a nerve branch to the coxo-subalar muscle 56. Nerve IIN4 continues dorsally and innervates the tergo-pleural muscles 41 and 44, the pleuro-axillary muscles 45 and 46, and the tergal remotor muscles 54 and 55. The posterior tergo-sternal muscle 38 of the mesothorax is innervated by nerve branch IIN6a'. Nerve branch IIN6a is the posterior wing nerve (PWN, Fig. 9). It is noteworthy that the posterior tergo-sternal muscle 17 of the prothorax is innervated by a nerve branch IIN2a of the anterior wing nerve IIN2. The pleuro-subalar muscle 47 is innervated by a nerve branch formed by the coalescence of IIN10 + IIN11 and nerve branch IIN6a//r. Ventral muscle 3 is innervated by a nerve branch from the IIN10 + IIN11 nerve (Fig. 9). Metathorax: The metathorax is the shortest thoracic segment in the male of the annual cicada, Tibicen chloromera (Walker). The entire notum is reduced to a narrow band behind the scutellum of the mesonotum (Fig. 2). Innervation of the metathoracic segment is achieved by three pairs of nerves: IIN5, IIN6, and IIN1Q + IIN11 (Fig. 9). Nerve IIN5 arises from the lateroposterior surface of the thoracic-abdominal ganglionic mass and passes mesad to the mesofurca. After receiving nerve branch IINlla, nerve IIN5 continues posteriorly and ramifies into five nerve branches (Fig. 9). The anterior nerve branch, IIN5, is the dorsal nerve and innervates the tergal promotor muscle 75, the anterior tergo-sternal muscle 67, and terminates in the median dorsal muscles 65. The next nerve branch origi- nates at the base of IIN5 and bifurcates into a nerve branch which enters the integument and nerve branch IIN5a which innervates the pleural promotor muscle 76 and the pleural abductor muscle 83. A nerve branch originating between IIN5 and IIN5b provides a nerve to the integument before passing over the anterior basal rim of the metathoracic coxa and into the leg. Nerve 38 New York Entomological Society [Vol. LXXIV IN9 Fig. 12. Posterolateral view of the nerves and muscles of the right side of the fourth abdominal segment of the male of Tibicen chloromera (Walker). Second stage of dissection. March, 1966] Vasvary: Morphology or Annual Cicada 39 branch IIN5b passes under the posterior mesof ureal arm and provides a nerve branch to the pleural depressor muscle 86 before innervating the tergal depressor muscles 84 and 85. The remaining nerve branch innervates the sternal promotor muscle 77, the sternal depressor muscle 87, and the sternal remotor muscle 81. Nerve IIN6 passes under the posterior mesofurcal arm and forms nerve branch IIN6a which passes around the posterior tergo-sternal muscle 38 of the mesothorax providing the nerve branch, IIN6a' to muscle 38. Nerve branch IIN6a" passes around the tergal depressor muscle 84 and proceeds along the posterior edge of the second phragma (2 Ph) and provides a nerve branch to the membranous sac of the abdominal air chamber (MS) before terminating in the integument along the posterolateral edge of the metatergum (Fig. 9). Nerve branch IIN6ar// coalesces with nerve IIN10 + IIN11. Nerve IIN6a is the posterior wing nerve and passes along the anterior edge of the second phragma and enters the base of the metathoracic wing. Nerve IIN6 passes under the posterior metafurcal arm and proceeds dorsally along the posterior edge of the tergal remotor muscle 79. Nerve IIN6 provides a nerve branch which enters the integument of the epimeron before innervating the following muscles: coxo- subalar 80, posterior tergo-sternal 68, tergal remotors 78 and 79, tergo-pleural 69 and 70, and the pleural-axillary muscles 71 and 72. Nerve IIN6b provides a nerve branch to the metathoracic leg before innervating the sternal remotor muscles 82 and the coxo-subalar muscle 80. Nerve branch IIN6c innervates the furco-entopleural muscle 74. Nerves IIN10 and IIN11 arise from the posterior dorsolateral surface of the thoracic abdominal ganglionic mass and coalesce to form nerve IIN10 + IIN11. The anterior branch of nerve I1N10 + IIN11 passes mesad to the ventral muscle 3 and provides a nerve branch to muscle 3 prior to passing mesad to the pro- furcal arm and the posterior tergo-sternal muscle 17 of the prothorax. Nerve IIN10 + IIN11 continues anteriorly passing over the large trachea of the mesothoracic spiracle and provides two nerve branches which coalesce with a nerve branch from IN 2 + INI + SN9 + SN6a that innervates the anterior internal tergo-sternal muscle 15 of the prothorax. A nerve branch from SN9a also joins nerve IIN10 + IIN11. The pattern of axon distribution resulting from this fusion requires histological clarification. However, it appears that the anterior intersegmental muscle 15, which attaches to the anterior dorso- lateral region of the protergum and the ventrolateral cervical sclerite, does, in part, receive its innervation from nerve IIN10 + IIN11. The IIN10 + IIN11 nerve continues anteriorly and passes under the tentorial bridge. Nerve IIN11 provides a short nerve branch, IINlla, that coalesces with nerve IIN5. The posterior branch of IIN10 + IIN11 passes dorsally over the caudal portion of the thoracic-abdominal ganglionic mass and bifurcates into a dorsal branch and ventral branch. The dorsal branch divides forming two 40 New York Entomological Society [Vol. LXXIV nerve branches. One nerve branch passes mesad to IIN6a' and enters the large trachea originating from the metathoracic spiracle. The second nerve branch passes laterad to IIN6a' and coalesces with IIN6a/r/ to innervate the sterno-pleural muscle 7 3, the pleuro-subalar muscle 47 of the meso thorax, and the occlusor muscle 88 of the metathoracic spiracle. The ventral branch, IIN10 + IIN11, continues posteriorly and innervates the longitudinal ventral muscle 66, then forms a loop which proceeds anteriorly and provides a nerve branch which coalesces with nerve IIN7. The IIN10 + IIN11 nerve proceeds in a posterior direction and enters the first abdominal segment which contains the sound mechanism. Further description of the innervation pattern of the IIN10 + IIN11 nerve is presented under the section entitled “The Musculature and Innervation of the Sound Mechanism.” Nerve IIN7 innervates the muscles located in the anterior portion of the first abdominal segment while nerve IIN8 innervates the muscles located in the posterior portion of the first abdominal segment. The IIN9 nerves innervate the remaining pregenital abdominal segments by providing a pair of lateral nerve roots to each consecutive segment. Discussion: The thoracic nervous system in the male of the annual cicada, Tibicen chloromera (Walker), presents a perplexing enigma regarding the determination of nerve homologies. This problem is due to the coalescence of the mesothoracic, metathoracic, and abdominal ganglia into a single ganglionic mass located in the mesothorax. Condensation of the ventral nerve cord has presumably resulted in the coalescence of lateral nerve branches thereby produc- ing apparent variations in the nerve distribution pattern in Tibicen when com- pared to nerve patterns described in other insects. Schmitt (1962) suggests utilization of the dorsal longitudinal muscles as a starting point in establishing nerve homologies. The dorsal longitudinal muscles are innervated by the dorsal nerves of each consecutive segment. Therefore, from a descriptive standpoint, it is usually easy to identify the dorsal nerve as it issues from its ganglion. However, Niiesch (1954) has shown that some of the axons which supply the dorsal longitudinal muscles also originate from the immediately anterior ganglion. The dorsal muscles of the prothorax in Tibicen is innervated by nerve IN2 + INI + SN9 + SN6a. Nerve SN9 may be the anterior ganglionic connective of the dorsal nerve and has adhered to the interganglionic connective. However, it cannot be determined without recourse to histological examination if nerve INI or nerve IN2 is the prothoracic dorsal nerve. The dorsal longitudinal muscles of the mesothorax are innervated by the I INI nerves which arise from the very short interganglionic connectives between the prothoracic ganglion and the thoracic-abdominal ganglion mass. Anterior ganglionic connectives are not visible in Tibicen. Niiesch (1954) has demon- strated in Telea that motor axons from the prothoracic ganglion pass through March, 1966] Vasvary: Morphology of Annual Cicada 41 the anterior ganglionic connectives to the mesothoracic dorsal nerve. Anterior ganglionic connectives are not visible in Chauliodes , as described by Maki (1936), nor in Agulla , as described by Matsuda (1956). However, Schmitt (1962) proposes that if the findings of Niiesch regarding the innervation of the dorsal longitudinal muscles are applicable to other Neopterygota, it is probable that the fibers of the anterior ganglionic connectives are also present in Chauliodes and Agulla , but are incorporated in the interganglionic connectives to their connec- tion with the dorsal nerves. Nerve IIN5 in Tibicen is the dorsal nerve of the metathorax and provides in- nervation to muscles located in the anterior portion of this segment and the dorsal longitudinal muscles. In the Neopterygota, wing nerves may also be useful in establishing nerve homologies. The wing nerve enters the wing cavity and is associated with sensory structures at the base of the wing. In Tibicen there are two wing nerves. The anterior wing nerve IIN2 arises from the anterolateral surface of the thoracic-abdominal ganglionic mass. Two nerve branches arise from the base of the anterior wing nerve (IIN2a and IIN2b) and a third nerve branch IIN2c coalesces with nerve IIN3 (Fig. 9). Nerve IIN2 then continues as a completely independent nerve providing sensory nerve branches to the integument below the tegula, the tegula, and the integument in the region of the third axillary sclerites before entering the wing cavity. Maki (1936) describes a similar condition in Chauliodes , where a separate nerve which he labeled “fourth root” arises from the mesothoracic ganglion and passes directly into the wing. In Dissosteira , Schmitt (1959) found that in addition to innervating the dorsal muscles of the mesothorax, the dorsal nerve also provides an anterior nerve branch which enters the tegmen anteriorly and a posterior branch, entering the same wing posteriorly. In Agulla , Matsuda (1956) also found that a branch of the dorsal nerve enters the wing. It appears that the wing nerves of Tibicen and Chauliodes are homologous to the wing nerves of Dissosteira and Agulla despite their association with dorsal nerves in the latter two insects. The posterior wing nerve in Tibicen arises as a nerve branch, IIN6a, from nerve IIN6 (Fig. 9). Nerve IIN6a provides three nerve branches, IIN6a', IIN6a", IIN6a"', before continuing as a separate nerve and passing directly into the metathoracic wing cavity. It is interesting to note that the thoracic legs receive their innervation from two pairs of nerves, one entering the coxae anteriorly and the other posteriorly. The prothoracic legs are innervated by nerve branches from nerves IN2 and IN4 and the mesothoracic legs by nerve branches from IIN3 and IIN4 while nerve branches from IIN5 and IIN6 enter the metathoracic legs. In Tibicen no median nerves are visible between the subesophageal ganglion and the prothoracic ganglion nor between the prothoracic ganglion and the thoracic-abdominal ganglionic mass. However, the median nerves may be 42 New York Entomological Society [Vol. LXXIV included within the interganglionic connectives. The transverse or lateral nerves to the occlusor muscles of the mesothoracic spiracles arise from the dorsal surface of the prothoracic ganglion. The mesothoracic occlusor muscle 64 is innervated by a nerve resulting from the coalescence of nerves IN6, IN 7, and IN8 (Fig. 9). Case ( 1957) has shown that in the cockroach the axons to muscles of a thoracic spiracle leave the anterior ganglion by way of the median nerve, passing to the transverse nerve and then to the muscles. Hoyle (1959) has reported a similar axon path in the thorax of Schistocerca gregaria. It appears then that the nerve formed by the coalescence of IN6, IN7, and INS is in part the transverse nerve since it terminates in the occlusor muscle of the mesothoracic spiracle. In many instances the transverse nerves from the prothoracic ganglion coalesce with the mesothoracic dorsal nerves. This was found to be true in Agulla and Blattella by Matsuda, in Carausiaus by Marquardt (1939), in Chauliodes by Maki (1936), in Dissosteira by Schmitt (1959), in Periplaneta by Pipa and Cook (1959), in Perla by Wittig ( 1955), and in Telea by Niiesch (1957). Schmitt (1962) mentions that no explanation has been offered for the coalescence of dorsal nerves to the transverse nerves and presumably the transverse nerves provide other functions in addition to exercising control over the spiracles. The IIN10 + IIN11 nerves may contain axons of a transverse nerve, since a nerve branch from IIN10 + IIN11 coalesces with nerve branch IIN6a//r and the resulting nerve terminates in the occlusor muscle 88 of the metathoracic spiracle. 4. THE MUSCULATURE AND INNERVATION OF THE SOUND MECHANISM General: The musculature of the sound mechanism of Tibicen chloromera (Walker) is shown in Figs. 2 to 4 and a list of these muscles with their muscle numbers and attachments is presented in Table 8. Fig. 10 shows that the innervation of the first abdominal segment is achieved by nerves IIN7, IIN8, and IIN10 + IIN11. Nerve branch IIN8a is the auditory or tymbal nerve since it innervates the large tergo-sternal muscle 94 or tymbal muscle of the sound mechanism. Nerve IIN10 + IIN11 provides nerve branches to IIN8a and the tymbal muscle. The Musculature of the Sound Mechanism: The musculature of the sound mechanism of Tibicen chloromera (Walker) is contained within the first ab- dominal segment. The tergo-sternal muscle 94 (Fig. 2), or tymbal muscle, is the largest muscle of the sound mechanism and has its attachments on the basal portion of the first abdominal sternum and a sclerotized terminal plate which attaches to the tymbal by a tendon. The tymbal muscles and the tymbals are the essential elements of the sound-producing apparatus (Myers, 1928). The first abdominal sternite has become modified into a sclerotized V-shaped structure which provides attachments and support for the large tymbal muscles. March, 1966] Vasvary: Morphology of Annual Cicada 43 dim Fig. 13. Diagram of the nerve pattern of the right side of the fourth abdominal segment of the male of Tibicen chloromera (Walker) viewed mesally. Carlet (1876), Vogel (1923), and Myers (1928) have established that this sclerotized V-shaped structure is a modification of the first abdominal sternum. A sternal canal is present within the base of this structure and provides a passageway for two pairs of nerves, IIN8 and IIN9. The “wings” or “arms” 44 New York Entomological Society I Vol. LXXIV Table 8. Musculature of the sound mechanism of Tibicen chloromera (Walker). Muscle Muscle Origin number (or attachment) Insertion (or attachment) Dorsal muscle 89 Anterior intersegmental fold Tergal ridge Dorsal muscle 90 Tergal ridge Posterior intersegmental fold Ventral muscle 91 Metafurca Anterior edge of first abdominal sternite Ventral muscle 92 Metafurca Lateral apodemal arm of anterior edge of sternum Ventral muscle 93 Posterior edge of first ab- dominal sternum Posterior intersegmental fold Tergo-sternal muscle (Tvmbal muscles) 94 Basal portion of first ab- dominal sternum Terminal plate which attaches to tymbal by a tendon Tergo-sternal muscle 95 Anterolateral edge of ter- gum Lateral apodemal arm of ante- rior edge of sternum Tergo-sternal muscle 96 Anterolateral edge of ter- gum ventrad to 95 Lateral apodemal arm of ante- rior edge of sternum Tergo-sternal muscle 97 Anterolateral edge of ter- gum ventrad to 96 Lateral apodemal arm of ante- rior edge of sternum Tergo-sternal muscle 98 Anteroventral edge of tym- bal Lateral apodemal arm of ante- rior edge of sternum Tergo-sternal muscle 99 Tergum along lateral edge of mirror Lateral apodemal arm of poste- rior edge of sternum Tergo-sternal muscle 100 Posterior intersegmental fold Lateral apodemal arm of poste- rior edge of sternum Occlusor of spiracle 101 Lateral apodemal arm of posterior edge of sternum Ventral end of spiracle Occlusor of spiracle 102 Lateral apodemal arm of posterior edge of sternum Base of spiracle of the sclerotized V-shaped structure attach to the tergal ridge of the first abdominal segment. A comparison of the musculature of the sound mechanism of Tibicen chloro- mera with Huechys sanguinea var. philaemata described by Maki (1938) and Cicada {— Tibicen ) plebeia described by Berlese (1909) is presented in Table 9. The musculature of Tibicen differs from that found in the two other species of cicadas compared in Table 9 by the presence of tergo-sternal muscles 96, 97, and 98. Maki’s ventral muscle 76 in Huechys was not described by Berlese (1909) in Cicada (= Tibicen) plebeia ; however, it is present in Tibicen chloro- mera. In Tibicen , the dorsal muscles 89 and 90 have attachments on the tergal ridge (tr) of the first abdominal segment (Fig. 2). Berlese (1909) shows in his figure 542 similar points of attachments for the dorsal muscles 37 and 28-29 in Cicada (= Tibicen) plebeia. However, the dorsal muscle 74 of Huechys san- guinea var. philaemata , as shown by Maki (1938) in his figure 24, has its attachment on the anterior and posterior intersegmental folds of the first abdominal segment. Innervation of the Sound Mechanism: The innervation of the sound mech- anism is shown in Fig. 10. Nerves IIN7, IIN8, and IIN10 + IIN11 provide innervation to the muscles of the first abdominal segment which contains the March, 19661 Vasvary: Morphology or Annual Cicada 45 sound mechanism. Nerve IIN7 issues from the posterior portion of the thoracic- abdominal ganglionic mass and proceeds posteriorly and passes over the meso- furca. Nerve IIN7 receives a nerve branch from the IIN10 + IIN11 nerve before forming the nerve branch IIN7a. Branch IIN7a provides innervation to the ventral muscle 92, the membranous sac of the abdominal air chamber, and occlusor muscle of the first abdominal spiracle 101 before coalescing with the dorsal nerve IIN7. Nerve IIN7 is the anterior dorsal nerve of the first abdomi- nal segment since it terminates in the dorsal muscles 89. Nerve IIN7 provides a nerve branch to the ventral muscle 91 and passes under muscle 91, around the posterior arm of the metafurca, and continues along the anterior edge of the first abdominal segment. Nerve IIN7 provides a sensory nerve branch to the integument and coalesces with nerve IIN10 + IIN11 before innervating the tergo-sternal muscles 95, 96, and 97. Nerve IIN7 then provides a nerve branch to the membranous sac surrounding the tymbal muscle 94 and coalesces with nerve branch IIN7a before providing a nerve branch to the tergo-sternal muscle 98 and dorsal muscle 89 (Fig. 10). Nerve IIN8 issues from the posterior portion of the thoracic-abdominal gan- glionic mass and proceeds posteriorly over the mesofurca. Nerve IIN8 then divides into a dorsal nerve branch IIN8a which terminates in the tergo-sternal muscle 94, and a ventral nerve IIN8 which passes between the ventral muscles 91 and enters the sternal canal. Nerve branch IIN8a is the auditory nerve and proceeds dorsally and for a portion of its length adheres to the IIN8a nerve from the opposite side. Nerve branch IIN8a then receives a nerve branch from the IIN10 + IIN11 nerve and proceeds in a dorso-oblique path over the mesal surface of the large tergo-sternal muscle 94. Nerve IIN8a then passes around the dorso-posterior edge of muscle 94 and enters this muscle along its lateral surface. Nerve IIN8 passes through the sternal canal and provides nerve branches to the integument and ventral muscle 93. Nerve IIN8 then passes along the posterior edge of the first abdominal segment and provides nerve branches to the occlusor muscle of the spiracle 102, tergo-sternal muscles 99 and 100, and the dorsal muscle 90. Nerve IIN8 is the posterior dorsal nerve since it terminates in the dorsal muscle 90 located in the posterior portion of the first abdominal segment. The IIN10 + IIN11 nerve enters the first abdominal segment after supplying a nerve branch to nerve IIN7 and proceeds posteriorly in a dorso-oblique path and provides a nerve branch which coalesces with nerve IIN8a. The IIN10 + IIN11 nerve continues dorsolaterally and provides a short nerve branch to the tymbal muscle 94 before it loops in an anterior direction and coalesces with nerve IIN7 (Fig. 10). The IIN10 + IIN11 nerve is the sympathetic nerve of Voskresenskaya and Svidersky (1960), who report that without the innervation of the sympathetic nerve the sound-producing system cannot function normally. Therefore, both the auditory nerve, IIN8a, and the sympathetic unpaired nerve, 46 New York Entomological Society [Vol. LXXIV Table 9. Comparison of the musculature of the sound mechanism of Tibicen chloromera, with Huechys sanguined var. philaemata (Maki, 1938) and Cicada (— Tibicen) plebeia (Berlese, 1909) . Muscles Tibicen chloromera Huechys sanguined var. philaemata (Maki, 1938) Cicada ( = Tibicen ) plebeia (Berlese, 1909) Dorsal muscles 89 37 Dorsal muscles 90 — 28-29 Dorsal muscle — 74 — Ventral muscle 91 75 35 Ventral muscle 92 76 — Ventral muscle 93 — 14 + 15 Tergo-sternal muscle 94 77 XXVI Tergo-sternal muscle 95 78 — Tergo-sternal muscle 96 — — Tergo-sternal muscle 97 — — Tergo-sternal muscle 98 — — Tergo-sternal muscle 99 — XVII Tergo-sternal muscle 100 83 XVII First interpleural muscle — — LIII Occlusor of spiracle 101 79 — Occlusor of spiracle 102 85 - IIN10 + IIN11, are necessary for the rhythmic “singing” of the cicada. It is interesting to note that Hagiwara and Watanabe (1956) concluded that the paired tymbal muscles receive alternate impulses from the ganglion, and this alternate activity of the two tymbals may give a double sound vibration frequency. 5. THE MUSCULATURE AND INNERVATION OF THE FOURTH ABDOMINAL SEGMENT General: The abdominal musculature of adult and larval insects conforms to a simple fundamental pattern which is repeated with only minor variations in each of the pregenital segments (Snodgrass, 1935). The major groups of ab- dominal muscles found in insects are the dorsal muscles, ventral muscles, lateral muscles, transverse muscles, and spiracular muscles. The dorsal and ventral muscles in most insects occur in two layers and thereby form dorsal internal and external muscles and ventral internal and external muscles. In the male of Tibicen chloromera (Walker) the dorsal external and ventral external muscles are absent. The writer observed a similar condition in all of the typical pre- genital abdominal segments. Another common form of diversification affecting dorsal and ventral muscles includes a more or less distinguishable grouping of the muscles into median and lateral sets (Snodgrass, 1935). In Tibicen the dorsal muscles can be classified into dorsal internal median and dorsal internal lateral muscle groups; however, the ventral internal muscles cannot be classified into median and lateral sets since there are no ventral transverse muscles or a wide separation between the ventral internal muscles. March, 1966] Vasvary: Morphology of Annual Cicada 47 The musculature of the fourth abdominal segment of the male of the annual cicada, Tibicen chloromera (Walker) is shown in Figs. 11 and 12 and a list of the muscles with their muscle numbers and attachments is presented in Table 10. The innervation of the abdominal musculature with the exception of the muscles of the first abdominal segment is achieved by nerves IIN9. The IIN9 nerves provide a pair of lateral nerve branches to each consecutive pregenital abdominal segment. Figs. 11 and 12 show the innervation of the fourth abdomi- nal segment. The Musculature of the Fourth Abdominal Segment: The musculature of the fourth abdominal segment of the male of Tibicen chloromera (Walker) can be classified into dorsal, ventral, lateral, and spiracular muscles (Table 10). The dorsal muscles are subdivided into dorsal internal median (103) and dorsal internal lateral (104) muscles, dorsal (105) and ventral (106) muscles of the apodeme, and dorsal transverse muscles (107). The dorsal internal median and dorsal internal lateral muscles have their attachments on the anterior and pos- terior intersegmental folds while the dorsal and ventral muscles of the apodeme have their attachments on the anterior edge of the apodeme and the posterior in- tersegmental fold. It is interesting to note that there is a complete absence of dorsal external muscles in the pregenital abdominal segments. The usual location of the dorsal external muscles is the posterior portion of the abdominal segment with their attachments on the posterior margin of the tergunr and the posterior intersegmental fold. In this position the dorsal external muscles serve as protractors of the abdomen. It appears that the protraction of the abdomen in Tibicen is achieved by the contraction of the dorsal and ventral muscles of the tergal apodeme. The dorsal transverse muscle, 107, has its attachments along the lateral edge of the dorsal vessel and the anterolateral intersegmental fold of the tergum. Eight closely associated sets of ventral internal muscles, 108, are present in Tibicen. The ventral internal muscles cannot be grouped into specific median and lateral muscle sets. Four pairs of lateral muscles are present in Tibicen : lateral internal 109, lateral external 110, lateral intrasegmental 111, and the dilator of the abdomen 112 (Figs. 11 and 12). The lateral internal and external muscles are tergo-sternal muscles and have their attachments on the anterior intersegmental fold of the tergum and the internal surface of the anterior sternal apodeme. The lateral intrasegmental muscle is tergo-sternal in its attachments and is located in the posterior portion of the segment. The dilator of the abdomen is attached to the anterolateral edge of the tergum and the external surface of the anterior sternal apodeme. The spiracle of the fourth abdominal segment is located in the anterolateral corner of the sternum. The occlusor muscle of the spiracle has its attachments 48 New York Entomological Society [Vol. LXXIV on the anterolateral portion of the sternum adjacent to the sternal apodeme and to the ventral edge of the spiracle. Maki (1938) in his figure 24 shows the musculature of the third abdominal segment of Huechys sanguined var. philaemata. The musculature of the fourth abdominal segment of Tibicen chloromera (Walker) is homologous to the third segment of Huechys sanguinea var. philaemata with the exception of the dorsal internal lateral muscles 104, the lateral intrasegmental muscle 111, and the dilator of the abdomen 112 of Tibicen (Table 11). The Innervation of the Fourth Abdominal Segment: The innervation of the fourth abdominal segment is achieved by a pair of lateral nerve branches which arise from the IIN9 nerves. The IIN9 nerves issue from the posterior end of the thoracic-abdominal ganglionic mass and pass over the mesofurca and meta- furca and between the ventral muscles into the sternal canal. After the IIN9 nerves pass through the sternal canal they provide a pair of lateral nerve branches to the second and remaining pregenital abdominal segments. The lateral nerve branch from nerve IIN9 divides into a dorsal nerve and ventral nerve prior to passing under the ventral internal muscles of the fourth abdominal segment. The innervation of the fourth abdominal segment is shown in Figs. 1 1 and 12. The ventral nerve (VNv) passes under the dorsal nerve (DNv) and terminates in the integument beneath the ventral internal muscles 108 (Fig. 12). The dorsal nerve provides a nerve branch to the ventral internal muscles which are innervated along their external surface. The dorsal nerve proceeds laterally and provides a nerve (TNv) to the occlusor muscle of the spiracle 113 (Fig. 13). Case (1957) presents experi- mental evidence that the median and transverse nerves provide a neural pathway connecting the spiracular mechanism with the central nervous system. Schmitt (1965) presents a comparative morphological study on the transverse nerves in the abdominal nervous system of insects and concludes that, in insects which apparently lack median and transverse nerves, these nerves have become in- corporated in the longitudinal connectives and lateral segmental nerves. In the majority of insects reviewed by Schmitt, the transverse nerve also innervates the alary muscles. In Tibicen chloromera (Walker) the writer could not determine the innervation of the alary muscles; however, it appears reasonable to con- clude that the innervation of the occlusor muscle of the spiracle of Tibicen is accomplished by fibers of the transverse nerve which have become incorporated in the dorsal nerve. After providing a nerve branch to the occlusor muscle of the spiracle, the dorsal nerve ramifies into three nerve branches. The anterior nerve branch in- nervates the lateral internal muscle 109, lateral external muscle 110, and the dilator of the abdomen 112. The posterior nerve branch divides into a sensory nerve which terminates in the integument and a nerve branch which innervates the lateral intrasegmental muscle 111 (Figs. 11 and 12). March, 1966] Vasvary: Morphology of Annual Cicada 49 Table 10. The musculature of the fourth abdominal segment of Tibicen chloromera (Walker). Muscle Muscle number Origin (or attachment) Insertion (or attachment) Dorsal muscles Dorsal internal median muscles 103 Anterior intersegmental fold Posterior intersegmental fold Dorsal internal lateral muscles 104 Anterior intersegmental fold Posterior intersegmental fold Dorsal muscle of apodeme 105 Anterior edge of tergal apodeme Posterior intersegmental fold Ventral muscle of apodeme 106 Anterior edge of tergal apodeme Posterior intersegmental fold Dorsal transverse muscle 107 Anterior intersegmental fold Lateral edge of the dorsal vessel Ventral muscles Ventral internal muscles 108 Anterior intersegmental fold Posterior intersegmental fold Lateral muscles Lateral internal muscle 109 Anterior intersegmental fold of tergum Internal surface of sternal apodeme Lateral external muscle 110 Anterior intersegmental fold of tergum Internal surface of sternal apodeme Lateral intrasegmental muscle 111 Posterolateral portion of tergum Lateral edge of sternum Dilator of the abdomen 112 Lateral edge of tergum External surface of sternal apodeme Muscles of the spiracle Occlusor 113 Anterolateral portion of sternum adjacent to the sternal apodeme. Ventral edge of spiracle The dorsal nerve proceeds dorsally in an oblique-posterior direction over the tergal apodeme and supplies a nerve branch to the dorsal and ventral muscles (104 and 105) of the apodeme (Figs. 11 and 12). The dorsal nerve continues dorsally along the posterior portion of the tergum and passes over the dorsal internal lateral muscles 104 and provides nerve branches to these muscles. The dorsal nerve then divides into three nerve branches; two nerve branches pass laterally under the dorsal internal median muscles and terminate in the integu- ment while the dorsal nerve passes mesally over the dorsal internal median muscles supplying these muscles with nerve branches. The dorsal nerve termi- nates in the first set of dorsal internal median muscles (Fig. 11). The segmental nerve pattern in the male cicada, Tibicen chloromera (Walker), is notably abbreviated when compared to the innervation pattern of some families of Orthoptera, as described by Schmitt (1954), Chauliodes jormosanus as described by Maki (1936), the larva and adult of Hyalophora cecropa as described by Libby (1959 and 1961), and in Pteronarchys as described by Schmitt (1963). The abbreviated nerve pattern in Tibicen is largely due to the absence of the dorsal and ventral external muscles and by the condensation 50 New York Entomological Society [Vol. LXXIV Table 11. A comparison of the musculature of the fourth abdominal segment of Tibicen chloromera (Walker) to the musculature of the third abdominal segment of Huechys sanguined var. philaemata described by Maki (1938). Muscle groups Tibicen chloromera Huechys sanguinea var. philaemata (Maki, 1938) Dorsal muscles Dorsal internal median muscles 103 86 Dorsal internal lateral muscles 104 — Dorsal muscle of apodeme 105 S7 Ventral muscle of apodeme 106 88 Dorsal transverse muscle 107 89 Ventral muscles Ventral internal muscles 108 90 Lateral muscles Lateral internal muscle 109 91 Lateral external muscle 110 92 Lateral intrasegmental muscle 111 — Dilator of the abdomen 112 - Muscles of the spiracle Occlusors of the spiracle 113 93 of the ventral nerve cord which has resulted in the formation of a thoracic- abdominal ganglionic mass located in the mesothorax. With the condensation of the ventral nerve cord, the motor axons which supply the innervation to the typical pregenital abdominal segments have become incorporated within a single pair of nerves, IIN9. The IIN9 nerves supply a pair of lateral nerve branches to each consecutive abdominal segment after which there are no nerve connections between segments. The innervation pattern of the fourth abdominal segment of Tibicen is shown in Fig. 13. The dorsal nerve supplies innervation to the dorsal and ventral internal longitudinal muscles which are considered primitive muscle groups of the segmental musculature and are therefore useful in the establishment of a criteria of nerve homology. Schmitt (1954) shows that the dorsal and ventral internal muscles of Dissosteira , Acheta , and Periplaneta are innervated by nerve branches from the dorsal nerve. Libby (1959 and 1961) shows a similar innerva- tion of the same muscle groups in the larva and adult of Hyalophora. Further investigations of insects possessing a thoracic-abdominal ganglionic mass located in the thorax must be conducted before significant comparisons can be made with the segmental nerve pattern of Tibicen. SUMMARY AND CONCLUSIONS The musculature and innervation of the thorax, of the sound mechanism, and of a typical pregenital abdominal segment of the male of the annual cicada, Tibicen chloromera (Walker) are described. The musculature of the thorax and March, 1966] Vasvary: Morphology of Annual Cicada 51 abdominal segments of Tibicen is essentially homologous to the musculature of the male cicada Heuchys sanguined var. philaemata as described by Maki (1938). The ventral nerve cord consists of a subesophageal ganglion, prothoracic ganglion, and a thoracic-abdominal ganglionic mass. There are no ganglia pres- ent in any of the abdominal segments. The abdominal segments are innervated by lateral nerve branches arising from a pair of nerves that originate from the posterior portion of the thoracic-abdominal ganglionic mass located in the meso- thorax. Eight pairs of nerves arise from the subesophageal ganglion and supply innervation to the muscles associated with the feeding apparatus, the salivary glands, the lateral ducts of the salivary glands, and some of the muscles of the cervical area. The prothoracic ganglion and the anterior portion of the thoracic-abdominal ganglionic mass are covered dorsally by ventral muscles. The prothoracic ganglion supplies innervation to some of the muscles of the cervical area and the muscles of the prothorax. The thoracic-abdominal ganglionic mass provides innervation to the posterior tergo-sternal muscles of the prothorax, the muscles of the mesothorax, metathorax, and all the abdominal segments. No median nerves are visible between the subesophageal ganglion, prothoracic ganglion, and the thoracic-abdominal ganglionic mass. However, the median nerves are prob- ably included within the interganglionic connectives. Spiracular muscles of the thoracic segments are innervated by nerves which arise from the dorsolateral area of the prothoracic ganglion and the thoracic-abdominal ganglionic mass. The nerves to the spiracular muscles are apparently the transverse nerves of the “ventral sympathetic nervous system.” The sound mechanism is contained within the first abdominal segment. An invagination of the first abdominal sternite serves as an area for attachment and support for the large tymbal muscles. A sternal canal is located within the sternal invagination and permits the passage of two pairs of nerves. One pair of nerves innervates the muscles in the posterior portion of the first abdominal segment while the remaining pair of nerves provides innervation to the remain- ing abdominal segments. Each typical pregenital abdominal segment is innervated by a pair of lateral nerve branches which arises from a single pair of nerves originating from the posterior end of the thoracic-abdominal ganglionic mass and pass through the sternal canal. There are no nerve connections between the typical pregenital abdominal segments once the lateral nerves enter their respective segments. A single nerve branch from the dorsal nerve innervates the occlusor muscle of the spiracle of the fourth abdominal segment. It appears that the innervation of the occlusor muscle of the spiracle is achieved by fibers of the transverse nerve which have become incorporated in the lateral nerve branches to the abdominal segments. 52 New York Entomological Society [Vol. LXXIV Acknowledgments I wish to express my gratitude to Dr. J. B. Schmitt of the Department of Entomology and Economic Zoology, Rutgers-The State University, for his assistance and guidance in the selection and suggestions for carrying out this morphological study. This paper is a portion of a thesis submitted to the Graduate School of Rutgers-The State University in partial fulfillment of requirements for the degree of Doctor of Philosophy. Literature Cited Beckel, W. E. 1958. The morphology, histology and physiology of the spiracular regu- lating apparatus of Hyalophora cecropia (L.) Proc. Intern. Congr. Entomol. 10th Meeting, Montreal, Que. 1956, 2: 87-115. Berlese, A. 1909. Gli Insetti. Vol. I. Milan. Binet, A. 1894. Contribution a l’etude du systeme nerveux sous intestinal des Insects. Journ. de l’Amat. de la Phys. 30 Ann., Nr. 5: 449-543. Brandt, E. 1878. Vergleichend-anatomische Untersuchungen fiber das Nervensystem der Hemipteren. Horae Soc. Ent. Ross., Tom. 14: 496-505. Carlet, G. 1876. Sur Panatome de l’appareil musical de la Cigale. C. R. 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The nervous system of certain abdominal segments and the innervation of the male reproductive system and genitalia in Hyalophora cecropia. Ann. Ent. Soc. Amer., 54: 887-896. Maki, T. 1936. Studies on the skeletal structure, musculature and nervous system of the alder fly, Chauliodes formosanus Peterson. Mem. Fac. Sci. and Agric., Taihoku Imp. Univ., 16 (3): 117-253. — . 1938. Studies on the thoracic musculature of insects. Mem. Faculty Sci. and Agric., Taihoku Imp. Univ., Formosa, Vol. 24, No. 1. Entomol. No. 10. Malouf, N. S. R. 1933. The skeletal motor mechanism of the “Stink bug”, Nezara viridula L. Bull. Soc. ent Egypte, Cairo, xvi (1932). 161-203. Marquardt, F. 1939. Beitrage zur Anatomie der Muskulatur und peripheren Nerven von Carausius ( Dixippus ) morosns Br. Zool. Jahrb. Anat., 66: 63-128. Matsuda, R. 1956. The comparative morphology of the thorax of two species of insects. Microentomology, 21: 1-63. Myers, J. G. 1928. Morphology of the Cicadidae (Homoptera). Proc. Zool. Soc. London, 25: 365-472, 74 figs. March, 1966] Vasvary: Morphology of Annual Cicada 53 Nijenhuis, E. D., and D. Dresden. 1955. On the topographical anatomy of the nervous system of the mesothoracic leg of the American cockroach ( Periplaneta americana) . Konikl. Ned. Akad. Wetenschap. Proc. Ser. C., 58: 121-136. Nuescil, H. 1954. Segmentierung und Muskelinnervation bei Telea polyphemus Cr. Rev. Suisse Zook, 61 : 420-428. — . 1957. Die Morphologie des Thorax von Telea polyphemus Cr. II. Nervensystem. Zool. Jahrb. Anat., 75: 615-642. Pipa, R. L., and E. F. Cook. 1959. Studies on the hexapod nervous system. I. The peripheral distribution of the thoracic nerves of the adult cockroach, Periplaneta americana. Ann. Entomol. Soc. Am., 52: 695-710. Pringel, J. W. S. 1954. A physiological analysis of Cicada song. Journ. Exp. Biol., 31: 255-560. Schmitt, J. B. 1954. The nervous system of the pregenital abdominal segments of some Orthoptera. Ann. Ent. Soc. Amer., 47: 677-682. . 1959. The cervicothoracic nervous system of a grasshopper. Smithsonian Inst. Pubis. Misc. Collections, 137: 307-329. . 1962. The comparative anatomy of the insect nervous system. Ann. Rev. of Entomol., 7: 137-156. . 1963. The abdominal nervous system in Pteronarcys. Jour. N. Y. Entomol. Soc., 71: 202-217. . 1965. Variations in the transverse nerve in the abdominal nervous system of insects. Jour. N. Y. Entomol. Soc. 73: 144-150. Snodgrass, R. E. 1927. Morphology and mechanism of the insect thorax. Smithsonian Misc. Coll., 80: No. 1, 108 pp., 44 figs. . 1935. Principles of Insect Morphology. McGraw-Hill Book Co., New York, N. Y. Swinton, A. H. 1880. Insect Variety: its propagation, and distribution, treating of odours, dances, colours, and music in all grasshoppers, cicadae and moths. London. Vogel, R. 1923. Uber ein tympanales Sinnesorgan, das mutmalsliche Hororgan der Singzikaden. Zeitschr f. wissensch. Zook, 120: 190-231. Voskresenskaya, A. K., and V. L. Svidersky. 1960. The role of the central and sympa- thetic nervous system in the function of the tymbal muscles of cicadas. Journ. Ins. Physiol., 6: 26-35. Weber, H. 1929. Kopf and Thorax von Psylla Mali Schmids. (Hemiptera-Homoptera) . Z. Morph. Oekol. Tiere, xiv, 59-165. Wittig, G. 1955. Untersuchungen am Thorax von Perla abdominalis Burm. (Larve und Imago). Zook Jahrb. Anat., 74: 491-570. Received for publication September 20, 1965 ABBREVIATIONS USED ON THE FIGURES AT — Anterior tentorial arm AWN — Anterior wing nerve Ba:i — Metathoracic basalare bp — Bristle plate CoeCon — Circumesophageal connective Con — Connective Cv — Cervix cv — Cervical sclerite Cxi — Coxa of the prothoracic leg Cx^ — Coxa of the mesothoracic leg Cx:; — Coxa of the metathoracic leg 54 New York Entomological Society [Vol. LXXIV dil — Dorsal internal lateral muscle dim — Dorsal internal median muscle dlra — Dilator muscle of the abdomen dlSyr — Dilator muscle of the salivary syringe dma — Dorsal muscle of the tergal apodeme DNv — Dorsal nerve Ful — Mesofurca F u3 — Metafurca GI — Prothoracic ganglion GII — Thoracic-abdominal ganglionic mass L— Leg Lb — Labium le — Lateral external muscle li — Lateral internal muscle lis — Lateral intrasegmental muscle LmNv — Labral nerve M — Mirror of the sound mechanism mlb — Muscles of labium mr — Muscles of the rod MS — Membranous sac of the sound mechanism mxb — Maxillary bristle Op — Operculum osp — Occlusor muscle of the spiracle pi — Tymbal muscle plate PlAi — Prothoracic pleural arm PlAs — Mesothoracic pleural arm PlAs — Metathoracic pleural arm pmdb — Protractor muscle of the mandibular bristle PT — Posterior tentorial arm PWN — Posterior wing nerve r — Rod rmdb — Retractor muscle of the mandibular bristle rmxbi — Internal retractor muscle of the maxillary bristle rmxbs — External retractor muscle of the maxillary bristle Sai — Subalare of the mesothorax Sa;! — Subalare of the metathorax SLD — Salivary duct SLGL — Salivary gland SoeGng — Subesophageal ganglion Sp2 — Mesothoracic spiracle Spy — Metathoracic spiracle sp — Fourth abdominal spiracle Ti — Prothoracic tergum To — Mesothoracic tergum Ty — Metathoracic tergum tapd — Tergal apodeme TB — Tentorial bridge tg — Tegula tn — Tendon between tymbal plate and tymbal TNv — Transverse nerve March, 1966 | Vogel: Spiders from Pennsylvania 55 Tr — Trachea tr — Tergal ridge TYM — Tymbal Vi — Ventral internal muscles Vma — -Ventral muscle of the tergal apodeme Vnv — Ventral nerve wg — Wing of sclerotized V-shaped structure of first abdominal segment lPh — First phragma 2Ph — Second phragma I — First abdominal segment ISp — First abdominal spiracle II — Second abdominal segment lpmxb — Protractor muscle of the maxillary bristle 2pmxb — Protractor muscle of the maxillary bristle Spiders from Powdermill Nature Reserve Beatrice R. Vogel Biology Department, Yale University Abstract: This paper is a list of 150 species of spiders collected during a 2 -week study of the fauna of Powdermill Nature Reserve in Pennsylvania. Local and regional faunal lists are the backbone of ecological and zoogeograph- ical studies. Such lists provide the detailed information on local faunas necessary for syntheses of a broader scope. The spider fauna of Pennsylvania has been sadly neglected. Distribution maps in recent taxonomic revisions almost invari- ably show a lack of locality records from this state. There has been only one list of Pennsylvania spiders published during the last half century (Truman, 1942): a list of spiders from Presque Isle, Erie County. This present paper is also a local faunal list. The Powdermill Nature Reserve of Carnegie Museum is an area of about 1,500 acres in the Ligonier Valley of Westmoreland County, Pennsylvania. The reserve includes a variety of woodland and open (chiefly old field) habitats. This collection was primarily made during a 2 -week study in June and July, 1965, with a few additional specimens collected at other times of the year. This list must, of necessity, be regarded as preliminary. With the exception of duplicates retained by the author, the specimens are deposited in Carnegie Museum. The collection consists of over 1,000 specimens of mature spiders, representing about 150 species. The list from Presque Isle also includes about 150 species, but the two lists have only half their species in common. These lists, along with scattered reports, bring the published number of Pennsylvania species between 200 and 250. Judging by the fauna of New York state, there should eventually be more than 500 species of Pennsylvania spiders. 56 New York Entomological Society LVol. LXXIV I wish to thank C. J. Goodnight, of Western Michigan University, for identifi- cation of the opilionids; W. Ivie, of the American Museum of Natural History, for identification of the erigonids; and W. J. Gertsch, Curator of Spiders at the American Museum of Natural History, for identification of Clubiona and for help with some of the difficult species. I am also indebted to M. Graham Netting, Director of Carnegie Museum, for making my fieldwork at Powdermill Nature Reserve possible. ORDER ARANEIDA Suborder MY GALOMORPHAE ANTRODIAETIDAE Antrodiaetus unicolor (Hentz) Suborder ARANEAMORPHAE AMAUROBIIDAE Amaurobius bennetti (Blackwall) DICTYNIDAE Dictyna cruciata Emerton Dictyna joliacea (Hentz) Dictyna frondea (Hentz) Dictyna sublata (Hentz) Lathy s foxi Marx ULOBORIDAE Hyptiotes sp. (immature) SEGESTERIIDAE Ariadna bicolor (Hentz) TETRAGNATHIDAE Leucauge venusta (Walckenaer) Mimognatha foxi (McCook) Pachygnatha autumnalis Keyserling T etragnatha elongata Walckenaer T etragnatha straminea Emerton T etragnatha versicolor Walckenaer THERIDIIDAE Achaearanea globosum (Hentz) Achaearanea tepidariorum (Koch) Ancylorrhanis hirsutum (Emerton) Asagena americana Emerton Crustulina altera Gertsch and Archer Ctenium frontata (Banks) Ctenium pumilis (Emerton) Dipoena nigra (Emerton) Enoplognatha tecta (Keyserling) Euryopis funebris (Hentz) Steatoda borealis (Hentz) Teutana triangulosa (Walckenaer) Theridion albidum Banks Theridion differens Emerton Theridion frondeum Emerton Theridion lyricum Walckenaer Theridion murarium Emerton Theridion sexpunctatum Emerton Theridion spirale Emerton Theridula opulent a Walckenaer ERIGONIDAE Ceraticelus bulbosus (Emerton) Ceraticelus fissiceps (Cambridge) Ceraticelus laetabilis (Cambridge) Ceratinopsidis formosa (Banks) Ceratinopsis inter pres (Cambridge) Ceratinopsis nigriceps Emerton Collinsia oxypaederotipus (Crosby) Cornicularia minuatus Emerton Cornicularia vigilax (Blackwall) Eperigone macndat a (Banks) Erigone autumnalis Emerton Grammonota trivittata ? Banks Hypselisthes florens (Cambridge) Maso sarcocuom (Crosby and Bishop) Maso sundevalli (Westring) Origanates rostratus (Emerton) Scylaceus pallida (Emerton) LINYPHIIDAE Bathyphantes albiventris (Banks) Lepthy phantes subalpina (Emerton) Linyphia waldea Chamberlin and Ivie Meioneta fabra (Keyserling) Pity ohy phantes costatus Hentz Tennesseellum formica (Emerton) ARANEIDAE Araneinae Acacesia hamata (Hentz) Acanthepeira stellata (Walckenaer) March, 1966 Vogel: Spiders from Pennsylvania 57 Araneus attestor Petrunkevitch Araneus cornutus Clerck Araneus marmoreus Clerck Araneus solitarius (Emerton) Araneus trifolium (Hentz) Araniella displicata (Hentz) Cyclosa conica (Pallos) Cyclosa turbinata (Walckenaer) East ala anastera (Walckenaer) M angora gibber osa (Hentz) Mastophora bisaccata ? (Emerton) (imma- ture) Neoscona arabesca (Walckenaer) Singa praetensis Emerton Singa sp. aff. variabilis Argiopinae Argiope trifasciata (Forskal) Gea heptagon (Hentz) Gasteracanthinae Micrathena gracilis (Walckenaer) Micrathena sagittata (Walckenaer) Theridiosomatinae Theridiosomma radiosum McCook AGELENIDAE Agelenopsis pennsylvanica (Koch) Cicurina arcuata Keyserling Cicurina brevis (Emerton) Cicurina idahoana Chamberlin Cicurina pallida Keyserling Coras medicinalis (Hentz) Wadotes sp. (immature) HAHNIIDAE Antistea brunnea ? (Emerton) (immature) Hahnia cinerea Emerton OXYOPIDAE Oxyopes salticus (Hentz) PISAURIDAE Dolomedes scriptus Hentz Dolomedes tenebrosus Hentz Dolomedes triton sexpunctatus Hentz Dolomedes urinator Walckenaer Dolomedes vittatus Walckenaer Pisaurina brevipes (Emerton) Pisaurina mira (Walckenaer) Pisaurina mira var. subinflata LYCOSIDAE Arc.tosa virgo (Chamberlin) Lycosa frondicola Emerton Lycosa gulosa Walckenaer Lycosa helluo Walckenaer Lycosa rabida Walckenaer Pardosa distinct a (Blackwall) Pardosa lapidicina Emerton Pardosa milvina Hentz Pardosa moesta Banks Pardosa saxatilis (Hentz) Pirata insularis Emerton Pirata maculatus Emerton Pirata minutus Emerton Pirata montanus Emerton Schizocosa avida (Walckenaer) Schizocosa crassipes (Walckenaer) Schizocosa saltatrix (Hentz) GNAPHOSIDAE Drassyllus fallens Chamberlin Sosticus insularis (Banks) Zelotes duplex Chamberlin ANYPHAENIDAE Any phaenella saltabunda (Hentz) CLUBIONIDAE Castianeira sp. (immature) Chiracanthium inc.lusum (Hentz) Clubiona abboti Koch Clubiona kastoni Gertsch Clubiona obesa Hentz Clubionoides pallens (Hentz) THOMISIDAE Misuminae Misumenoides formosipes (Walckenaer) Misumenops asperatus (Hentz) Misumenops oblongus (Keyserling) X ysticus elegans Keyserling Xysticus ferox (Hentz) X ysticus f rat emus Banks Xysticus funestus (Keyserling) Xysticus triguttatus Keyserling Philodrominae Philodromus placidus Banks Philodromus rufus Walckenaer Thanatus sp. (immature) 58 New York Entomological Society [Vol. LXXIV Tibellus maritimus (Menge) Tibellus oblongus (Walckenaer) SALTICIDAE Evarcha hoyi (Peckham) Habrocestum pulex (Hentz) Habronattus decorus (Balckwall) Hasarius adansoni (Audouin) lcius harti Emerton Maevia incle mens (Walckenaer) Marpissa lineata (Koch) Marpissa undata (DeGeer) Metaphidippus galathea (Walckenaer) Neon nelli Peckham Paraphidippus marginata (Walckenaer) Peckhamia scorpiona (Hentz) Phidippus clams Keyserling Phi dip pus prince ps (Peckham) Phlegra fasciata (Hahn) Salticus scenicus (Linnaeus) Sittacus jloridanus Gertsch and Mulaik Synemosyna formica Hentz Zygoballus bettini Peckham ORDER OPILIONIDA Leiobunum nigropalpi Wood Leiobunum ventricosum Wood Leiobunum verrucosum Wood Literature Cited Included is a brief list of works for studying Pennsylvania spiders. Many of these con- tain bibliographies of more specialized papers. Kaston is probably the most useful single work for identifying species. Bonnet, P. 1945-1959. Bibliographia Araneorum. Toulouse, 1, 2: i-xvi, 1-832; 1-5058. Crosby, C. R., and S. C. Bishop. 1928. Araneae. In A list of the Insects of New York. Cornell Univ. Agr. Exper. Sta., Mem. 101 : 1034-1074. Kaston, B. J. 1948. Spiders of Connecticut. State Geological and Natural History Survey, Bull. 70: 1-874. Levi, H. W. 1957. The spider Genera Enoplognatha , Theridion and Paidisca in America north of Mexico (Araneae:Theridiidae) . Bull. Amer. Mus. Nat. Hist., 112 (1): 1-123. Truman, L. C. 1942. A list of spiders collected in western Pennsylvania. Proc. Penn. Acad. Sci., 16: 25-28. Received for Publication November 29, 1965 Recent Publications The Natural History of Mosquitoes. Marston Bates, Harper and Row, New York, $2.45 (paper) 378 pp., 1965. A Systematic Revision of the Amenidae (Diptera: Calliphoridae) , R. W. Crosskey, Bull. Brit. Mus. (Nat. Hist.), Entomology, 16: 2, about $5.00, 107 pp., 1965. The Culicoides of New York State (Diptera: Ceratopogonidae) , Bull. # 399. Hugo Jamn- back, New York State Museum and Science, $1.00, 154 pp., 24 plates, 1965. Mierolepidoptera of Juan Fernandez Island. J. F. Gates Clarke, Proc. U. S. Nat. Mu- seum 117 No. 3508, Smithsonian Institution, Washington, D. C., 106 pp., 1965. A Revision of the Nodini and A Key to the Genera of Eumolpidae of Africa (Cole- optera: Eumolidae), B. J. Selman, Bull. Brit. Mus. (Nat. Hist.), Entomology, 16, No. 2, about $1.90 (paper), 31 pp., 1965. Review of the Genus Cerceris in America North of Mexico (Hymenoptera: Sphecidae), Herman A. Scullen, Proc. U. S. Nat. Museum, 116, No. 3506, Smithsonian Institution, Washington, D. C., 2121 pp., 1965. Defensive Secretion of a Caterpillar (Papilio). Thomas Eisener, and Yvonne C. Mein- wald, Science, 150, Dec. 1965, pp. 1733-1738, illus. March, 1966] O’Brien: Aedes aegypti 59 Origin and Structural Function of the Basal Cells of the Larval Midgut in the Mosquito, Aedes aegypti Linnaeus1 James F. O’Brien2 Biological Laboratories, Fordham University, Bronx, New York 10458 Abstract: This study of a series of midgut whole mounts of larval and pupal Aedes aegypti shows that basal or regenerative cells first appear as a distinct cell type in the mosquito midgut at about the ninth hour of larval life. These cells seldom take part in forming the epithelial lining of the larval midgut. After their appearance, frequent mitotic divisions occur in the basal cells throughout the larval instars resulting in the presence of a large number of these cells in the prepupal midgut. During metamorphosis in the pupal stage, the basal cells remain to form the epithelial layer of the imaginal midgut. Relatively little is known about the cytological development of the midgut in the mosquito, Aedes aegypti Linnaeus. Christophers’ (1960) description of the Aedes digestive tract indicates that the midgut has received little attention from cytologists. Among the three types of cells comprising the larval midgut of Aedes , the regenerative or basal cells remained somewhat of a mystery as to their origin. Christophers stated that the origin of the basal cells is unknown. Berger (1938) reported finding regenerative cells in the larval midgut of the mosquito Culex pipiens but offered no explanation as to their origin. The fairly constant size, active divisions, and increasingly larger numbers of these cells during the larval stages indicate that they must perform some function other than to replace epithelial cells in the larval midgut. The question regarding the origin of the basal cells as well as the fact that such a large number of these cells is present in the later larval instars indicated the need for further cytological study of the midgut of A. aegypti. While the investigation is principally concerned with the larval midgut, pupal and adult midguts were also studied to determine the origin and structural function of the basal or regenerative cells. MATERIALS AND METHODS The Aedes larvae used in this study were obtained from the colony main- tained in this laboratory (O’Brien, 1965). Beginning about 6 hours after hatch- ing and at intervals of from 1 to 4 hours throughout the larval and pupal stages, the midguts were dissected from the specimens. They were then prepared as whole mounts, stained with the Feulgen reaction and counterstained with Orange G. The dissections, the fixation, and the staining procedure were performed on depression slides to eliminate loss or damage to the tissue (O’Brien, 1965). 1 A portion of the author’s dissertation submitted in partial fulfillment of the require- ments for the degree of Doctor of Philosophy at Fordham University. The author wishes to acknowledge the assistance and encouragement given him during this study by Prof. C. A. Berger, S.J., of the Fordham Biological Laboratory. This work was supported in part by an Educational Assistance Grant from the Arthur J. Schmitt Foundation. -Present address: Regis College, Willowdale, Ontario, Canada. 60 New York Entomological Society I Vol. LXXIV Fig. 1. Photomicrograph of portion of stomach area of 17-hour larva showing three potential regenerative cells (arrows). X 1,290. RESULTS In 6-hour larval midguts, only two cell types are present, the longitudinal and circular rows of muscle cells and larger cells forming the epithelial lining of the midgut. At about the twelfth hour of larval life, growth of the midgut has resulted in an increase in size of the epithelial cells, making the epithelial cells easily distinguishable from the smaller regenerative cells which have ap- peared by this time. Examination of whole mounts of midguts between 6 and 12 hours old shows that at about 9 hours, some of the original midgut epithelial cells are undergoing mitotic division. Such a division gives rise to two cells that are smaller than the neighboring cells. These smaller cells are regenerative cells. Up to this time, the midgut wall is only two cell layers thick, the outer cells being the rows of muscle cells and the inner layer the epithelial cells. The division of some of the initial epithelial cells results in the formation of the smaller cells that lie on the basement membrane, at the bases of the epithelial cells — hence the term “basal” cells. Study of the cells of the midguts obtained from larvae between 6 and 9 hours old reveals the presence of large, uniformly sized epithelial cells resting on the basement membrane. A few of these cells exhibit nuclei that appear to be in early prophase of mitotic division, while the neighboring cells contain normal “resting” nuclei. Since all the cells are of about the same size, the cells appear- ing to be in early prophase must be the potential basal cells (Fig. 1). The origin of the basal cells from epithelial cells was confirmed when mitosis was observed March, 1966] O’Brien: Aedes ciegypti 61 Fig. 2. Photomicrograph of portion of a pouch of gastric ceca of 24-hour larva showing one of the primordial epithelial cells in mitotic prophase. X 1,290. in epithelial cells of the gastric ceca (Fig. 2) where regenerative cells appear at a later stage and in fewer numbers than in the stomach area of the midgut. All divisions of the basal cells are normal mitotic divisions, exhibiting the somatic pairing of homologous chromosomes characteristic of dipteran cells (Figs. 2, 3, 4). After the basal cells appear in the midgut, their number increases rapidly by repeated divisions. These cells lie at the bases of the large primordial epithelial cells which continue to grow larger during the larval instars and never divide after about 24 hours of larval life. By the fourth instar, the regenerative cells form almost a complete layer of cells, intermediate in size between the large primordial epithelial cells and the smaller muscle cells, against the basement membrane. The number of basal cells found in the gastric ceca is considerably smaller than in the stomach area of the midgut. Examination of the pupal midgut shows that the regenerative cells form the new epithelial lining of the imaginal midgut. During the larval instars, few of the basal cells help to form the epithelial layer of the midgut. But, after the onset of pupation, the primordial epithelial cells quickly separate from the basement membrane, are sloughed off into the lumen of the midgut, and begin to disintegrate. The basal cells that have been increasing in number throughout larval life continue their divisions and soon form the epithelial lining of the imaginal midgut. Since the adult midgut contains no structure similar to the pouches of the larval gastric ceca, the basal cells that formed in the region of 62 New York Entomological Society [ Vol. LXXIV Fig. 3. Photomicrograph of portion of gastric ceca of 17-hour larva showing a large epithelial cell in mitotic prophase. X 1,290. the gastric ceca during the larval instars combine with the cells of the cardiac region and those of the anterior portion of the stomach area to form the epithe- lium of the anterior region of the adult midgut. DISCUSSION The results of this study indicate the need for revising some statements based upon earlier findings. Berger ( 1938) reported that the cells comprising the epithelial lining of the larval mosquito midgut (the “primordial” epithelial cells) never undergo mitotic division but rather only increase in size during larval life. The basal cells were thought to function primarily as replacement cells for the worn-out epithelial cells in the larval midgut. But the findings here presented show that the early first-instar midgut contains only muscle cells and primordial epithelial cells and that very few of the basal cells function as replacement cells in the larval instars. Therefore, it seems that some of the primordial epithelial cells in the young larva become potential basal or regenerative cells soon after hatching. Once larval feeding and growth begin, these potential basal cells cease functioning as epithelial cells, undergo mitotic division, and become basal cells. In the process of this transformation, their places in the epithelial layer are taken by the nearby primordial epithelial cells which do not divide, but rather enlarge to fill the space left in the epithelial lining. Thus, some of the primordial epithelial cells do undergo division, but only during the early hours of larval life. The factor determining the time of transformation from primordial epithelial cells to potential basal cells is not known. Since so few of the basal cells in the larval midgut function as replacement March, 1966] O’Brien: Aedes aegypti 63 Fig. 4. Photomicrograph of portion of stomach area of 24-hour larva showing a basal cell in mitotic prophase. X 1,290. cells in the epithelial coat, the main role of these cells must be to form the epithelial lining of the imaginal midgut. Therefore, the formation of the adult midgut does not take place principally in the pupa. Both the muscular coat for the midgut, basically that present in the prepupa (O’Brien, 1965), and the epithelial lining of the imaginal midgut, derived from the basal cells of the larval midgut, have been steadily developing throughout the larval stages. SUMMARY Regenerative or basal cells in the larval midgut of A. aegypti first appear about 9 hours after hatching. The basal cells generally take no part in forming the epithelial lining of the larval midgut. After their appearance in the early larval midgut, the basal cells undergo frequent mitotic divisions, resulting in the presence of a large number of basal cells in the prepupa. Early in the pupal stage, the primordial epithelial cells of the larval midgut are sloughed off into the midgut lumen and the basal cells remain to form the epithelial lining of the imaginal midgut. Literature Cited Berger, C. A. 1938. Multiplication and reduction of somatic chromosome groups as a regular developmental process in the mosquito, Culex pipiens. Pub. 496. Carnegie Institution of Washington. Christophers, S. R. 1960. Aedes aegypti (L.). Cambridge Univ. Press. O’Brien, J. F. 1965. Development of the muscular network of the midgut in the larval stages of the mosquito Aedes aegypti Linnaeus. Jour. N. Y. Ent. Soc. 73(4): 226-231. 64 New York Entomological Society [Vol. LXXIV BOOK REVIEWS TWO BOOKS FOR YOUNG NATURALISTS Monarch Butterflies. Alice L. Hopf. Illustrated by Peter Burchard. Thomas Y. Crowell Company, 1965, 135 pp., price $3.75. Fireflies in Nature and the Laboratory. Lynn and Gray Poole. Illustrated by Christine Sapieha. Thomas Y. Crowell Company, 1965, 149 pp., price $3.95. These two little books are valuable additions to the young naturalist’s library. Mrs. Hopf’s book will have greater appeal, probably, since it is based on personal experiences and trans- mits the author’s enthusiasm for field work and dedication to the Monarch Butterfly. The Poole book brings together a large amount of information about luminescence which might be difficult for the young person to ferret out by himself. It is of broader scope than its title implies. Both books will arouse the reader’s interest in getting outdoors and “swinging a net.” Monarch Butterflies describes the work that has been done in tagging butterflies for the purpose of gathering information on flight and migration and offers practical suggestions for the young collector who would like to cooperate in this scientific study. It discusses the collecting, rearing, and photographing of Monarchs and gives detailed and practical suggestions which are applicable to many other species of insects. In touching briefly upon the studies that have been made to test the Monarch’s protection against predation it shows the reader how he, too, may make observations which might be of value. Fireflies includes a discussion of the vocabulary of luminescence and describes luminous dinoflagellates, annelid worms, molluscs, mycetophilids, bacteria, and fungi. The chapters about the early research on luminous organisms, and the various authors’ accounts of them, may not hold the readers’ interest, but those on recent and current work and on the methods of collecting fireflies, shipping them to scientists, and exchanging them with other young collectors for natural history specimens from their particular area will certainly elicit an enthusiastic response. The black and white illustrations are enchanting. Unfortunately the book is marred by poor editing: scientific names are sometimes italicized, sometimes not; genus names are sometimes capitalized, sometimes not; and the arachnid daddylonglegs is called an insect. The price of these books seems high for text that is scarcely more than magazine article length. But the books are attractive; they are printed in large, clear type; and they seem to have been written not just to give information but to encourage young people to be active. Elsie B. Klots / V tt-* J ■ V -(■-{ ' C; 7 4%;'’^ a ;c. ' f ] :. ) A a A- 1 vy 1 a : , , 1 x , \ 1 "' / ■ ■/ ;• r ^ yy: <■( ’ ^ L ); ‘rx- 1 A A A/ Ai|4:, t'V'4 A-y 4'as A "4 a vV'y! ,) • 4 / 4 ;a 4' Vol. LXXIV JUNE 1966 No. 2 yt KC) j - " | Sr ;)• t > ■ ' f m vv, , y • ■■ I M ' . | ; f?; • y i ; | mmwmMmm !\ 4 . I , Vy " j -A ! T\4 I rr .4-.' . \ '\K„. /,' i || ‘ i m iXS\TV ’Tt'/-; Lwtt^'v i \ ivv- V . u1 - m w ^X 'X' :a r,x&i ;4'^At -fiy^ y 4.444*1 i m -f- 1 /■ * , ■ ' A ■ .,\ A 4T4A . v , 4 - V ■M 14 l/ f 4 ■4 4; ' . A f /A., 7 '■•■ 4 o , 444 ni i- \ ■ ■ i . 44:4 1.4 :if , A • ; ■% ' ' (fj. 4 > , . )' / M r ;-t 4A4 1 v,( \ tl~ ' 1 1 V •■’r (' V 'J ■' ■ Devoted to Entomology in General '4 ' f. 44 3$ 7 1 M) AN- 1 ■ A 'vft \V' A ■ 4\v i . i li 4-4 yyu A A /v>V ,'i '.N I f 44- ..4 tl '4 (,4 4... ' ,/w-4 A\ t : Imm u m i ii ■ \ . \i ii~ . 4 4tl!i 4A r.!ir\ ;< r-W :;v ; 4 7: a ay, 4 v ' r: . a r.Ai-i , 4.- • > /f . -4 4 ,a- • 4 -v '*<•• x-r } ■ y i-,0- o Ay Hy. A. .-X .1 V> 'V P> v,. / .V /. A ) • S$\ \; } .fK.1 .: 44 4a > The New York Entomological Society Organized June 29, 1892- — Incorporated February 25, 1893 / ' ' . ; y ^ ■ Reincorporated February 17, 1943 -r- tj) \W'x The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St., & Central Park W., New York 24, N. Y. \ Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. Members of the Society will please remit their annual dues, payable in January, to the Treasurer. V , ,v 77 7 '7 Officers for the Year 1966 President , Dr. Richard Fredrickson College of the City of New York 10031 Vice President, Dr. Kumar Krishna American Museum of Natural History, New York 10024 £ Secretary, Mrs. Lucy Heineman 115 Central Park West, New York 10023 Assistant Secretary, Mr. Albert Poelzl 230 E. 78th Street, New York 10021 Treasurer , Mr. Raymond Brush American Museum of Natural History, New York 10024 Assistant Treasurer , Mrs. Patricia Vaurie 4- z'-' r - £ American Museum of Natural History, New York 10024 > Trustees A- X ■ y v* /Xj . N\; .. ' y i/7 j- Iv \ ■ " X/ • »‘i(y ✓ / \ 1 Year Term Dr. Alexander B. Klots Dr. John B. Schmitt 2 Year Term Mr. Robert Buckbee 1 \ Dr. Jerome Rozen, Jr. V, '-V -V- S' 7; Mailed June 29, 1966 r The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press Inc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, /Kansas. \ - 7 X Journal of the New York Entomological Society Volume LXXIV June 29, 1966 No. 2 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen Columbia University College of Pharmacy 115 West 68th Street, New York, N. Y. 10023 Associate Editor James Forbes Fordham University, New York, N.Y. 10458 Publication Committee Dr. Pedro Wygodzinsky Dr. Asher Treat Dr. David Miller CONTENTS Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XII Charles P. Alexander 66 Notes on the Biology of Stelis ( Odontostelis ) bilineolata (Spinola), a Parasite of Euglossa cordata (Linnaeus) (Hymenoptera : Apoidea: Megachilidae) Frederick D. Bennett 72 Further Studies on the Internal Anatomy of the Meloidae (Coleoptera) . II. The Digestive and Reproductive Systems of the S.A. Blister Beetle, Picnoseus nitidipennis Fairmaire and Germain (Coleoptera: Meloidae) A. P. Gupta 80 Taxonomic Descriptions of the Immature Stages of the Parasitic Bee Stelis ( Odontostelis ) bilineolata (Spinola) (Hymenoptera: Apoidea: Megachilidae) Jerome G. Rozen, Jr. 84 Mature Larvae of the Old World Bee Genus Panurgus (Hymenoptera: Apoidea) Jerome G. Rozen, Jr. and Barbara L. Rozen 92 Melanism in Connecticut Panthea furcilla (Packard) (Lepidoptera : Noctuidae) Alexander B. Klots 95 An Apparent Association of Mites (Acarina) with the Rock Barnacle Balanus Richard W. Fredrickson 101 Bylaws of the New York Entomological Society Book Reviews Notes — Help for Ailing Caterpillars? Membership of the New York Entomological Society Recent Publications Proceedings Necrology Invitation to Membership 103 109 Alice L. Hopf 111 112 116 117 122 123 66 New York Entomological Society I Vol. LXXIV Undescribetl Species of Crane Flies From the Himalaya Mountains (Diptera: Tipulidae) , XII1 Charles P. Alexander Amherst, Massachusetts Abstract: Six new species of the Eriopterine genera Ormosia and Erioptera are described, including Ormosia ( Oreophila ) licina n. sp., from Kashmir and Kumaon, and Ormosia ( Par ormosia ) atrotibialis n. sp., Ormosia ( Ormosia ) subpulehra n. sp., and 0. (0.) um- bripennis n. sp., from Sikkim: Erioptera ( llisia ) diadexia n. sp. and E. (I.) epicharis n. sp., from Sikkim. Part XI of this series of papers was published in the Journal of the New York Entomological Society, 73: 163-167, 1965. The materials upon which the new species are based were collected by Dr. Fernand Schmid, of Ottawa, to whom 1 express my deepest appreciation for this outstanding series of Asiatic Tipulidae. Doctor Schmid collected insect specimens in India and adjoining countries between 1953 and 1961 as a member of the Swiss Zoological Expedition. His insect collections were restricted to certain groups, where they proved to be of paramount importance in making known the exceedingly rich fauna of the region. A summary of the stations visited, as they pertain to the crane flies, is given in a paper by the writer (Philippine Jour. Sci., 90: 163; 1961), covering the period between 1953 and 1960. Between February and October, 1961, still further collections were made by Doctor Schmid in the Kameng Frontier Division of the North East Frontier Agency (NEFA), Assam. In the crane fly materials several hundred new species were included that have been discussed and presently are being described in a long series of papers that are summarized herewith in order to assist other students of the subject: Philippine Jour. Sci. (chiefly Tipulinae and Limoniini) Ann. and Mag. Nat. Hist. (London) (chiefly Tipulinae and Eriopterini) Proc. Royal Ent. Soc. London (Pediciini) Trans. Royal Ent. Soc. London (Hexatomini; Phyllolabis) Bull. Brooklyn Ent. Soc. (Tanyderidae, Ptychopteridae, Trichoceridae) Jour. N. Y. Ent. Soc. (chiefly Eriopterini) Ent. News (Hexatomini) Trans. Amer. Ent. Soc. (Eriopterini) Ormosia ( Oreophila ) licina n. sp. General coloration of mesonotum light brown, sparsely pruinose, pleura yellow; antennae moderately long; wings yellowed, very restrictedly patterned with pale brown, vein 2nd A sinuous; male hypopygium with the outer dististyle black, coarsely spinulose; lateral margins of gonapophyses produced into two or three acute points. 1 Contribution from the Entomological Laboratory, University of Massachusetts. June, 1966 I Alexander: Himalayan Crane Flies, XII 67 male: Length about 4.8-5 mm; wing 5.8-6 mm; antenna about 1.5-1 .6 mm. female: Length about 5 mm; wing 6 mm. Rostrum light yellow; palpi pale brown. Antennae of male moderately long, scape obscure yellow, the remainder black; flagellar segments oval to long-oval, shorter than their verticils. Head light gray. Thorax light brown, sparsely pruinose, pronotum more yellowed, pretergites clear yellow. Pleura and lateral prescutal borders light yellow. Halteres yellow. Legs with coxae and trochanters yellow; remainder of legs yellowish brown, femora more yellowed basally, tarsi brownish black. Wings yellowed, very restrictedly patterned with pale brown, includ- ing the stigma, cord, outer end of cell 1st M2 , and small spots at Sc2 and origin of Rs. Vena- tion: Sci ending nearly opposite the fork of R2+ s+i, Sc2 far retracted, about opposite one-third to two-fifths Rs ; vein R» close to fork of R2+-m', cell 1st M» elongate, subequal to distal section of M ]+2; vein 2nd A sinuous. One wing of the holotype has cell M2 open by atrophy of the basal section of Ms. Abdominal tergites brown, basal sternites paler. Male hypopygium with the dististyles slightly subterminal, broadly united basally; outer style blackened, relatively short and stout, the outer half with numerous strong spinules; inner style pale, relatively short, the outer margin with a strong lobe before midlength. Phallosome with lateral margins of outer apophyses produced into two or three acute points; aedeagus short, black. holotype 3, Dakwani, Pauri Garhwal, Kumaon, 9,300-11,000 feet, August 5, 1958 (Schmid). Allotopotype, 9, pinned with type. Paratypes, 8 9, Gangrea, Pauri Garhwal, 7,500-10,000 feet, June 12, 1958; 13, Tales, Kashmir, August 13, 1954 (Schmid). Ormosia ( Oreophila ) licina is most similar to O. (O.) hutchinsonae Alex- ander, which differs in the coloration, venation, as the short straight vein 2nd A, and in the structure of the hypopygium, especially the phallosome and the elongate dististyle. Ormosia ( Parormosia ) atrotibialis n. sp. Generally similar and closely allied to Ormosia ( Parormosia ) leucoplagia Alexander, differing in the coloration of the legs in the male. male: Length about 4.5-5 mm; wing 5. 2-5. 6 mm. female: Length about 5 mm; wing 5.8 mm. Antennae light yellow, in cases with the intermediate flagellar segments bicolored, their bases narrowly dark brown, the outer two-thirds to three-fourths yellow. Mesonotal prescutum obscure yellow with a more or less distinct capillary brownish black median line ; scutum brown, scutellum and postnotum darker. Legs black in both sexes, the tips of the femora narrowly yellow, including about the outer tenth of segment, extreme tibial bases more narrowly yellowed. In leucoplagia the tibiae and basitarsi of male yellow, of the female black, as in the present fly. holotype 8, Lachen, Sikkim, 8,900 feet, June 13, 1959 (Schmid). Allotopotype, 9 . Paratopotypes, 48 9; paratypes, 1 8, 2 9 9 , Lachung Sikkim, 8,610 feet, July 2, 1959 (Schmid). Ormosia ( Ormosia ) subpulolira n. sp. Allied to puchra ; general coloration of thorax gray, prescutum with a broad brown 68 New York Entomological Society [Vol. LXXIV central stripe, humeral region yellowed; femora yellow with two subequal broad brownish black rings, the outer one nearly apical ; wings whitened, with conspicuous pale brown clouds; male hypopygium with both dististvles extended into acute blackened points; gona- pophvses appearing as a massive black triangular head, its outer margin with three strong spines. male: Length about 4.5 mm; wing 5.3 mm. Head broken. Pronotal scutum dark brownish gray, scutellum testaceous yellow. Mesonotal prescutum gray, with a broad brown central stripe that is narrowly darker medially ; humeral region, including the pseudosutural foveae, yellow, tuberculate pits very reduced; posterior sclerites of notum dark brown, sparsely pruinose. Pleura dark gray, the yellow setae of the posterior pteropleurite very long. Halteres broken. Legs with coxae dark brown; trochanters obscure yellow; femora yellow, each with two subequal broad brownish black rings that are about equal to the pale base or intervening interspace, the tip narrowly yellow; re- mainder of legs light brown. Wings with the ground color whitened, with conspicuous pale brown clouds chiefly in the outer three-fourths, stigma darker; whitened marginal spots in cells R>, R,, and Ri, less evident in cells R5 and 2nd Mz, larger in cells Ms, Cu and the anals; cells basad of cord more extensively whitened; veins brown, prearcular field and Sc, R, and Cu more yellowed. Venation: R2 at fork of R2+ 3+4; cell 2nd M2 square at base; vein 2nd A strongly sinuous, close to border on outer end. Abdomen, including hypopygium, brownish black. Male hypopygium with apical end of tergite short and broad, the lobes low. Both dististyles extended into acute blackened points. Gonapophysis appearing as a massive blackened triangular head, the basal stem relatively slender, outer margin with three strong spines, with a further series of about four microscopic denticles on lower margin near the stem. holotype a broken S , mounted on microscope slide, Zema, Sikkim, 9,100 feet, June 14, 1959 (Schmid). Ormosia ( Ormosia ) siibpulchra is related to O. (O.) kashrniri Alexander and O. (O.) pulchra (Brunetti), all three species differing among themselves chiefly in important characters of the male hypopygium. Ormosia ( Ormosia ) umbripennis n. sp. General coloration of head and thorax brownish black; palpi, antennae, halteres, and legs black; wings strongly infuscated; Sc2 beyond midlength of Rs, cell 1st M2 shorter than vein Mi, vein 2nd A gently sinuous. female: Length about 6 mm; wing 6.5 mm; antenna about 1.6 mm. Rostrum and palpi black. Antennae black throughout ; flagellar segments long-oval, with dense white setae, the verticils longer than the segments. Head brownish black. Thorax uniformly very dark brown to brownish black, the surface of mesonotum subniti- dous; prescutum and scutellum with a few long setae. Halteres brownish black, base of stem obscure yellow. Legs black. Wings strongly infuscated, especially the prearcular and costal fields and the stigma; veins brown. Venation: ending opposite the oblique R2, Sc2 moderately retracted, about opposite three-fifths the long Rs; R2+ 3+i shorter than basal section of R:,; cell 1st M-> shorter than vein Ah; m-cu at fork of M, perpendicular and slightly sinuous; vein 2nd A gently wavy. Abdomen brown, the outer segments more blackened. Ovipositor with cerci horn yellow, long and slender, gently upcurved to the acute tips. June, 1966] Alexander: Himalayan Crane Flies, XII 69 holotype 9, Namnasa, Sikkim, 10,000 feet, July 1, 1959 (Schmid). The only other generally similar regional species is Ormosia ( Ormosia ) nyc- topoda Alexander, of Pakistan, which similarly has the legs black but with the wings pale and having the venational details distinct. Erioptera ( Ilisia ) diadexia n. sp. Allied to asymmetrical general coloration of thorax gray, the prescutum with two diffuse brown stripes; antennae black; femora brownish yellow, tips brownish black; wings brownish yellow, conspicuously patterned with brown spots and dots, the latter on all veins excepting Sc and Cu ; male hypopygium with the outer dististyle bilobed, inner style broad, yellow, and the tip very obtuse; gonapophyses with the two arms virtually identical, appearing as straight blackened rods, the tip microscopically toothed. male: Length about 5-6.5 mm; wing 5.8-8 mm. Rostrum gray, palpi black. Antennae relatively long, black; flagellar segments long-oval to fusiform, basal segments with long verticils, all with further dense pale setulae. Head brownish gray. Prothorax brownish gray; anterior pretergites obscure yellow. Mesonotal prescutum brownish gray, the interspaces more infuscated to form two diffuse stripes; posterior sclerites of notum brownish gray, central area of scutum narrowly brown. Pleura brownish gray. Halteres with stem yellow, knob weakly infuscated. Legs with coxae brownish gray ; trochanters brownish yellow ; femoral and tibiae brownish yellow, tips brownish black, the tibiae slightly enlarged and darkened beyond bases; basitarsi light brown, remainder of tarsi black. Wings brownish yellow, conspicuously patterned with brown spots and dots, the former including about five costal areas, the second over Sc2, the third largest, over tip of Sci and R2, fourth area at tip of i?i+2; smaller marginal spots at ends of all longitudinal veins; narrow brown seams over cord, m, arculus, and at near midlength of Cu\ ; paler brown spots on all longitudinal veins excepting Sc and Cu, those basad of cord paler; veins yellow in the ground areas, brown in the patterned markings. Venation: R>+ 3+y P - " i ' : T\ Y r y( , * 1 , ’/ ' The New York Entomological Society Y;Ky l 'YY Y ^ - ; ir / yj: J ' vYY Organized June 29, 1892 — Incorporated February 25, 1893 Reincorporated February 17, 1943 % r - rt&'N'' V:w’ ^ \-f{- vr , v The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St., & Central Park W., New York 24, N. Y. Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. Members of the Society will please remit their annual dues, payable in January, to the Treasurer. y V y -■ sir. President, Dr. Richard Fredrickson Officers for the Year 1966 ' ■ ■- -fA College of the City of New York 10031 Vice President, Dr. Kumar Krishna American Museum of Natural History, New York 10024 Secretary, Mrs. Lucy Heineman 115 Central Park West, New York 10023 Assistant Secretary, Mr. Albert Poelzl 230 E. 78th Street, New York 10021 Treasurer, Mr. Raymond Brush American Museum of Natural History, New York 10024 Assistant Treasurer, Mrs. Patricia Vaurie I V. American Museum of Natural History, New York 10024 C? Yx •4/.,- sf 1 Year Term Dr. Alexander B. Klots 2 Year Term Dr. Jerome Rozen, Jr. Trustees * P 0 C:& Dr. John B. Schmitt 0 Mr. Robert Buckbee Mailed September IS, 1966 The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press Inc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas. ii -■ r \ : " V';J. Journal of the New York Entomological Society Volume LXXIV September 15, 1966 No. 3 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen Columbia University College of Pharmacy 115 West 68th Street, New York, N. Y. 10023 Associate Editor James Forbes Fordham University, New York, N.Y. 10458 Publication Committee Dr. Pedro Wygodzinsky Dr. Asher Treat Dr. David Miller CONTENTS David Bruce (1833—1903) and Other Entomological Collectors in Colorado F. Martin Brown 126 Vitamin Synthesis by the Symbionts in the Fat Body of the Cockroach, Peri- planeta americana (L.) Daniel Ludwig and Margaret R. Gallagher 134 Life History Notes on Lagoa laceyi (Barnes and McDunnough) (Lepidoptera : Maegalpygidae) Alexander B. Klots 140 A New Blattisocius (Acarina: Mesostigmata) from Noctuid Moths Asher E. Treat 143 Proceedings 160 Recent Publications 164 126 New York Entomological Society [Vol. LXXIV David Bruce (1833—1903) and Other Entomological Collectors in Colorado* F. Martin Brown Fountain Valley School, Colorado Springs, Colo. Abstract: Brief comments upon the men who collected insects before Bruce: W. S. Wood, Jr., W. J. Howard, Jas. Ridings, A. A. Allen, Lt. MacCauley, and T. L. Mead preface a biographical sketch of David Bruce, well-known as a collector of Colorado insects from 1883 to 1897. The information about Bruce was garnered from letters written by him to Herman Strecker and newspaper articles published at the time of his death in Brockport, New York. He is best known for his cooperation with W. H. Edwards in studies of the life histories of high altitude butterflies. Thirty-five years ago, when my interest in the butterflies of Colorado was aroused, two names were prominent as early collectors of specimens for the students of these insects. These were Theodore Lutrell Mead and David Bruce. Mead had spent the summer of 1871 in the mountains of central Colorado as a quasi-member of the Wheeler Survey party. Later he prepared the text for the portion of Volume 5 of the reports of that Survey that is devoted to Lepidoptera. In 1934 and again in 1956 I published notes about Mead’s work in Colorado, including a fairly detailed itinerary of his travels based upon his collection. In my continuous search for information about the early naturalists who worked in Colorado I failed to discover much that was useful about David Bruce. I did find bits and pieces about other early naturalist-explorers — William S. Wood, Jr. who visited this part of Kansas Territory in 1859 (Brown 1957a) ; Winslow J. Howard, a jeweler-naturalist who followed the mining camps of the west and lived in Denver City and Central City during the early 1860’s (Brown 1957a); James Ridings who was here in the summer of 1865; A. A. Allen in 1871 (Brown 1957b); and Lt. McCauley who performed a reconnaisance of the extreme southwestern portion of the state in 1877 (Brown 1958). There are others about whom I have gathered a few notes, but not enough to say more than that they visited the state. Wood was a youngster when he was commissioned by the Entomological Society of Philadelphia to explore and collect in the Rocky Mountains, and his insects are so-labeled, “Rocky Mts.” It was through examination of his bird- skins and their documentation that I discovered where in the Rocky Mountains he had spent the summer of 1859. He ranged perhaps no more than thirty or thirty-five miles from Denver spending much of his time in the foothills to the west and southwest of the budding city. He collected insects for the Society’s * This study was supported by N.S.F. Grant GS-969. The original paper was presented to the Ghost Town Club of Colorado Springs on 28 January 1966. September, 1966 I Brown: David Bruce 127 cabinet and members and bird-skins for the Academy of Natural Sciences of Philadelphia. Howard had been employed by Tiffany in New York as a jeweler and watch- maker. In 1860 he appeared in Denver. In the Western Mountaineer for July 19 of that year appeared this notice: “Watches and Jewelry — We solicit your special attention to the advertisement of W. J. Howard, Esq., which appears in this issue. Mr. Howard was formerly in the leading establishment in his line on the continent — that of Messers Tiffany & Co., New York City — and we are able to assure our readers from personal knowledge that any work entrusted to him will be skillfully and properly done. He has a rare collection of the natural curiosities of the Rocky Mountains, which will be found very entertaining to those interested in natural science. Give Mr. Howard a call, and if you have any interesting specimens of the mineral wealth of the country, take them with you.” Howard’s place of business was on the east corner of Larimer and F. Streets in Denver. He probably moved to Central City in late 1861. He gave that city as his address in his application for membership in the Entomological Society of Philadelphia in March of 1862. In Central City he established the firm of How- ard and Colony, manufacturing jewelers. Apparently Howard returned to the East in 1865. A note in the Rocky Mountain News of February 25, 1866, stated that he was living in Brooklyn and had married. In the fall of that year, according to the News for October 15, Howard passed through Denver on the way to Montana. Then I lose him until the 1870’s when he was established in Prescott, Arizona. Recently I came across a lead to him in Leadville at a later date, 1879, but as yet have not been able to pin down his activities. Ridings was a member of the Entomological Society of Philadelphia, an Englishman who by vocation was a house builder and cabinet maker. Apparently he was successful. In Cresson’s history of the society this appears: “rapid in- crease . . . made it necessary to procure more convenient and commodious quarters . . . This need was promptly supplied by James Ridings, who generously erected for the sole use of the society, a two story brick building on the northwest corner of 13th and Rodman Streets. . . .” There is no evidence in the treasurer’s accounts that the Society paid anything for the erection of the building or for rent of it. The journey into Colorado was made by stage up the Platte River. Ridings was passenger in one of the fewT coaches that passed through unmolested by the Indians in 1864. By the time that he returned to the East the troops had the Indians in control along the Platte. While in Colorado Ridings’ activities took him west to Empire City and north to Burlington, as Longmont then was called. One result of Ridings’ collecting in Colorado was the first published summary of knowledge of the butterflies of Colorado written by Tryon Reakirt and published in 1865. 128 New York Entomological Society [Vol. LXXIV The first collector to venture deeply into the mountains was Theodore Mead in 1871. At the expense of his family and of his future father-in-law, W. H. Edwards, Mead joined the Colorado party of the Wheeler Survey. His wander- ings carried him west to the Independence Pass area, north through Middle Park and south to Canyon City. The result of his work about doubled our knowledge of the butterflies of Colorado, if not of all of the Rocky Mountain Region. Outside of Colorado, only Constantin Drexler, a taxidermist from the Smith- sonian, had previously done any collecting in southwestern Wyoming; and John Pearceall, a member of the Entomological Society of Philadelphia, who had accompained the Mullen Expedition in the Bitterroot Mountain region, had con- tributed to our knowledge of these insects. The two had spent time in the late 1850’s collecting everything that they could lay their hands upon from minerals and fossils to plants and animals. Lt. Charles McCauley was dispatched in the summer of 1877 to make a survey of the roads in southewestern Colorado. At S. F. Baird’s suggestion that “natural history collections made would be of interest” he sampled the area from Tierra Amarilla to the site of Durango travelling via the old Spanish road. It will be noticed that none of these naturalists spent more than a few months in Colorado. It was not until David Bruce arrived on the scene in 1883 that we find a man who, year after year, searched the state for moths and butterflies. This he did until the turn of the century. It is only in the last few months that I have found any thing about Bruce except that he had collected this or that specimen. In the “Strecker hoard” in Chicago, which I am studying for the Chicago Natural History Museum with National Science Foundation support, I found 103 letters written by Bruce to Herman Strecker. What I retail from now on has been gleaned from those letters and odds and ends that I have picked up elsewhere. Bruce wrote “newsy” letters to Strecker, although he had never met the man. The correspondence between the two started in 1882 and ended in 1897. David Bruce was born in Perth, Scotland, on June 13th, 1833; this he told Strecker in a letter dated January 30, 1883. In it he stated “my family removed to the City of Norwich, Norfolk, England, when I was less than a year old. I have since been knocking about in different parts of the world for 49 years.” He early developed an interest in birds and butterflies and painting. In 1861, he fortuitously met one of the “greats” of English entomology, William C. Hewitson, noted for his beautiful precise illustrations of insects. Hewitson urged Bruce to develope his art and to devote his life to scientific illustration. Bruce did not wholly follow the advice. Later in 1861 Bruce set off for New Zealand. Let me quote him (10iv83) in reply to Strecker’s query about the insects of those islands: “I am sorry to say I never captured any insects except fleas and bedbugs. I have no pleasurable recollections connected with my journey there. I went there first simply because my girl and her people went, but after being in the same vessel September, 1966] Brown: David Bruce 129 with her for three months I came to the conclusion I didn’t want her, so went to Australia where I didn’t stay long for I was anxious to get back.” “My second voyage was just after the death of my first wife. My brother was located in New Zealand. I collected birds only and done a little Agency in fine colors and paperhangings.” Later on there appears another tid-bit linking Bruce to New Zealand. “I perhaps mentioned I had a son in New Zealand. My brother there had seven daughters (his second wife went in for twins) he implored me to send him one of my boys, he would send two girls for it in the way of trade. As my eldest son was willing I sent him but as I had some of my own declined the girls. Well, the luck of the family clung to poor Teddy. The vessel was lost and nothing heard of him for 15 months when he was brought back to London, having been picked up by another ship and been around the world. His passage was renewed without additional expense and he went out without any other adventures.” Bruce married Rachel Marshall at Graves End, England, in 1871 and was in Paris at the time of the Seige. When he arrived in this country I do not yet know. I suspect about 1880. He settled in Brockport, New York, and estab- lished a business in which his sons joined him. Bruce put to work his painting ability and journeyed around western New York painting frescoes for churches, hotels and mansions. He was a good business man, and did not object to doing just straight interior house painting. He was successful enough in this area to be financially free to take annual trips to Colorado to study and collect. The first of these trips was in 1883. His ticket, first class, from Rochester to Denver on the Rochester & Pittsburg and the Burlington cost $50 for the round trip. He was able to stay only 8 days since he was called back early in July to do the interior of the Brockport Episcopal Church. In that short time he prepared 200 bird skins and caught several hundred butterflies and moths. This short first stay was made at Buffalo Creek in the Platte Canyon where he lived with an English family summering there. They were the W. G. Smith family. En route to Colorado Bruce had stayed a short time in Red Cloud, Nebraska. He had been injured falling from a scaffold about a month before his departure from the East and needed to rest en route. He made a similar stop on his hurried trip home. In late July he was again at Buffalo Creek! This time he stayed through August. In the family were young son and daughter who went with Bruce on his collecting trips. He left collecting gear with the teen-agers when he finally returned to the east. During the winter Bruce bought a copy of Mead’s report on the butterflies he had collected in Colorado in 1871. This provided him with information that he had previously lacked and he began making plans to return to our state. In mid-July, 1884, he wrote from Denver, “I returned to Denver yesterday after a sojourn of a couple of weeks in the hills. The season is very backward this year. The roads in the mountains are impassible from deep snows, yet on the whole 130 New York Entomological Society [Vol. LXXIV I don’t think I have much to grumble about with my success in collecting Lepidoptera. My business venture in Colorado is at present at zero or so near a failure that I can hardly hope much of it, in fact everything here is very dull, the mining prospects are poor and nothing goes down with the monied men but the cattle business ... 1 go up to the mines again on Thursday and shall stay probably two or three weeks at a high elevation (10 to 12,000 feet) and shall put in all the time 1 can collecting.” He returned from the high country on July 28. He had been on the summits of the Hayden Mountains at 12,000 feet. Now he planned to work the lower country at about 8,000 feet, to the west of Denver. On the 19th of August, just before he set out for home, he wrote Strecker, “I had the most cursed luck last week imaginable, for I and a friend borrowed a horse and wagon for a few days to go off on an exploring expedition for about 25 miles. On the second day out we drowned the horse and almost ourselves in crossing a stream. Had to walk 7 miles in wet clothes over the most devilish road in an awful storm of thunder, lightning, wind and hail. Had to pay 80 dollars for the horse.” The return address for this letter was “c/o H. Tammen, Rocky Mountain Museum, 454 Larimer Street, Denver, Colo.” During this stay Bruce’s base was “two miles from Denver, right by the foothills.” On this trip Bruce had met with a rancher operating on the Cache la Poudre who was a kindred soul in loving the out-of-doors and collecting specimens. He quoted part of a letter from this cattleman-nimrod, “My friend is one of the best and most fearless hunters living and would run himself nearly to death to catch a good butterfly or shoot a rare bird for me, but he cannot get hold of the names — he tells me he shot a splendid ‘White Pilgrim’ the other day. That is as near as he gets to Pelican. But as long as he lets me have them he can call them what he likes.” During the summer of 1884 when in the high country Bruce made his head- quarters at or near the Whale Mine in Hall Valley. When writing about plans for the next summer he told Strecker “The proprietor of the silver mine there refers in glowing terms to my visit and hopes to see me again early next summer when he will try “and make things pleasant.” I visited Bruce’s cabin above the Whale Mine several times in the 1930’s and caught there many of the species first described from those barren highlands from Bruce’s specimens. On March 16th, 1885, Bruce wrote to Strecker, “My son in Cheyenne has a contract that will oblige him to visit all of the Forts on the Mexican and Cana- dian borders during the next two summers. He has invited me to go with him which I have made up my mind to do as I shall get lots of free riding and liesure to entomoligize. . . . We start in middle April.” Bruce’s wonderful summer was doomed. In a letter of November 10 we read, “This year to me has been an utter blank entomologically and worse than that personally and financially. My son died June 13 of pneumonia. T returned from Cheyenne to find my wife had September, 1966] Brown: David Bruce 131 fallen down the cellar stairs from stumbling on a kitten and hurt herself severely.” Shortly after this Bruce and Strecker had a set-to, as appears usual with all of Strecker ’s correspondents. From here on there no longer are gossipy and newsy letters. Bruce collected in Colorado during 1886 and 1887, principally for W. H. Edwards who quoted Bruce extensively in “Butterflies of North America,” a sumptuous three-volume work. Bruce had been so successful collecting in the high country of Colorado that his material now is found in the principal museums of the world. For Edwards he collected eggs and larvae and between the two of them we know more about high altitude butterflies of Colorado than of any other high country in the world. In 1888 Bruce did not visit the state but stayed at home busy at his decorating business. In 1889 and through the 1890’s Bruce’s letter-drop in Denver was George Eastwood at Taylor’s Free Museum on Larimer Street. He wandered all over the western half of the State. Dr. Alexander Shaw of Denver and with interests in the D & RG Railroad saw to it that Bruce had 1,000-mile passes to carry him about in Colorado and Utah. In return Bruce built “pictures” composed of tropical butterflies mounted behind glass for Shaw. In 1892 Bruce was com- missioned to gather an exhibit of moths and butterflies of the State to be part of the Colorado State exhibit at the Chicago World Fair. This he did, being given free travel and a good salary for his work. The World Fair committee paid half of these costs and Colorado Agricultural and Mechanical College at Fort Collins the other half. One letter written March 26, 1891, gives us a verbal picture of Bruce. Strecker had written to him asking for a photograph. Bruce replied “have not had my ‘picter’ taken in America at all — but soon will- — I am old and grey (56) but very active, eyes and teeth as good as ever — 5 Vh — weigh 200 pounds — fresh ruddy complection yet long grey beard — now you ought to know me when I drop on you as I shall one day.” Bruce never did visit Strecker. During 1892 and 1893 while working at Glenwood Springs Bruce became very much interested in what he believed to be natural hybrids that he was catching. This was a biologically moot point that many naturalists denied occurring. Later, in 1894, his patron W. H. Edwards joined him in this study at Glenwood Springs and the facts were proven conclusively. In connection with these studies Bruce wrote Strecker, who questioned natural hybridism, “I am afraid hybridism is common in Colorado. Whoring is a recognized institution in all mining districts and the insects have taken to it as well as the genus Homo.” Early in 1893 Bruce sold his private collection to the University of Wisconsin at $100 per thousand specimens. This was Bruce’s going price to all comers. The size of the Wisconsin purchase made no difference. Material poured from Brockport to Madison until in late 1895 the University called a halt. They had run out of room in the museum! Two families were yet to be shipped, the 132 New York Entomological Society [Vol. LXXIV smaller Noctuids and all of the Geometrids. I wrote to Dr. Shenefelt at Wisconsin to learn more about this collection and to find out whether or not they had received the rest of it. He had the University archivist, Mr. J. E. Boell, look into the matter. His reply to Dr. Shenefelt was, “We have searched high and low for information on this collection, but the only thing that we could find was that the Regents authorized expenditure of funds for a wire partition up in Science Hall to hold the butterfly collection. We could find nothing in the financial rec- ords that indicated a payment to Bruce for this collection. We have no letters between Bruce and Owen.” From the forgoing it sems probable that Owen him- self was paying for the collection and that it rests, unmarked, among the Owen Collection. The Owen Collection no longer is at the University of Wisconsin. A recent letter from Mr. William Sieker, of Madison, Wisconsin, reads in part “When I came to school here in 1931, Owens Collection was being shipped to the U. S. National Museum. I was hired (at about 50^ an hour) to pin the insects more securely into the boxes. I was pretty green then, and was overwhelmed with the size of his collection. It was big — but lacked labels by the thousands, as Owen, I guess, was not too particular about data. This I gathered from what others have said and what short opportunity I had to observe his collection.” Owen probably used a collection method that was in vogue during the late 19th Century. This was to put all of the data on a general label at the head of each series and none on the specimens themselves. A variant of this was to label the first specimen of a series with a pin-label containing the locality data and follow this specimen with the rest of the series without labels. Once such a collection is disturbed it is hopeless to try to label the specimens correctly. Bruce was now in his sixties. He did not take to the field in 1895 nor in 1896. He did return to Colorado in the following year and joined forces with John T. Mason. This proved unsatisfactory to Bruce in many ways. He did not get on well with Mason in the field and the two men had totally different ideas about how to split the monetary rewards for the work. The Mason Collection is in the Denver Museum of Natural History. I know little of Bruce from this time on until his death on September 24, 1903. Fifty years after that event Mr. A. E. Elwell, well on in his eighties, wrote about Bruce for the Brockport Republic-Democrat of November 25, 1954. This article stresses Bruce’s ability as a taxidermist and artist. From it I gather that the now almost universally used “habitat group” method for exhibiting specimens in Museums was a creation of Bruce, not Ackley. Bruce’s death was a sudden one. The Brockport Republic for October 1, 1903, published, “Soon after entering the yard of Mrs. John Sheplar on the Moscow Road in Hamlin, Thursday afternoon, David Bruce fell to the earth and expired before being found. He was seen to enter the yard and a moment later when the family looked September, 1966] Brown: David Bruce 133 out, Mr. Bruce was discovered on the ground and examination showed that he had died.” Bruce is buried in the Lake View Cemetery in Brockport, N. Y. Literature Cited Brown, F. Martin 1934. “The localities of T. L. Mead’s collection of butterflies from Colorado in 1871.” j. N. Y. Ent. Soc. 42: 155-162. . 1956. “Itineraries of the Wheeler Survey Naturalists, 1871 — Theodore L. Mead.” Lepidopterists’ News, 9: 185-190, map. . 1957a. “Two Early entomological collectors in Colorado.” Ent. News 68: 41-47. . 1957b. “J. A. Allen’s trip to Colorado, etc, in 1871.” Lepidopterists’ News 10: 209-212. . 1958. “The McCauley Expedition to the San Juan region.” J. N. Y. Ent. Soc. 55: 139-146. Through the courtesy of Mrs. Willis Knapp, Chairman of the Brockport Museum Com- mittee, I received typed copies of the obituary for Bruce and of Mr. Elwell's article cited in the text. Received for publication May 23, 1966 134 New York Entomological Society [Vol. LXXIV Vitamin Synthesis by the Symbionts in the Fat Body of the Cockroach, Periplaneta americana (L. ) Daniel Ludwig and Margaret R. Gallagher Department of Biological Sciences, Fordham University Abstract: Determinations were made on the vitamin content of the fat bodies of normal and aposymbiotic cockroaches. Of the 10 vitamins studied (ascorbic, folic, nicotinic and pantathenic acids, biotin, cvanocabalamin, inositol, pyridoxine, riboflavin and thiamine), only 3 (ascorbic, folic and pantathenic acids) were present in considerably larger amounts in the normal fat body. Cultured symbionts were able to synthesize them. The lighter cuticular color, sluggishness and reduced reproductive ability of the aposymbiotic insect may be explained by the absence of these vitamins. Blochmann (1888), working with the cockroach, Blatta orientalis , was prob- ably the first to observe intracellular bacteroids in the fat body of an insect. Glaser (1920, 1930) isolated the organisms, successfully cultured them and classified them as bacteria belonging to the genus Corynebacterium. Trager ( 1952), Peklo (1953), Brooks and Richards (1955a, b and 1956) all agreed that the bacteroids are intracellular symbionts. Wigglesworth (1929) suggested that the role of the symbionts may be the synthesis of vitamins. He thought that the intracellular microorganisms in the fat body of the tsetse fly, Glossina , may synthesize vitamins necessary for growth. Evidence to support this view was given by Fraenkel and Blewett ( 1943a and b), Blewett and Fraenkel (1944), Pant and Fraenkel (1950, 1954) and Keller ( 1950), when they showed that insects with intracellular microorganisms did not, and those without them did, require most of the B vitamins in their diet. In addition to the B vitamins, there is evidence that the symbionts might be responsible for the synthesis of ascorbic acid. Filosa ( 1955) and Cordero (1956) demonstrated that homogenates of the cockroach, Periplaneta americana, can synthesize ascorbic acid using most of the D-sugars as substrates. Lisa (1958) observed that homogenates of the cockroach, Leucophaea maderae , synthesized ascorbic acid from D-mannose, and Pierre ( 1962), that the symbionts present in the fat body of this insect are responsible for this synthesis. Noland, Lilly and Baumann (1949) reported that the symbionts in the fat body of the cockroach, Blatella germanica, are largely responsible for the production of folic acid. The present investigations, which consist of a comparison of the vitamin con- tent of fat bodies of normal and aposymbiotic insects, were undertaken to determine whether vitamins are synthesized by the symbionts of the cockroach, P. americana. September, 1966 1 Ludwig and Gallagher: Vitamin Synthesis in Cockroach 135 Table 1. Methods used for the quantitative determination of vitamins in the fat bodies of normal and aposymbiotic cockroaches. Vitamin Methods of assay Ascorbic acid Spectrophotometric method of Roe and Kuether (1942, 1943) with modi- fications of Lowry, Lopez and Bessey (1945) and by Mills and Roe (1947). Biotin Microbiological method of Pennington, Snell and Williams (1940), modi- fied by the use of Lactobacillus arabinosus as given by Strohecker and Henning ( 1965) . Paper chromatographic method of Radhakrishnamurthy and Sarma (1953). Cyanocobalamin Microbiological method using Lactobacillus leichmanii ATCC 7830, out- lined by Strohecker and Henning (1965). Folic acid Microbiological method of Capps, Hobbs and Fox (1948). Inositol Microbiological method of Stokes, Larsen, Woodward and Foster (1943). Paper chromatographic method of Hough, Jones and Wadman (1948). Nicotinic acid Microbiological method of Snell and Wright (1941). Paper chromatographic method of Kodicek and Reddi (1951). Pantothenic acid Microbiological method of Pennington, Snell and Williams (1940). Pyridoxine Microbiological method of Stokes, Larsen, Woodward and Foster (1943). Paper chromatographic method of Snyder and Wender (1953). Riboflavin Microbiological method of Snell and Strong (1939). Chemical method of Scott, Hill, Norris and Hensen (1946). Thiamine Microbiological method of Sarett and Cheldelin (1944). Chemical method of Hennessy and Cerecedo (1939). MATERIALS AND METHODS The technique employed for rendering the cockroaches aposymbiotic was that of Brooks and Richards ( 1955a), except that they used a 0.1% antibiotic diet and the insects became aposymbiotic in the second generation; whereas in the present experiments, a 10% antibiotic diet was fed and they became aposymbiotic 120 days from the beginning of treatment. The diet consisted of 80% Gaines’ dog pellets, 5% Brewer’s yeast, 5% dextrose and 10% of a mixture of aureo- mycin and terramycin in a 1:1 ratio. The dog pellets were powdered and then mixed with the other ingredients. This food preparation was changed every 5 days to insure the freshness of the antibiotics. Controls were maintained on a diet of Gaines’ dog pellets and water. Sub-groups of insects were cultured on diets which were deficient in the specific vitamin to be tested. Histo- logical sections of the fat body were prepared at the end of 60, 90, 100 and 120 days to determine aposymbiosis. Cultures of symbionts were obtained from the fat body according to the techniques of Begg and Sang (1950) and of Pant, Nayar and Gupta ( 1957). These cultures were maintained in lactose broth at 30° C. 136 New York Entomological Society [Vol. LXXIV Table 2. Amount of different vitamins found in the normal and aposymbiotic fat bodies of the cockroach. Values are given as amount/gram of fat body. Each is an average of 10 determinations. Vitamin Normal Aposymbiotic Micro- , , . , . i Chemical biological ,, j , method method Chromato- graphic method Micro- biological method Chemical method Chromato- grraphic method Ascorbic acid 0.2 mg. 0.03 mg. Biotin 48.0 nifig. 42.0 mMg. 45.0 niMg- 39.0 mMg. Cyanocobalamin 28.0 m/rg. 26.0 mMg. Folic acid 62.0 Mg- 9.6 Mg- Inositol 126.8 Mg. 120.0 Mg- 159.0 Mg- 131.0 Mg. Niacin 408.0 Mg- 305.0 Mg- 528.0 Mg- 420.0 Mg- Pantothenic acid 74.0 Mg- 0.0 Mg- Pvridoxine 67.6 mg. 61.0 mg. 61.0 mg. 62.0 mg. Riboflavin 70.0 Mg- 70.0 Mg- 68.0 Mg- 68.0 Mg- Thiamine 66.0 Mg- 71.0 Mg- 62.0 Mg- 69.0 Mg- All analytical procedures were carried out on homogenates of fat bodies from normal and aposymbiotic nymphs. Five per cent homogenates were made in 0.2 molar phosphate buffer at a pH of 6.8, except for the determinations of riboflavin, nicotinic acid and thiamine, in which cases the fat bodies were homogenized in sterile distilled water. The various methods used to assay each vitamin are given in Table 1. Details of each are given by Gallagher (1962), and descriptions of the various methods for vitamin assays by Strohecker and Henning (1965). OBSERVATIONS Organisms fed an antibiotic diet did not become completely aposymbiotic until 120 days of treatment. One manifestation of aposymbiosis was a change in the color of the cuticle from mahogany to a light tan. This change began approximately 80 days after the insect was placed on antibiotics. They also appeared less active, demonstrated a slower response on exposure to light and less speed in avoiding capture as compared to normal insects. They were also of smaller size and molted less frequently than normal insects. The results of the vitamin assays are summarized in Table 2. The table shows that in all cases there is a close agreement in the results obtained by different methods for each of the vitamins. Of the 10 vitamins assayed, only 3 were present in smaller amounts in the fat bodies of the aposymbiotic than in those of the normal insect. They are ascorbic, folic and pantothenic acids. It appears that these vitamins are synthesized by the symbionts. Additional experiments, using cultures of isolated symbionts, verified this conclusion. September, 1966] Ludwig and Gallagher: Vitamin Synthesis in Cockroach 137 DISCUSSION The fading of the cuticular color in the aposymbiotic insect may be associated with the absence of ascorbic acid. In the normal insect, melanin is formed from the oxidation of tyrosine by tyrosinases. Ascorbic and pantothenic acids are activators of tyrosinase (Levine, Dann and Marples, 1943). In vertebrates, de- fective tyrosine metabolism can be corrected by the administration of either folic or ascorbic acids (Rodney, Swendseed and Swanson, 1947). If the reactions involving ascorbic, folic and pantothenic acids are similar in insects to those in vertebrates, a deficiency of any or all of them could produce a fading of the cuticular color. The present experiments demonstrate that they are all produced by the symbionts cultured from the fat body and are absent from the fat body of the aposymbiotic cockroaches. Henry (1962) reported that another deficiency of the aposymbiotic cockroach, Blatella germanica , is the inability to synthesize certain amino acids, including tyrosine, from glucose. Thus in insects without symbionts, the substrate from which melanins are formed is also lacking. The decrease in reproductive capacity noted in the aposymbiotic insect may be caused by a deficiency of folic acid. Berger (1944) gave the first cytological evidence of the necessity of this vitamin for cell division when he showed that sulfanilamide, a folic acid antagonist, caused metaphase arrest in onion roots. Hindmarsh (1949) found that this inhibition of mitosis could be reversed with p-aminobenzoic acid, a precursor of folic acid. Goldsmith and Grank (1952) induced sterility in the vinegar fly, Drosophila melanogaster , by inhibiting mitosis in the germ cells with aminopterin, a folic acid antagonist. Mitlin, Butt and Shortino ( 1957) prevented oviposition in the house fly, Musca domestica, by feeding aminopterin. A microscopic examination of the ovaries showed inhibited ovarian growth and the eggs contained much less yolk than those of normal flies. Gersdorff and Mitlin ( 1954) showed that the addition of folic acid to the rearing medium reversed the antagonism of aminopterin in house fly larvae. Literature Cited Begg, M. and J. H. Sang. 1950. A method of collecting and sterilizing large numbers of Drosophila eggs. Science, 112: 11-12. Berger, C. A. 1944. Experimental studies on the cytology of Allium. Torreya, 44: 41. Blewett, M. and G. Fraenkel. 1944. Intracellular symbionts and vitamin requirements in insects. Proc. Roy. Soc. London, 132: 212-222. Blochmann, F. 1888. Uber des regelmassige Vorkommen von bakterienahnlichen Gebilden in den Geweben und Eieren verscheidener Insekten. Zeitschr. f. Biol., 24: 204-224. Brooks, M. A., and A. G. Richards. 1955a. Intracellular symbiosis in cockroaches. I. Production of aposymbiotic cockroaches. Biol. Bulk, 109: 22-39. . 1955b. Intracellular symbiosis in cockroaches. II. Mitotic division of the mycetocytes. Science, 122: 242. — — — . 1956. Intracellular symbiosis in cockroaches. III. Reinfection of aposymbiotic cockroaches with symbionts. J. Exp. Zook, 132: 447-465. 138 New York Entomological Society [Vol. LXXIV Capps, B. F., N. L. Hobbs and S. H. Fox. 1948. A dehydrated experimental medium for the microbiological assay of folic acid. J. Bact., 55: 869-870. Cordero, S. M. 1956. The synthesis of ascorbic acid in the cockroach, Periplaneta americana (Linnaeus). M. S. Dissertation, Fordham University, New York. Filosa, M. 1955. A synthesis of ascorbic acid by nymphs of the cockroach, Periplaneta americana (Linnaeus). M. S. Dissertation, Fordham University, New York. Fraenkel, G. and M. Blewett. 1943a. Vitamins of the B group required by insects. Nature, 151: 703. . 1943b. Intracellular symbionts of insects as a source of vitamins. Nature, 152: 506. Gallagher, M. R. 1962. Vitamin synthesis by the symbionts in the fat body of the cockroach Periplaneta americana. Ph.D. Dissertation, Fordham University, New York. Gersdorff, W. A. and N. Mitlin. 1954. The relative toxicity to house flies of the methyl and ethyl analogs of allethin. J. Econ. Ent., 46: 945-948. Glaser, R. W. 1920. Biological studies on intracellular bacteria. Biol. Bull., 39: 133-144. . 1930. On the isolation, cultivation and classification of the so-called intracellular “symbiont” or “rickettsia” of Periplaneta americana. J. Exp. Med., 51: 59-81. Goldsmith, R. D. and I. Grank. 1952. Sterility in the female fruit fly, Drosophila melanogaster, produced by the feeding of a folic acid antagonist. Amer. J. Physiol., 171: 726-729. Hennessy, D. and L. R. Cerecedo. 1939. The determination of free and phosphorylated thiamine by a modified thiochrome assay. J. Amer. Chem. Soc., 61: 179-183. Henry, S. M. 1962. The significance of microorganisms in the nutrition of insects. Trans. N. Y. Acad. Sci., Ser. II, 24: 676-683. Hindmarsh, M. M. 1949. Effects of sulphanilamide and p-aminobenzoic acid on mitosis. Nature, 163: 610. Hough, L., J. K. N. Jones and VV. H. Wadman. 1948. Applications of paper chroma- tography to the separation of the sugars and their derivatives on a column of powered cellulose. Nature, 162: 448-449. Keller, H. 1950. Die Kultur der intrazellularen Symbioten von Periplaneta orientalis. Zeitschr. f. Naturforsch., 56: 269-273. Kodicek, E. and K. K. Reddi. 1951. Paper chromatographic determination of nicotinic acid and its derivatives. Nature, 168: 475-477. Levine, S. Z., M. Dann and E. Marples. 1943. A defect in the metabolism of tyrosine and phenylalanine in premature infants. III. Demonstration of the irreversible conversion of phenylalanine to tyrosine in human organisms. J. Clin. Invest., 22: 551-552. Lisa, J. D. 1958. Enzymes of the fat body of the cockroach, Leucophaea maderae (Fabricius). Ph.D. Dissertation, Fordham University, New York. Lowry, O. D., H. A. Lopez and O. A. Bf.ssey. 1945. The determination of ascorbic acid in small amounts in blood serum. J. Biol. Chem., 160: 609-612. Mills, M. B. and J. H. Roe. 1947. A critical study of proposed modifications of the Roe and Kuether method for the determination of ascorbic acid, with further contributions to the chemistry of this procedure, j. Biol. Chem., 170: 159-163. Mitlin, N., C. Butt and J. Shortino. 1957. Effect of mitotic poisons on house fly oviposition. Physiol. Zook, 30: 133-136. Noland, J. L., J. H. Lilly and C. A. Baumann. 1949. Vitamin requirements of the cockroach Blatella germanica (L.). Ann. Ent. Soc. Amer,, 42: 154-164. Pant, N. C. and G. Fraenkel. 1950. The function of symbiotic yeasts of two species of insects, Lasioderma serricorne and Stegohium ( Sitodrepa ) paniceum. Science, 112: 498. . 1954. On the functions of intracellular symbionts of Oryzaephilus surinamensis. J. Zool. Soc. India, 6: 191-192. September, 1966] Ludwig and Gallagher: Vitamin Synthesis in Cockroach 139 , J. K. Nayar and P. Gupta. 1957. On the isolation and cultivation of the intra- cellur symbionts of Oryzaephilus surinamensis. Experimentia, 13: 241. Peklo, J. 1953. Microorganisms or mitochondria? Science, 118: 202. Pennington, D., E. E. Snell and R. J. Williams. 1940. Effect of diet on pantothenic acid content of chick tissues, j. Biol. Chem., 135: 212-222. Pierre, L. L. 1962. Synthesis of ascorbic acid by the normal fat body of the cockroach, Leucophaea maderae (F.), and by its symbionts. Nature, 193: 904-905. Radhakrishnamltrthy, R. and P. S. Sarma. 1953. Symposium on chromatography. 11: 10-46. Rodney, G., M. E. Swendseed and A. L. Swanson. 1947. Tyrosine oxidation by livers in rats with sulfasuxidine-induced pteroylglutamine acid deficiency. J. Biol. Chem., 168: 395-396. Roe, J. H. and C. A. Kuether. 1942. A color reaction for dehvdroascorbic acid useful in the determination of vitamin C. Science, 95: 77-84. . 1943. The determination of ascorbic acid in whole blood and urine through the 2, 4-dinitrophenylhydrazine derivative of dehvdroascorbic acid. J. Biol. Chem., 150: 609-613. Sarett, H. P. and V. H. Cheldelin. 1944. Use of Lactobacillus fermentum 36 for thiamine assay. J. Biol. Chem., 155: 153-160. Scott, M. L., F. W. Hill, L. C. Norris and G. F. Hensen. 1946. Clinical determination of riboflavin. J. Biol. Chem., 165: 65—7 1 Snell, E. E. and G. Strong. 1939. Microbiological assay of riboflavin. Indust. Engin. Anal., 11: 346-348. and L. D. Wright. 1941. A microbiological method for the determination of nicotinic acid. J. Biol. Chem., 139: 675-686. Snyder, J. W. and S. H. Wender. 1953. Separation and determination of pyridoxine, pyridoxal and pyridoxamine by paper chromatography. Arch. Biochem. Biophys., 46: 465-469. Stokes, E., A. Larsen, H. G. Woodward and R. Foster. 1943. Neurospora assay for pyridoxine. J. Biol. Chem., 150: 17-24. Strohecker, R. and H. M. Henning. 1965. Vitamin Assay. The Chemical Rubber Co., Cleveland. Trager, W. 1952. Mitochondria or microorganisms? Science, 116: 332. Wiggles worth, V. B. 1929. Digestion in the tse-tse fly. Parasit., 21: 316. Received for publication May 17, 1966 140 New York Entomological Society [Vol. LXXIV Life History Notes on Lagoa laceyi (Barnes & McDunnough) (Lepidoptera: Megalopygidae) Alexander B. Klots* Abstract: Descriptions are given of the egg and larvae. The mature larva is figured. The mature larva is strongly aposematic in coloration. On 24 July 1959 a number of small larvae were collected in Big Canyon, Guadalupe Mts., Eddy Co., New Mexico, feeding on a scrubby oak, probably Quercus gambeli. Big Canyon, just north of the Texas-New Mexico border, runs from extremely arid, creosote bush and mesquite desert up into the timbered interior of the mountain range. The larvae were found at about 5500 ft. elevation in a zone characterized by alligator-barked juniper (J uni perns deppeana ) and the lower fringes of yellow pine ( Pinus scopulorum) . During the summer’s field work they were taken to the Southwest Research Station of the American Museum of Natural History near Portal, Arizona, where they fed freely on Quercus emoryi ; and eventually to Connecticut, where they fed freely on Q. ilici folia and coccinea. By early September they had entered the last instar, and by the end of September had all enclosed themselves in cocoons. Twelve adults (6 3 3 and 6 $ $) emerged 14-29 April 1960. After being bred to one of the males, one of the females laid about 180 eggs, nearly all of which hatched. The larvae of this Fi generation were reared on various species of eastern Quercus, at first by the author and then, while he was out of the country, by Miss Alice Gray of the American Museum of Natural History. Considerable material of various larval instars, cocoons and adults has been preserved and is in the American Museum of Natural History and the United States National Museum. Three 3 S and three 2 $ , one of each with the genitalia dissected, were compared with the type material of Logoa laceyi (Barnes and McDunnough) in the U. S. National Museum by Dr. Don Davis, and later by the author. Both Dr. Davis and the author consider them identifiable as laceyi. However, in the absence of any modern systematic work on the group it would be unwise to say what laceyi (type locality Texas) is — a distinct species or a subspecies or form of something else, especially since neither the genitalia nor the color and pattern show clear-cut distinguishing characters, and adequate material is lacking. At present, therefore, it seems best merely to record the characteristics of this material for the benefit of some future student. * Department of Biology, The City College of New York, and Department of Entomology, American Museum of Natural History. September, 1966] Klots: Lagoa laceyi Notes 141 Fig. 1. Lagoa laceyi (Barnes & McDunnough) mature larva, lateral aspect (bead to left) X 3. eggs Length 2.2-2 .5 mm., width about 1 mm. Bluntly ovoid, somewhat flattened. Laid in rows, with the sides contiguous, and thickly covered with hairs and hair-like scales of the female’s vestiture. Hatching period: 7-9 days. immature larva Vestiture, except in color, as described below for mature larva; almost wholly white, only the urticating setae being brownish and, in penultimate instar, some of the medium length plumose setae being faintly brownish. mature larva (Fig. 1) Length 20-30 mm. Skin creamy to slightly pinkish white. Prothorax greatly expanded cephalad and ventrad, forming a hood enclosing head; largely naked, with a fringe of hairlike setae around cephaloventral margin. Remainder of body with short, inconspicuous hairs arising from small patches around and above leg and proleg bases, and corresponding regions in legless segments; but prominent vestiture arising in tufts from flat, only slightly projecting verrucae. Vestiture of each verruca as follows: centrally a group of short, stiff, sharply pointed, smooth, brownish urticating setae; in a zone around these many very long, finely plumose, delicate hairlike setae; in a zone around these many shorter, stiffer, finely plumose setae. The urticating setae are more or less brownish. The very long plumose setae are white on the meso- and metathorax and abdominal segments 1-7, but brick red on abdominal segments 8-10. The shorter plumose setae are mostly dark to blackish except that on the mesothorax they tend to be paler brown, or even in part whitish. On the mesothorax there are four verrucae on each side. The most ventral, and largest, is just posterior and slightly ventral to the prothoracic spiracle. The other three, somewhat smaller and nearly equal to each other, lie farther caudad on the segment, and extend in a line dorsad. On the metathorax and abdominal segments 1-8 there are only 3 verrucae on each side, forming 3 longitudinal series, subdorsal, supraspiracular and subspiracular ; of 142 New York Entomological Society I Vol. LXXIV these the supraspiracular ones arc the largest. On the 9th abdominal segment on each side the verrucae of the subdorsal and supraspiracular series are like those of these series anterior to them; but the most ventral one is much smaller and only slightly ventrad and considerably posterad of the one above it. The last segment is largely naked dorsally, with a fringe of long, plumose setae around the caudal margin and a tuft above each proleg. In the mature larva many of the very long, white setae of the thorax tend to droop cephalad and ventrad; the more dorsal ones of the anterior abdominal segments stand up almost straight dorsad, forming a conspicuous crest. There is a similar, but less conspicuous middorsal crest on the posterior abdominal segments. The shorter plumose setae vary con- siderably in individuals from a medium brown to almost black; these are most conspicuous laterally, especially those of the subspiracular verrucae. In the immature larvae the long setae show no such arrangement, protruding randomly. cocoon Length 18-22 mm. Parchment-like, formed of brown silk and other secretions, in which are intermingled most of the soft, red, white and black larval setae but few, if any of the urticating ones. Near the anterior end is a dorso-ventrally diagonal, flat, very hard and stiff partition. Anterior to this the cocoon is very thin and delicate, with an especially abundant mass of the larval setae filling the anterior space. During eclosion the pupa pushes against the hard partition and is led by its slant to the surface of the cocoon away from the solid object to which the cocoon is fastened; this corresponds to the ventral surface of the pupa. The edge of the stiff partition here breaks easily away from the wall of the cocoon, forming a subterminal slit through which the pupa emerges for at least the length of its head and thorax. SIGNIFICANCE OF THE LARVAL APPEARANCE It is perfectly possible that the all-white, fluffy appearance of the smaller larvae has a protective function, making them resemble the tangled masses of cottonwood ( Populus ) down that is almost omnipresent in the Southwest at this stage of the larval life, floating thickly in the air and accumulating in masses on nearly everything. The similarity of the larvae to this down was, in fact, noted when they were collected. The mature larvae must be regarded as definitely aposematic, their black, white and red coloration making a distinctive recognition pattern. They are, of course, well protected by their urticating setae. Another point of interest is the similarity to these and other protected megalopygid larvae of the larvae of some of the metalmark butterflies (Rio- dinidae), occurring in the same environments, which also have long, drooping white hairlike setae. The metalmark larvae may benefit from their resemblance to cottonwood down, and may also benefit, as Batesian mimics, from their re- semblance to the megalopygid larvae. The author, in fact, thought that the very small laceyi larvae were metalmarks when he first saw them. The author is greatly indebted to Mr. Bruce Harris of the New Mexico Department of Game and Fish for information and aid about collecting places in the Guadalupe Mts.; to Dr. Don Davis of the U. S. National Museum for comparing specimens with the type of L. laceyi ; and to Miss Alice Gray of the American Museum of Natural History for rearing the Ft generation when the author was unable to do so. Received for publication June 20, 1966 September, 1966] Treat: A New Blattisocius from Noctuid Moths 143 A New Blattisocius (Acarina: Mesostigmata) from Noctuid Moths Asher E. Treat The City University of New York and The American Museum of Natural History Abstract: Blattisocius patagiorum is distinguishable from previously described species by the slender, edentate form of the movable cheliceral digit in nymphs and females. Males possess an accessory organ lateral to each peritreme. Behavior suggests facultative parasitism upon noctuid moths. The six previously known species of the ascid genus Blattisocius have been recorded (Chant, 1963) from a great variety of habitats, including association with insects in stored grains. Evans ( 1958) reported B. dentriticus (Berlese) from the thorax of a noctuid moth, Caradrina morpheus (Hiifn.), taken in Darlington, Yorkshire, England, but he gave no details regarding its relationship to the host. The species here described has been found on several noctuids under circumstances that provided an unusual opportunity for detailed observations on certain aspects of its behavior and reproduction. Genus Blattisocius Keegan, 1944 Blattisocius patagiorum n. sp. This species differs from others of the genus in the slender, wholly edentate form of the movable digits of the chelicerae in nymphs and females. The peritremes of the female are a little longer than those of B. keegani Fox, but shorter that those of B. tar salis (Berlese). The length of the fixed cheliceral digits is also intermediate as between these species, being longer than that of B. tarsalis, but shorter than that of B. keegani. female In the six specimens at hand, the length of the dorsal shield varies from 530 to 570 u, averaging 546. Its variation in width is from 258 to 280, with an average of 267 /x. It is lightly reticulated in all areas, and bears 33 pairs of setae. The average length of seta ]6, which is typical of the dorsocentral series, is 48 ix. Setae J4, J5, and Z5 are very finely serrate; the others are simple. Dorsally, the soft integument bears 19 pairs of setae (Fig. la). The tritosternum (Fig. 2a) is about 72 fx long and is undivided in its basal three fourths; the laciniae are finely plumose. The sternal shield is lightly reticulate. The fourth sternal setae are on the membrane. Anteromedial to them are minute metasternal plates bearing pores. The genital shield is about as wide as the sternal shield. Its side margins are concave and its rear margin truncate. The genital setae are on the edges of the shield, the paragenital “pores” in small plates at its sides. There are two pairs of elongate metapodal plates. The ventrianal shield is roughly rectangular and lightly reticulate. It bears three pairs of preanal setae. Five pairs of setae are based upon the soft ventral integument. The peritremes extend to about the middle of coxae III. The peritremal shields are broadly joined to the exopodal plates embracing coxae IV. There are prominent expodal plates flanking coxae II and III, but endopodal plates are lacking. The spermathecae are as shown in Fig. 2b. The tectum or epistome is smooth and convex anteriorly (Fig. lb). The movable digits of the chelicerae taper smoothly to their pointed tips, and are without teeth (Fig. 2c). They average 33 /x in length. The fixed digits are about three fifths this length, smooth, and provided with a pilus dentilis. The corniculi (Fig. 2d) are slender and approximated. 144 New York Entomological Society [Vol. LXXIV b Fig. 1. Blattisocius patagiorum n. sp.; a, dorsal surface of idiosoma of holotype female; b, epistome (tectum), showing variation in form. Deutosternal denticles form a narrow series of seven “rows,” with a single denticle in each except the sixth, which may have two. The palpi are normal for the genus. Average leg lengths in microns are: I, 528; II, 422; III, 412; IV, 535. Setation conforms to that given for the genus by Lindquist and Evans (1965). Macrosetae are not present. September, 1966] Treat: A New Blattisocius from Noctuid Moths 145 The legs turn somewhat brown with age, but do not become so conspicuously tanned as in B. tarsalis. male In the five specimens studied the dorsal shield varies in length from 408 to 452 /x, and in width from 22 7 to 250. Average length and width are 430 and 236 ix. The reticular pattern resembles that of the female. There are 33 or 34 pairs of setae on the shield and 11 or 12 pairs on the soft dorsal integument. Setae J 3 , J4, J5, and Z5 are very slightly serrate (Fig. 3a). The tritosternum (Fig. 3b) is about 60 /x long, and is divided for about half its length. The sternogenital shield is elongate, with lateral projections anterior and posterior to coxae II. It bears four pairs of setae and is flanked by the genital pair near its posterior end. The ventrianal shield is broadly triangular and lightly reticulate. In some specimens it bears five, in others six pairs of preanal setae. The exopodal and peritremal shields are similar to those of the female, but dorsolateral and slightly anterior to each peritreme is a structure which I shall refer to as an accessory organ (Fig. 3c). In the most favorably oriented specimen, this appears to lie beneath or within a cuticular fold or pouch (Fig. 4a). The accessory organ is surrounded by a broadly oval plate with tapering anterior and posterior extensions that run parallel to and may join the peritremal plate toward their extremities. Enclosed by this plate is a cigar-shaped, transparent tube or trough with fine transverse ridges or folds projecting into its interior from its median border (Fig. 4b). It is about equal to the peritreme in length and width. Such a structure was men- tioned and figured by Oudemans (1929) and is figured, from Oudemans’ Plate 104, by Nesbitt (1951) in his drawing of the male of B. tarsalis (as tineivorus Oud.). Oudemans compares it to a piece of a peritreme, and says that he has never seen anything like it. Keegan (1944) also figured this part of the organ in his description of B. tarsalis (as trio dons) , but mentioned it no further than to say that in the male the “peritremal plate differs from that of the female.” In B. patagiorum, however, the transverse ridges produce a distinctly striated and not punctate appearance as figured in Nesbitt and by Keegan. The accessory organ differs in this respect from the peritreme, which does indeed appear punctate. As one focuses on the deeper, more dorsally situated parts of the organ, the ridges disappear, and the outlines change to a form which curiously resembles that of a canoe or gondola with elevated and projecting prow and stern (Fig. 4c). The “prow” and “stern” projections taper to blunt points, or in some specimens to apparently open ends, with the tapered portion at the anterior end occasionally showing some suggestion of coiling. The accessory organ seems to have no connection to the peritreme other than that of proximity. Its restriction to the male suggests a sexual function, possibly as a sensory organ or as a scent releaser. It occurs in the males of B. keegani, as well as in those of B. tarsalis, but is not found in B. dentriticus. I have not seen males of the other species of Blattisocius. The gnathosoma of the male resembles that of the female except for being relatively shorter and broader, and for having the corniculi more widely separated at their bases. The spermatodactyl is as shown in Fig. 3d, e. Leg lengths average 434, 343, 335, and 437 /x for legs I to IV respectively. Leg setation is like that of the female. early stages The eggs are laid singly and adhere only lightly to the substrate. They are smooth, firm, pearly white, and subcylindrical, measuring about 254 by 188 fx . Empty egg cuticula retain the shape of the egg. Larvae and nymphs are in most respects typical for the genus as described by Lindquist and Evans (1965). The larval cuticle is unstriated. The area corresponding roughly with that of the future peritremal shields is covered with coarse granulations or cuticular bosses (Fig. 5a, b). The movable digits of the larval chelicerae are short and broad based, but as in all subsequent stages, without teeth. There is a pair of trumpet-shaped organs in the ventral integument posterior to the third pair of sternal setae [Vol. LXXIV 146 New York Entomological Society lO/A. Fig. 2. Blattisocius patagiorum n. sp. ; a, ventral surface of idiosoma of holotype female; b, spermathecae, showing variations in form, that in the lower figure partly collapsed; c, right chelicera of female ; d, gnathosoma of female. September, 19661 Treat: A New Blattisocius from Noctuid Moths 147 Fig. 3. Blattisocius patagiorum n. sp.; a, dorsal surface of idiosoma of allotype male; b, ventral surface of idiosoma of male; c, left peritreme and accessory organ of allotype male (compare Fig. 4); d, tip of right chelicera of allotype male; e, spermatodactyl of another male at lower magnification and positioned so as to show ventral projection near tip. 148 New York Entomological Society I Vol. LXXIV Fig. 4. Blattisocius palagiorum n. sp.; dark phase contrast photographs of left accessory organ of allotype male at different focal levels; a, ventralmost level: the arc at the lower border of the striated integument in the upper part of the figure appears to be the lateral lip of a fold or pouch covering the deeper portions of the organ; b, intermediate level, showing fusiform portion of organ with transverse ridges or folds; c, deepest level, showing the canoe-shaped portion. The finger-shaped object below the accessory organ in a and b is the left peritreme. September, 1966] Treat: A New Blattisocius from Noctuid Moths 149 Fig. 5. Blattisocius patagiorum n. sp.; a, dorsal surface of idiosoma of larva; b, ventral surface of idiosoma of larva; c, tip of right chelicera of larva; d, dorsal surface of idiosoma of protonymph; e, ventral surface of idiosoma of protonvmph. 150 New York Entomological Society [Vol. LXXIV Fig. 6. Blattisocius patagiorum n. sp.; a, dorsal surface of idiosoma of deutonymph ; b, ventral surface of idiosoma of deutonymph; c, right chelicera of deutonymph. September, 1966] Treat: A New Blattisocius from Noctuid Moths 151 % m Jm i mm ; . •• - m mm- wmm m M H IB Fig. 7. Blattisocius patagiorum n. sp. ; chromosomes from aceto-orcein squash of an embryo of undetermined age. 152 New York Entomological Society [Vol. LXXIV (Fig. 5b). The protonymph (Fig. 5d, e) has the two setae typical of the genus on the palpal trochanter. The soft cuticle is striated in the nymphal and adult stages, and does not show the coarse granulations seen in the larvae. In the deutonymph (Fig. 6) the dorsal shields are united, but with some indication of the line of fusion. type material The above description is based upon 6 females, 5 males, 1 deutonymph, 12 protonymphs, 2 larvae, and 6 eggs, all collected or reared from moths taken in Tyringham, Berkshire County, Massachusetts. One male was found on 24 July, 1958, the other specimens during July, August, and September, 1965. Additional specimens have been taken from pinned moths collected in Giles County, Virginia, in 1956 and now in The American Museum of Natural History. These comprise 2 males, 2 deutonymphs, and 1 protonymph. Moths of the following species, all noctuids, have been found infested: 4 females and 1 male of Spaelotis clandestina Harris; 1 male and 1 female of Pseudospaelotis hams pica (Grote); 1 male of Amphipyra pyramidoides Guenee; 1 female of Septis lignicolora (Guenee). The number of mites per host varied from one to eight. The holotype is from a female of Pseudospaelotis hams pic a found among porch sweepings in Tyringham, Massachusetts, on 31 July, 1965. The allotype male is from the same host; it was observed in copula with a female (not the holotype) on 4 August, 1965. Both holotype and allotype are in The American Museum of Natural History. Paratypes well be sent to the United States National Museum, the Canadian National Collection in Ottawa, and the Institute of Acarology at Columbus Ohio. appearance and behavior The mites were found on the thorax of their hosts, typically facing forward and head down among the hairs and scales on or just behind the patagia. It was this that suggested the specific name patagiorum. As each mite pushes its way down among the hair and scale bases, it creates a temporary, funnel-shaped burrow, at the mouth of which the rear end of the mite can be seen. Adults and deutonymphs are yellow, as is the hemolymph of the host; the earlier stages, at least until feeding begins, are transparently white. The females are somewhat glossy when engorged. The dorsal shield is nearly flat, giving the living mites a rectangular profile in side view. In contrast to more heavily sclerotized ascids, these mites succumb quickly when placed in alcohol or lactic acid. One of the hosts, a female Spaelotis clandestina, survived for more than two months after its capture on the 19th of August, while its mites completed one whole reproductive cycle. This moth was kept at room temperature in a 9 cm plastic petri dish with about six square cm of bibulous paper, moistened occasion- ally to prevent excessive drying. Although I offered the moth a soaked raisin from time to time, I never saw it drink or take any food. It was active only when disturbed, and was probably uninseminated. Two other host moths oviposited during captivity, and in one instance the eggs proved viable. September, 19661 Treat: A New Blattisocius from Noctuid Moths 153 The mites, as a rule, moved about but little, spending many hours or days in a single “burrow.” At intervals ranging from a few seconds to a minute or more, there was a moment of activity for which I can think of no better term than “bustling.” It was impossible to see exactly what the mite was doing at such moments, because its fore parts were always hidden among the hairs of the host. There were leg movements and slight shifts of stance without any resulting change of location. I got the impression that the bustling mite was trying to push more deeply among the hair bases, perhaps seeking closer contact of the mouthparts with the host’s surface. At no time, however, did there appear to be any fixed attachment of the mite to the moth. When removed from its burrow and transferred to a glass observation tube, a mite would wander at random in a way somewhat similar to that of a moth ear mite, Dicrocheles phalaenodectes (Treat, 1965), but with slower and more deliberate gait. The forelegs were kept low and were used to palpate the substrate, only occassionally being lifted into the antennal position. A mite ex- perimentally transferred to a fresh host would soon start to burrow among the thoracic hairs, often with jerky, thrusting movements reminiscent of Dicrocheles. occasionally a mite would leave its burrow spontaneously and wander about the thorax for a time before making another burrow, usually not far from the first. Both sides of the moth were used freely. The act of defecation resembled that in the moth ear mite except that the anus being ventral or subventral rather than terminal, the fecal droplets were left on the floor of the burrow rather than upon objects directly rearward. Spherical white or pale yellow fecal pellets sometimes accumulated about the mouth of a burrow. These were dry and powdery after dehydration, not waxy or gummy as are those of the moth ear mite. Eventually these pellets disappeared, perhaps being dislodged by movements of the mite. A burrow that had been occupied for several days had its flooring hairs or scales lightly stuck together, though not matted or tangled. It may be some component of the feces that causes this stick- ing. Examined microscopically, the feces were seen to comprise a yellow, water soluble component and globular or twined guanine granules averaging about 1.5 [x in diameter. Although no controlled experiments were performed, the mites showed no obvious sensitivity to light. Bustling continued in bright light from a microscope illuminator as well as in the dimmest window light that would allow the mites to be seen. Heat and moisture sensitivity were not tested. On one occasion, within a period of less than eight hours, a mite that had been transferred to a fresh host in a separate petri dish found its way back to the original host. During this time the two dishes had been stacked in a dark box, with their covers raised at one edge by the thickness of a single sheet of bibulous paper. A surprising observation was that at times adult mites in their burrows reacted repeatedly and consistently to ultrasounds. This was first noted while I 154 New York Entomological Society [Vol. LXXIV was testing a host moth with a Galton whistle. The moth showed no reaction, but at each blast of the whistle the mite lurched forward and then made several leg movements. The reaction occurred regularly in tests made at various intervals over a period of several days. It was also tested and confirmed by another observer experienced in insect acoustics, Dr. K. D. Roeder of Tufts University. In one instance the mites continued responding to the sounds for several hours after the death of their host, thus eliminating the possibility that the response of the mites was secondary to some unobserved reaction on the part of the moth. Air turbulence as a possible artefact stimulus was ruled out by substituting an electrically driven Rochelle salt crystal for the Galton whistle. This produced pure ultrasound with no air blast or audible component. In the rated range of 32 to 44 Herz it proved an effective stimulus, while outside that range it evoked no response. It was not possible at the time to monitor this sound source or to check its intensity. No reaction was seen in mites that were already active at the time of stimulation, or in mites that had been placed upon a smooth sub- strate. In the absence of any known or suspected auditory organ, and with no obvious advantage to the mites in possessing such an organ, it seems reason- able to speculate that the effective stimulus for the observed responses was the acoustic displacement of some of the host’s setae in contact with the mite, and that the apparently auditory reactions were in fact mediated by primarily tactile receptors, possibly by the mites’ own setae. No such responses have seen, though often sought, in the moth ear mite. reproduction Living males are not easily distinguishable from females except under high magnification. Their general behavior is similar except that the males move from place to place a little oftener than the females. Encounters between one mite and another did not ordinarily evoke much observable reaction. Even mites of other species ( Dicrocheles phalaenodectes and Lasioseius sp.), when placed experimentally upon a moth infested with B. patagiorum, were allowed to enter a burrow and to climb over the occupant without opposition. I witnessed copulation three times: once (15 September) from its beginning, and twice (4 August and 13 September) when already in progress. A few apparent but un- successful attempts were also observed, in which a male climbed upon the back of a female but then dismounted and went elsewhere. On 15 September a male that had already been in copula with a mite on another moth was transferred to a second host carrying two mites, both probably virgin females though one might still have been a deutonymph at this time. The male approached a burrow on the left patagium, containing one of the mites, but then turned away to wander over the moth’s left tegula and forewing. He soon returned to the same burrow, but again left without entering. At 5:15 PM, five minutes after his transfer to the second host, the male found and entered a burrow on the right patagium, con- taining the second female. He immediately crept under her. embracing her September, 1966] Treat: A New Blattisocius from Noctuid Moths 155 opisthosoma with legs III and IV, his mouthparts at the level of her genital region. Except for slight movements the mites remained quietly in this position for at least three and a half hours. It was not possible to see whether or not a spermatophore was transferred. At 10:20 pm the male was seen leaving the dorsal side of the female, after which he wandered over the moth for a few minutes and was then transferred to alcohol. Five days later his mate, then fully engorged, had left the moth and was lost. She had laid no eggs. The previous mating of the same male with another female on the earlier host had been followed by oviposition within 36 hours. In this case the female had been the only occupant of the moth from its discovery on 19 August until 12 September, when the male was transferred to it from a moth of another species (Aniphipyra pyramidoides) . On the following day the mites were seen in copula, and on 15 September the female laid the first of about 30 eggs. The last egg was laid on 28 September, but the female survived until the death of the host, three weeks later, at which time the mite was mounted for study. Intervals between successive eggs varied from about four to more than twelve hours, the average being probably about eight hours. The temperature varied considerably during the period of oviposition; at the time of four-hour intervals it was about 30° C. On 18 September I watched, under 42. 5X magnification, the laying of the tenth egg, and made the following notes. uAt 3:30 pm bustling movements were occurring every three or four seconds, but they became less frequent until by 3:50 the mite was quiet for a minute or more at a time. She was well engorged, with no depression of the ventrianal plate. Her white, nodular malpighian tubules showed intermittent undulations, some beginning proximally (nearest the rectum, which was full of white matter) and some distally, the former being the more frequent. There were also elongations and shortenings of the malpighian tubules, but no translational movements of the nodes. At intervals of a minute or more the ventrianal plate was deeply depressed, most markedly on the left side, where also, a large ovoid white mass could be seen through the dorsal sur- face. I thought at first that this mass was the tenth egg, but this proved incorrect, for it was still there after the tenth egg had been laid. It may have been the eleventh. Twice there were movements that suggested compressional straining. At 4:10 pm the gnathosomal end of the mite was slowly lifted up as the egg was passed forward. This egg emerged more slowly than a Dicroc heles egg but was free within about five seconds after the movement began. The ventrianal plate was deeply depressed at this time, and remained so until about 4:30, when the opisthosoma was regaining its distended form through re-engorgement or other- wise. Immediately after the egg was free, the mite caressed it a few times with her forelegs and palpi, but then moved aside slightly and began a series of jerky, thrusting movements toward the depths of the burrow, which had the effect of shifting the egg rearward along her left side. She then probed deeply into the 156 New York Entomological Society [Vol. LXXIV burrow and became quiet, possibly feeding, until about 4:30 pm, at which time the usual bustling was resumed.” The eggs adhered only lightly to the host, and if not removed experimentally were lost from the moth within a few hours. Their surface was dry, and they were often electrostatically repelled by the needle when I tried to pick them up. They hatched in from one to three and a half days, probably depending upon the temperature. The last four shrivelled and failed to hatch. I squashed ten of the eggs in aceto-orcein, but although the chromosomes stained fairly well I could not determine the chromosome number unequivocally. In many of the cells there were two short, straight chromosomes, two straight ones of intermediate length, and two long V- or C-shaped bodies, which, if these last were single units, would give a chromosome total of six, but if double (i.e., actually two chromosomes each), a total of eight. I think six is the more likely number. Some cells, however, appeared to have only three, and some four chromosomes, while others seemed to be polyploid. All of the embryos yielded similar squashes; there was no sign of ‘‘commas” or sex chromatin masses as in males of Dicrocheles (Treat, 1965). The larvae were water white. They moved with a rhythmic, swinging gait, with legs I in the antennal position. When placed on a moth, some larvae, after a momentary freeze of ten seconds or more, began to wander superficially over the scale tips. These were soon brushed or flicked off by sudden movements of the moth. Other larvae burrowed among the scales much as do the adults, but farther back on the thoracic disc. These remained on the host and within a few hours transformed into protonymphs, leaving their exuviae on the floor of their burrows. Evidently feeding is not necessary in the larval stage, because protonymphs were produced from larvae kept in glass vials without food. The protonymphal stage varied in duration from a few hours to two days, and in the longer period at least, involved some feeding. The deutonymphs be- came yellow and engorged, and in this condition were not easily distinguished from adults. In one instance, transformation to the adult occurred after a deutonymphal stage of four days, the total time from egg to adult in this case being ten days. Molting was not observed directly, but in all instances the cast skins were left on the floor of the burrow. DISCUSSION The details given above raise questions with regard to the relationship between these mites and their noctuid hosts. Are the mites to be considered parasites, or are they not? And if not, what then? Certainly the association involves something more than phoresy. The long survival period, the ability of the female to produce many viable eggs, and of the offspring to reach adulthood on the original host, all indicate a source of food either in or on the host itself, although the September, 1966] Treat: A New Blattisocius from Noctuid Moths 157 failure of the eggs to adhere to the host suggests that in nature the earliest stages, at least, may be passed elsewhere. For the instars actually associated with moths, whether regularly or only occasionally, commensalism in the strict sense is unlikely, since the moths under observation took no food and were not dusted with pollen. The remaining possibilities are parasitism and phagophily — the use of other symbionts as food. If these mites were phagophiles, they certainly did not feed upon other mites, since none was present except when one or two were placed upon the moths ex- perimentally, and these were ignored by the Blattisocius. Conceivably the food was some kind of microorganism. To be sure, the long-surviving host (numbered 85 for identification) occasionally had small patches of white mycelial growth upon its thorax. The hyphae were septate and bore spores of various sizes on short conidiophores. But this bloom was apparently ignored by the mites, and it disappeared when the humidity was reduced. The patagia of some arctiid moths give out a repugnatorial secretion, but no such secretion has been seen or described in the noctuids with which we are concerned. Some months after moth number 85 had been injected with alcoholic Bouin’s solution, I denuded the patagia and examined them microscopically. Along their dorsal margins, in places previously occupied by the mites, were several minute, dark brown discolorations. Under high magnification these appeared to be limited to the goblet-like bases of individual scale sockets. The bustling activities of the mites, the stylet-like shape of their movable chelae, and the appearance of their midgut and rectal contents suggest that the food is hemolymph which exudes from minute punctures in the host’s cuticle, possibly through the scale bases. The bustling movements might be concerned with removing plugs of coagula and releasing a fresh supply of the liquid. This notion is, of course, wholly speculative and may prove quite incorrect. According to Lindquist and Evans (1965), “No ascid mites are known to be truly parasitic.” If B. patagiorum were shown to be a true parasite, the questions would still remain whether its parasitism is facultative or obligate, and whether the choice of hosts is restricted to moths. Other species of the genus have been reported from many different hosts and habitats including lizards, birds1 nests, mammals, and various kinds of moths and other insects, particularly those infesting stored grains (Hughes, 1961). I have found B. dentriticus on the noctuid Pseudaletia adult era (Schaus) from Pelotas, Brazil, and also on a notodontid, Datana ministra (Drury) from New Jersey. I have found B. keegani on this same species of notodontid, on the noctuids Folia contigua (Schiff.) from Kyoto, Japan, and Zale lunata (Drury) from Charleston, South Carolina, and on a tineid, Tineola biselliella (Hum.) from Pittsburgh, Pennsylvania. I have taken B. tarsalis from the noctuids Crymodes devastator (Brace) from Salt Lake City, Utah, and Epizeuxis aemula (Hbn.) from Tyringham, Massachusetts. In several 158 New York Entomological Society [ Vol. LXXIV instances (e.g., Rivard, 1960) Blattisocius species have been shown to be predators on other mites, though capable of living also upon molds. It is noteworthy that the hosts of B. patagiorum as recorded on page 152, though representing two more or less divergent noctuid subfamilies, have this in common — that they characteristically rest by day in crevices under the bark of dead trees or in dead wood, often in the joints and crevices of buildings. Such situations favor mite populations of various kinds, and could be expected to yield occasional examples of disjunctive or facultative association between some of the regular occupants and casually intruding moths. I have come across other instances of such association, involving various gamasines, particularly ascids of the genera Proctolaelaps and Lasioseius , which I hope to report elsewhere. It is interesting to note that notwithstanding the latitude in the selection of host species suggested by these records, there is considerable restriction in a given species of mites with regard to the part of the host’s body that is occupied. Blattisocius patagiorum, for example, is recorded only from the thorax of the host, and usually from its dorsal surface. My specimens of Proctolaelaps and Lasioseius , by contrast, regardless of the moth species on which they were discovered, have almost invariably been found between the palpi, under the base of the proboscis. This consistency in site selection might argue some degree of regularity in the association of the mites with moths, but it could also be merely the result of inherent differences in responsiveness to tactile or other stimuli, which, though perhaps adaptive in some other context, might lead to relatively meaningless differences in the sites occupied on casually or accidentally boarded hosts. Many more collection records and behavioral studies will be needed to resolve such problems. In any event, it seems unlikely that the mites in question significantly reduce the life span or population density of their noctuid associates. acknowledgments: I thank Dr. Evert E. Lindquist of the Canada Department of Agriculture for critically examining both specimens and manuscript, and for calling my attention to details that I should otherwise have overlooked. Literature Cited Chant, D. A. 1963. The subfamily Blattisocinae Garman ... in North America, with descriptions of new species. Canadian Jour. Zook, 41: 243-305. Evans, G. O. 1958. A revision of the British Aceosejinae (Acarina: Mesostigmata) . Proc. Zool. Soc. London, 131: 177-229. Hughes, A. M. 1961. The mites of stored food. Technical Bulk No. 9, Ministry of Agriculture, Fisheries and Food. London: Her Majesty’s Stationery Office. Keegan, H. L. 1944. On a new genus and species of parasitid mite. Jour. Parasitok, 30: 181-183. Lindquist, E. E., and G. O. Evans. 1965. Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostig- mata). Mem. Entom. Soc. Canada, No. 47. September, 1966] Treat: A New Blattisocius from Noctuid Moths 159 Nesbitt, H. H. J. 1951. A taxonomic study of the Phytoseiinae (family Laelaptidae) predaceous upon Tetranychidae of economic importance. Zool. Verhandel. (Leiden), 12: 1-64. Oudemans, A. C. 1929. Acarologische Aanteekeningen, C. Entom Ber. Amsterdam, 8(170): 28-36. Rivard, I. 1960. A technique for individual rearing of the predacious mite Melichares dentriticus (Berl.) (Acarina: Aceosejidae) , with notes on its life history and be- haviour. Canadian Entomologist, 92: 834-839. Treat, A. E. 1954. A new gamasid . . . inhabiting the tympanic organs of phalaenid moths. Jour. Parasitol., 40: 619-631. . 1965. Sex-distinctive chromatin and the frequency of males in the moth ear mite. Jour. New York Entom. Soc., 73: 12-18. Received for publication June 20, 1966 160 New York Entomological Society I Vol. LXXIV Proceedings of the New York Entomological Society (Meetings held in Room 129 of the American Museum of Natural History unless otherwise indicated) Meeting of February 1, 1966 President Richard Fredrickson presided; 14 members and 2 guests were present. Miss Margaret Pogany was elected to membership and Mr. Howard Topoff, a graduate student at the City University, was proposed for student membership. Dr. Rozen introduced Dr. Herbert Ruckes, Jr., the son of our recently deseased Dr. Ruckes. He is a specialist in the Anobiidae (Coleoptera) . Dr. Asher Treat proposed him for membership. program. Blackflies of Western South America. Dr. Pedro Wygodzinsky of the Museum staff discussed the biogeography of blackflies and the attempts by others and himself to find primitive genera in Western South America. Available evidence indicates that the more primitive forms are limited to the Northern Hemisphere. Thus, either the group originated in the Northern Hemisphere and radiated southward, or the primitive forms have died out in South America; this latter explanation does not seem likely. The talk was illustrated with specimens and slides. David C. Miller, Sec. pro tem. Meeting of February 15, 1966 Dr. Fredrickson presided; 25 members and 4 guests were present. Mr. John Pallister presented the report of the Auditing Committee for the year 1965 and stated that the Society’s financial records are in proper order. Dr. Herbert Ruckes, Jr., and Mr. Howard Topoff were unanimously elected to full and student membership respectively. Mr. Aaron Nadler, a specialist in the Psocoptera who has done a great deal of collecting for the Museum, was proposed for membership. A note from Mrs. Herbert Ruckes, Sr. was read thanking the Society for the memorial resolution and the expression of sympathy which was sent to her on her husband’s death. Miss Joan Todd, a grade school Biology teacher, was introduced as a guest. program. A World Without Butterflies, and One Man’s Fight to Delay It. Dr. Kurt Gohla, Professor of German, Fordham University was the speaker of the evening. (An abstract follows.) David C. Miller, Sec. pro tem. A World Without Butterflies, and One Man’s Fight to Delay It On a visit to Germany during the summer of 1965, a collecting trip to Tegernsee, a mountain resort in the foothills of the Bavarian Alps, was made expressly to obtain the Black Apollo butterfly, Parnassius mnemosyne L. In spite of fertile mountain meadows, neither this species nor any other Lepidoptera were seen. An effort to explain the diminishing of butter- flies and moths in this area is offered in a pamphlet issued by the Society for the Protection of Alpine Flowers and Animals. Three possible causes are under consideration by Doctor Max Dingier, Professor of Zoology at the University of Munich: Atomic contamination by radioactive dust in the atmosphere which may have a sterilizing effect upon the reproductive organs of insects in general; Electromagnetic sound waves which may interfere with the fine system of sense organs located in the antennae of the Lepidoptera; September, 19661 Proceedings 161 The use of artificial fertilizers which have caused some wild flowering plants, preferred by butterflies, to disappear. The disappearance of Lepidoptera from their customary mountain meadows and haunts constitutes a loss of ethical and esthetic values and would be an impoverishment of our entire social way of living. Color slides were shown demonstrating the work of an amateur lepidopterist, Mr. Walther Ender of Lage, Westphalia, who breeds Lepidoptera in great numbers and releases them in order to repopulate the area of the Teutoburg Forest in the northwestern part of West Germany. Kurt Gohla Meeting of March 1, 1966 President Fredrickson called the meeting to order; 28 members and 10 guests were present. Mr. Aaron Nadler was elected to membership. Dr. Edwin W. Teale read excerpts from a letter he had received from Mr. Roy Latham, now 85 years old, telling of his experiences with lights to attract moths at Orient Point, Long Island. Almost none came to the lights and those that did were common ones; only very few oher insects, such as Japanese beetles, are collected at lights. Dr. Pedro Wygodzinsky told of weevils from New Guinea that are covered with lichens and mosses in which mites are found. program. Zoological Collecting in New Guinea. Mr. Hobart M. Van Deusen, a curator in the Museum’s Department of Mammology and in charge of the Archbold Collections, opened his talk by showing the pelts of some of the few mammals that are found in New Guinea: a bat with a wing spread of five feet; an arboreal, giant rat, the largest specimen which is a trifle short of three feet; a spiny anteater, and a tree-climbing kangaroo. All of the animals are nocturnal which makes collecting rather difficult. Since 1933 the Archbold ex- peditions have returned to New Guinea every 3 or 4 years. The last one in 1964 explored the Huon Peninsula were rift valleys separate mountain peaks into what are virtually islands. Remarkable slides were shown which gave excellent views of the terrain, mountain peaks, plateaus, and caves, as well as the mammals, including one of a kangaroo with young in its pouch. Lucy M. Heineman, Sec. Meeting of March 15, 1966 Doctor Fredrickson presided ; 17 members and 9 guests were present. Mr. John A. Novak was proposed for student membership. Miss Alice Gray exhibited specimens of wingless scorpion flies collected by a former student now at Ithaca. Dr. Asher Treat questioned a statement in a story on Brachymeria intermedia , a parasite of the Gypsy Moth (New York Times, Sun- day, March 13, 1966), that these parasitic wasps do not sting humans. He reported having been stung several times by ichneumon wasps. Dr. Elsie Klots recounted a similar experience. The stings were painful but did not produce swellings or after-effects. He also called attention to an account by H. E. Hinton and M. S. Blum of the University of Bristol, England (New Scientist, Oct. 28, 1965, pp. 2 70-1) summarizing Hinton’s experience with the larvae of the chironomid fly, Polypedilum vanderplanki (Hint.) which is able to produce apparently normal adults when restored to water after total dehydration and exposure, in the dry state, to temperatures as low as -270 degrees and as high as 104 degrees centigrade. The ability to survive alternate hydration and dehydration in this and in many more primitive organisms has suggested to the authors that life may have originated not in the sea, as is generally supposed, but in rock crevices or similar situations on land. 162 New York Entomological Society [Vol. LXXIV program. The Importation of Foreign Plant Material. Mr. Charles A. Andrews of the Plant Quarantine Division of the U.S. Dept, of Agriculture discussed the need for restrictions on imported plants and plant materials, and he traced the history of our present regulations. He stressed the point that the Division has attempted to develop a plant pest protection program which will give us the maximum interference with commerce. The steps used in making inspections and the procedures in processing plants which enter our country from foreign propagators were outlined. Mr. Andrews showed slides depicting the carrying out of the restrictive provisions of the Division in the handling of tulip bulbs in Holland. Some showed the pests in bulbs and nuts, others were microscopic sections to explain how the identification of the pests are made. Lucy M. Heineman, Sec. Meeting of April 5, 1966 President Fredrickson called the meeting to order; 14 members and 4 guests were present. Mr. John A. Novak was elected to student membership and Mr. Robert Mesibov was proposed for membership. Miss Alice Gray demonstrated a fossil arthropod which showed up well when illuminated with ultra-violet black light. program. Fossil Roaeh-like Insects from the Carboniferous. Mr. Christopher Durden of the Biology Department of Yale University discussed the distribution and the classifica- tion of the numerous roach-like fossils which are now available. Wing venation and their manner of folding are important features. Some had wing margins that might have been used for stridulation. Most of the Carboniferous roach fossils are about as large as our present roaches. The talk was illustrated with slides. Albert J. Poelzl, Assistant Secretary Meeting of April 19, 1966 President Fredrickson presided; 31 members and 7 guests were present. Mr. Robert Mesibov was elected to student membership, and Miss Alice Gray proposed Mr. Kenneth Friedman and Mr. David F. Kanter for student memberships. Dr. Alexander Klots introduced Dr. and Mrs. Traub of Bethesda, Maryland. Dr. Traub, a former student at C.C.N.Y., a retired Army colonel, is an authority on fleas as typhus carriers. Miss Anne Birdsey called attention to an article on the science page of the Sunday New York Times written by Norton T. Novitt, a Denver, Colorado amateur scientist, which proposed that flying saucers may be electrified flying ants. She also showed a paperback copy of “1001 Answers About Insects” by Alexander and Elsie Klots. Dr. Klots announced that the Honorable Miriam Rothschild is now in the United States. Unfortunately, she was not able to stay in New York for tonight’s meeting. She is currently engaged in a project concerning repellant insecticides which necessitates her using many specimens of the moth, Diacrisia virginica. She would appreciate having egg masses of this moth air-mailed to her at Elsfield Manor, Oxford, England. Miss Alice Hopf is anxious to obtain specimens of the viceroy butterfly in any of its stages. Program. Termites and Evolutionary Processes. Dr. Alfred E. Emerson, Professor Emeritus of the University of Chicago, a Research Associate in the Dept, of Insects of the Museum for many year, discussed regressive evolution, recapitulation, convergent evolution, and the evolution of behavior as illustrated by termites. He stressed that the unit of natural selection in these insects is the entire colony rather than the indivdual. The king and the queen are the only individuals in the colony capable of reproducing, and the genes controlling structural and adaptive characteristics which are manifested in the sterile castes, September, 1966] Proceedings 163 the workers and the soldiers, are transferred through these reproductives ; though they, themselves, do not manifest these characteristics. The talk was illustrated with slides. Lucy M. Heineman, Sec. Meeting of May 3, 1966 Dr. Fredrickson called the meeting to order; 15 members and 9 guests were present. Several guests were introduced: Mr. Harry Steen; Mrs. Michal Emsley of the New York Zoological Society Research Station, “Simla,” in Trinidad; Dr. and Mrs. Leon Cahen who are active members of the Explorers Club. Mr. Kenneth Friedman and Mr. David Kanter were elected to student membership. Mr. Kennith Watson, Mr. H. Steen, and Dr. Philip Spear were pro- posed for membership. Dr. Fredrickson mentioned that progress is being made on the pro- posed merger of the N.Y. and the Brooklyn Entomological Society. The lawyers for the two societies are in consultation and an agreement has been drawn up . The members will be kept informed about future developments and will be required to vot on the agreement after it has been approved by the Executive Committee of our Society. The President, also, announced that our member, Dr. Edwin Way Teale, received a Pulitzer Prize for his book, “Wandering Through Winter,” the final volume of his history of the four seasons in America. The Secretary was instructed to convey to Dr. Teale the hearty congratulations of the Society on the receipt of this well-deserved honor. Program. Entomology and the National Pest Control Operators. Dr. Philip Spear, Technical Director for the National Pest Control Association, explained that Pest Control operators are concerned with pests in and around structures, within the contents of the structures, and with the use and the problems of pesticides. Insects occupy a large part of the time and energy of the operators. Thus, entomology in all its phases is an important study in this industry. About 5,000 firms are members of the Association, and approxi- mately 30,000 workers are employed in the field. They deal, usually, with emergency situations, but they do prefer to operate on a preventative basis. His talk was illustrated with many slides which showed the scope of work done in structures, the damages done by various pests, and the pests. Lucy M. Heineman, Sec. Meeting of May 17, 1966 President Fredrickson presided; 27 members and 21 guests were present. One of the guests present was Dr. John Vandenburg of the New York University Medical School, Dept, of Preventive Medicine. Mr. Kennith Watson, Mr. Harry Steen, and Dr. Philip Spear were elected to membership. Dr. James Forbes, Associate Editor of the Journal, reported on the 10th Annual Meeting of the Council of Biological Editors which was held May 3-4 in the Center for Continuing Education on the campus of Notre Dame University. Dr. Forbes represented the Society at this meeting. Dr. Klots read a letter from the Edwin W. Teales in which he thanked the Society for its good wishes and described happenings on their trip through England. Dr. Fredrickson reported that on the proposed merger of our Society with the Brooklyn Entomological Society it will probably be necessary to call for a special meeting in order to vote on the final agreement. Notices will inform the membership. Program. National Geographic Society Motion Picture on the work being done by Dr. L. S. B. Leaky in finding human fossil remains in the Oldvai Gorge in Africa; filmed by Baron Hugo van Lavick. This was accompanied by a recorded, running commentary by Dr. Leaky. It was a fascinating film showing the work camps, the terrain, and how the fossils were found. It supported Dr. Leaky’s theory that there were two contemporary 164 New York Entomological Society [Vol. LXXIV types of man, herbivorous and carnivorous. The film demonstrated differences between the mouthparts and the feeding of these two types of animals. The photography of insects and the feeding of the different forms was superb. Lucy M. Heineman, Sec. Recent Publications Aspects of Insect Biochemistry. 1965. Biochemical Society Symposium (London), T. W. Goodwin, Ed. Academic Press, New York, 119 pp., illus., $6.00. Seven papers: “Active Transport in Insects” by J. E. Treherne; “Formation of the Specific Structural and En- zymic Pattern of the Insect Flight Muscle” by Th. Bucher; “Some Distinctive Features of Insect Metabolism” by F. P. W. Winteringham ; “Intermediary Metabolism and the Insect Fat Body” by B. A. Kilby; “The Metabolism of Aromatic Compounds” by P. C. J. Brunet; “Hormones Controlling Growth and Development in Insects” by V. B. Wiggles- worth; and “Skeletal Structure in Insects” by K. M. Rudall. Pesticides in Clinical Practice, Identification, Pharmacology and Therapeutics. 1966. Royal L. Brown. Charles C. Thomas, 504 pp., $15.75. The Entomology of Radiation Disinfection of Grain. 1966. Edited by P. B. Cornwall. Pergamon Press, Long Island City, New York, 256 pp., $9.50. Ticks of the Genus Ixodes in Africa. 1966. Don R. Arthur. University of London Press, London: Oxford University Press, New York, 365 pp., $11.20. Reviewed in Science 152: No. 3723, p. 750. Polymorphism in Some Nearctic Halictine Bees. 1966. G. Knerer and C. E. Atwood. Science 152: 1262-1263. Classification of the Bees of the Australian and South Pacific Regions. 1965. Charles D. Michener. American Museum of Natural History Bulletin, 130: 1-362, $10.00. Termites (Isoptera) of Thailand. 1965. Muzaffer Ahmad. American Museum of Natural History Bulletin, 131: 1-114, $2.00. Insect Aerodynamics: Vertical Sustaining Force in Near Hovering Flight. 1966. Leon Bennet. Science, 152: 1263-1266. A Revision of the Neotropical Genus Metamasius (Coleoptera: Curculionidae, Rhyn- chophorinae) : Species Groups I and II. 1966. Patricia Vaurie. American Museum of Natural History Bulletin, 131 : 211-338, $5.00. Contributions Towards a Revision of Myrsidea Waterson I (Mallophaga: Menoponi- dae). T. Clay. British Museum (Natural History) Bulletin: Entomology, 17: 327-395, 1£ 10s. A Revision of the British Aleyrodidae (Hemiptera: Homoptera). L. A. Mound. British Museum (Natural History) Bulletin: Entomology, 17: 397-428, (14s). The Interrelationships of Three Gall Makers and Their Natural Enemies, on Hack- berry ( Celtis occidentalis L.). John Conrad Moser. 95 pp., $1.00. A Handbook for the Identification of Insects of Medical Importance. 1965. John Smart with chapters by Karl Jordon and R. J. Whittick. British Museum (Natural History), London, 4th ed., 340 pp., <£3. Reviewed in Science, 152: 748-749. INVITATION TO MEMBERSHIP The New York Entomological Society was founded in 1892 and incorporated the following year. It holds a distinguished position among scientific and cultural organizations. The Society’s Journal is one of the oldest of the leading entomological periodicals in the United States. Members and subscribers are drawn from all parts of the world, and they include distinguished professional naturalists, enthusiastic amateurs, and laymen for whom insects are only one among many interests. You are cordially invited to apply for membership in the Society or to subscribe to its Journal which is published quarterly. Regular meetings are held at 8:00 P.M. on the first and third Tuesdays of each month from October through May at the American Museum of Natural History, the headquarters of the Society. A subject of general interest is discussed at each meeting by an invited speaker. No special training in biology or entomology is necessary for the enjoyment of these talks, most of which are illustrated. Candidates for membership are proposed at a regular meeting and are voted upon at the following meeting. CLASSES OF MEMBERSHIP AND YEARLY DUES Active member: Full membership in the Society, entitled to vote and hold office; with Journal subscription $9.00 Active member without Journal subscription 4.00 Sustaining member: Active member who voluntarily elects to pay $25.00 per year in lieu of regular annual dues. Life member: Active member who has attained age 45 and who pays the sum of $100.00 in lieu of further annual dues. Student member: Person interested in entomology who is still attending school; with Journal subscription 5.00 (Student members are not entitled to vote or to hold office.) Student member without Journal subscription 2.00 Subscription to Journal without membership 8.00 APPLICATION FOR MEMBERSHIP Date I wish to apply for membership (see classes above). My entomological interests are: If this is a student membership, please indicate school attending and present level. Name Address (Zip Code must be included) — Send application to Secretary — p Vol. LXXIV DECEMBER 1966 ? I Devoted to Entomology in General i ■f C-|f r '5 , r. The X 1 ' ~'\ ' f ■ Jr( , 'S f ' ' \ J New York Entomological Society ifk " h Organized June 29, 1892 — Incorporated February 25, 1893 Reincorporated February 17, 1943 ■{ r X' A, The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St., & Central Park W., New York 24, N. Y. ■> Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. Members of the Society will please remit their annual dues, payable in January, to the Treasurer. r 4 , v* Officers for the Year 1966 .P President , Dr. Richard Fredrickson College of the City of New York 10031 Vice President , Dr. Kumar Krishna American Museum of Natural History, New York 10024 Secretary , Mrs. Lucy Heineman 115 Central Park West, New York 10023 4V, Assistant Secretary , Mr. Albert Poelzl T' 230 E. 78th Street, New York 10021 Treasurer , Mr. Raymond Brush American Museum of Natural History, New York 10024 & Assistant Treasurer , Mrs. Patricia Vaurie American Museum of Natural History, New York 10024 Trustees 1 YearTerm ■ , . . Dr. Alexander B. Klots 2 Year Term Dr. Jerome Rozen, Jr. Dr. John B. Schmitt Mr. Robert Buckbee ‘ u Mailed December 29, 1966 nological Society is published q Inc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas. The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press :1a Journal of the New York Entomological Society Volume LXXIV December 29, 1966 No. 4 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen Columbia University College of Pharmacy 115 West 68th Street, New York, N. Y. 10023 Associate Editor James Forbes Fordham University, New York, N.Y. 10458 Publication Committee Dr. Pedro Wygodzinsky Dr, Asher Treat Dr. David Miller CONTENTS The Nerves of the Thoracic Segments of the Larva of Prodenia litura (Lepidoptera: Noctuidae) J. Bahadur and B. B. L. Srivastava 168 Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XIII Charles P. Alexander 180 The Larva of Amblyscirtes samoset (Scudder) (Lepidoptera: Hesperii- dae) Alexander B. Klots 185 Studies on Parasitic Mites of New Jersey Jack R. Manischewitz 189 Structure of Gastric Apex as a Subfamily Character of the Formicinae (Hymenoptera: Formicidae) __ Akev C. F. Hung and William L. Brown, Jr. 198 Xenillidae, A New Family of Oribatid Mites (Acari: Cryptostigmata) Tyler A. Woolley and Harold G. Higgins 201 Pieris narina oleracern (Harris) in New Jersey (Lepidoptera: Pieridae) Cyril F. dos Passos 222 Two North American Spiders (Araneae: Linyphiidae) Wilton Ivie 224 Notes 188 Book Reviews 228 Index of Scientific Names, Volume LXXIV 231 Index of Authors, Volume LXXIV iii 168 New York Entomological Society [Vol. LXXIV The Nerves of the Thoracic Segments of the Larva of Prodenia litura (Lepidoptera: Noctuidae) J. Bahadur and B. B. L. Srivastava School of Studies in Zoology, Vikram University, Ujjain (India) Abstract. The nervous system of the thoracic segments of the larva of Prodenia litura is described. The dorsal and transverse nerves remain connected to each other at three points through three connectives. But in the prothoracic segment, there is no transverse nerve, so that the dorsal nerve establishes a connection with subconnective nerve by a plexus. Ordinarily no connection is found between the dorsal and ventral nerve but in the pro- thoracic segment, such a connection is established at one point. INTRODUCTION The studies on the nervous system started with the work of Lyonet ( 1762). Since then many aspects of it have been dealt with. Du Porte (1915) described the nervous system in Sphida and Ruckes (1919) studied the innervation of the male genital organs in certain lepidopterans. However, the real work on the nerve pattern started with Maki (1936) who described it in the alderfly, Chauliodes jormosanus. Nesbitt (1941) studied the nerve patterns in Orthop- tera and other related orders. Schmitt ( 1954, 1959) studied the nervous system of cervicothoracic and the pregenital abdominal segments in some orthopterans. With these studies it was realized that there exists a basic segmental nerve pattern in insects. Whether such a homology can be traced in widely separated orders as Orthoptera and Lepidoptera, is yet to be seen. Libby (1959, 1961), however, investigated the nerve pattern of certain abdominal segments of the larva and adult of the moth, Hyalophora cecropia and tried to establish homol- ogy with other insect nerve patterns. The short review shows that the thorax has not been tackled so far in detail. To fill up this lacuna and to establish how far there exists a basic homology with the thorax of other insect orders, the authors undertook a very detailed study of the nerves of the thoracic seg- ments of the larva of Prodenia litura. MATERIAL AND TECHNIQUES The full grown larvae were directly collected from the cabbage fields and kept in the laboratory. For the studies on the distribution of the nerves, 1% methylene blue in normal salt solution was injected into the body cavity of the larva. After a few hours, the insect was etherized and dissected in normal saline (0.65%). Sometimes, instead of injecting the solution into the body cavity, the dye was directly poured over the dissected animal and allowed to stay for 2 to 4 hours to secure better staining of finest motor nerves. Further dissection was done in normal saline. To destain the adjoining tissues, acid water was sometimes used. Normally, all the nerves of a particular segment December, 1966] Bahadur and Srivastava: Lepidopteran Nerves (Prodenia) 169 could not be traced in one day and hence the dissection used to be kept in normal saline with a few drops of formalin. In such preservation, the blue colour of the nerves disappears but they remain quite distinct because of the milky white appearance which they attain. All the dissections were carried out under the stereoscopic binocular microscope in artificial light. The diagrams are purely diagrammatic. OBSERVATIONS The thorax is composed of three segments with their ganglia. The prothoracic ganglion remains connected to the suboesophageal ganglion by a pair of stout but short connectives which lie free throughout their entire length. The con- nectives between the other ganglia lie united anteriorly for about one fifth of the distance and then diverge gradually, continuing their course separately until they enter the anterior border of the succeeding ganglion. The two sepa- rated connectives enclose between them some space within which the diagonal muscles cross each other near their point of insertion. The enclosed space is smaller in the prothoracic segment but larger in the other two segments. NERVES OF THE PROTHORACIC GANGLION (Fig. 1) The prothoracic ganglion gives rise to two pairs of lateral nerves and a pair of subconnective nerves. The lateral nerves are designed as the dorsal and the ventral nerves. From the median portion of the ganglion arises a pair of sub- connective nerves. The Dorsal Nerve : The dorsal nerve (DN) leaves the ganglion and runs obliquely outwards and upwards over the ventral median muscles and ventral internal lateral muscles to reach the subconnective nerve (SN) with which it forms a plexus (px). It then sharply bends downwards, passes over the ventral internal muscles and extends to a considerable distance, giving branches at intervals. The first branch (ID) arises over the ventral internal longitudinal muscle and divides into two branches; the inner branch (a) passes downwards and curves slightly inwards and bifurcates to innervate the tracheae and the tracheoles. The outer branch (b) curves and bifurcates into b' and b". Whereas the former innervates the ventral internal lateral longitudinal muscle, the latter meets the longitudinal nerve of the dorsum (LND) which extends from the head up to the intersegmental fold of this segment. The main dorsal nerve proceeds further and after a short distance, besides receiving the sixth branch (6V) of the ventral nerve, itself gives rise to the second branch (2D). This branch divides into a number of branches to innervate the adjoining neck muscles and the tracheae. The third branch (3D) proceeds dorsally and gives rise to a number of branches which again innervate the various muscles and integument of the neck region. The fourth branch (4D) innervates the tergosternal muscle. The main dorsal nerve ulti- 170 New York Entomological Society [Vol. LXXIV Fig. 1. litura. Diagram of the nerve pattern of the prothoracic segment of the larva of Prodenia mately terminates into fine branches supplying the dorsal longitudinal muscles. the ventral nerve: The ventral nerve (VN) leaves the ganglion at the middle of the lateral margin and passes posteriorly below the ventral median muscle group and over the ventral external oblique muscles. It gives rise to a number of branches. The first branch (IV) runs obliquely downwards and bifurcates into a and b. The former passes deep into the prothoracic leg to innervate its muscles whereas the latter innervates the ventral internal and external oblique muscles. The second branch (2V) gives rise to a fine branch (a) which supplies the muscles of the leg and the other branch (b) innervates the ventral median muscle. The third branch of the ventral nerve (3V) subdivides into a number of fine branches to innervate the tracheae and ventral internal lateral muscles. The main ventral nerve after proceeding ahead for a short distance, curves anteriorly above the ventral internal lateral muscle and flattens. From the proximal part of this arises a small fourth branch (4V) which innervates the tracheae. The fifth branch (5V) innervates the tracheae and the ventral internal lateral muscles. The sixth branch (6V) extends to join the main dorsal nerve. Whereas the seventh branch (7V) proceeds as far as the prothoracic ganglion and innervates the ventral external oblique muscles, the eighth nerve (8V) December, 1966] Bahadur and Srivastava: Lepidopteran Nerves (Prodenia) 171 extends into the head to supply the tracheae and muscles of that region. The ninth nerve (9V) gives origin to a number of minute nerves which innervate the various ventral longitudinal and oblique muscles and tracheae of the neck and adjoining regions. the subconnective nerve : There is no median nerve in this ganglion so that the transverse nerves are also absent. From the mid antero-dorsal side of the prothoracic ganglion arises a single nerve which may be taken as the median nerve. It proceeds anteriorly for a very short distance and then bi- furcates above the suboesophageal ganglion to give rise to a pair of the so- called subconnective nerves (SN). Each nerve passes laterally to innervate the various muscles of the head but before taking a curve, a plexus (px) is formed between it and the adjoining dorsal nerve. NERVES OF THE MESOTHORACIC GANGLION (Fig. 2) the dorsal nerve: This nerve (DN) arises from the outer margin of the interganglionic connective, just a few millimeters above the mesothoracic ganglion. It passes laterally over the external and internal median muscles and after a short distance extends its first branch (ID) which passes over the ventral internal lateral longitudinal muscles and fuses with the transverse nerve. It, however, gives rise to a branch (a) which extends another three small branches to innervate the tracheae, ventral internal lateral muscles and the lateral internal oblique muscle. The main dorsal nerve proceeds ahead and gives rise to another connective branch (2D) which also fuses with the transverse nerve, just posterior to the spiracle. But before fusion, it gives rise to a branch at the point c' which proceeds antero-dorsally and divides into a number of minute branches. The branch cl extends posteriorly to innervate the two pleurosternal oblique muscles, the branch c2 innervates the tergosternal muscle and the branches of the lateral tracheal trunk and the branches c 3 and c4 proceed to innervate the dilator and occlusor muscles of the spiracle respectively. In addition to these, the branch 2D gives rise to two small branches (a and b) which innervate the lateral internal oblique and ventral external oblique muscles respectively. The main dorsal nerve passes dorsally above the lateral longitudinal tracheal trunk and gives rise to a connective (3D) which runs to fuse with the trans- verse nerve. The connective gives off two minute branches posteriorly and they innervate the dorsal and lateral external oblique muscles and tracheae. The fourth branch (4D) innervates the integument and the dorsal internal lateral muscle. The main nerve, by now, becomes thin and extends ahead into the dorsal region, giving off small branches. The branches 5D, 6D and 7D innervate the dorsal external oblique muscle and dorsal internal lateral muscle group. The main nerve ultimately terminates into a number of very fine branches which innervate the tracheae and the dorsal internal median and 172 New York Entomological Society [Vol. LXXIV Fig. 2 Fig. 2. Diagram of the mesothoracic nerve pattern of the larva of P. litura. oblique muscles. The longitudinal nerve of the dorsum (LND) is also present in between the intersegmental folds. It has its anterior attachment with the integument at the base of the dorsal internal lateral longitudinal muscles and its posterior attachment with the posterior intersegmental fold, beneath the insertion of the lateral internal oblique muscles. It remains connected to the dorsal nerve by a fine terminal branch of the dorsal nerve. the ventral nerve: The ventral nerve (VN) arises from the ganglion about the middle of its lateral margin and passes obliquely posteriorly over the ventral external oblique muscles. It gives rise to three branches, amongst which, the first branch (IV) extends two branches (a and b) to innervate the ventral external and lateral oblique muscles, tergosternal muscle and tracheae. The second branch (2V) bifurcates so that one branch (a) innervates the meso- December, 1966] Bahadur and Srivastava: Lepidopteran Nerves (Prodenia) 173 thoracic leg, ventral external oblique muscle and the integument, and the other branch (b) innervates the mesothoracic leg. Very near to the second branch, originates the third branch (3V) which innervates the ventral external and internal oblique muscles and tracheae. The fourth branch (4V) directly runs into the mesothoracic leg to innervate its muscles. The main nerve itself passes posteriorly to innervate the ventral external and internal oblique muscles. the transverse nerve i The unpaired median nerve (MN) of the mesothoracic segment arises from the fused intersegmental ganglionic connectives at the point where the connectives separate, that is, a very short distance posterior to the prothoracic ganglion. The median nerve travels about two thirds of the distance in between the interganglionic connectives and then gives off a pair of transverse nerves (TN). The transverse nerve receives three connective branches from the dorsal nerve as already stated. The two lateral connectives lie on the two sides of the lateral longitudinal tracheal trunk. The main trans- verse nerve which runs over this tracheal trunk, above the dorsal internal lateral muscles, terminates into the aorta to innervate it. postmedian nerve: A pair of fine nerves which may be designated as the postmedian nerves (PMN) arise from the transverse nerves, very near to the point of bifurcation of the median nerve. They proceed posteriorly and meet the interganglionic connectives at two points, a little above the mesothoracic ganglion. The two nerves communicate with each other by a pair of very short connectives. NERVES OF THE METATHORACIC GANGLION (Fig. 3) the dorsal nerve: The dorsal nerve (DN) arises from the outer margin of the interganglionic connective and passes over the ventral external oblique muscles for a short distance and then penetrates to run beneath the ventral internal lateral muscles. The first branch (ID) of the dorsal nerve extends to meet the transverse nerve but in addition gives rise to a branch (a) which subdivides into a number of minute branches to innervate the ventral external and internal oblique muscles and the ventral internal lateral longitudinal mus- cles. The main dorsal nerve subsequently sends off another connective branch (2D) which proceeds antero-laterally and fuses with the transverse nerve. But just near the fusion point, the branch 2D extends a branch anteriorly which innervates certain sternopleural muscles and tracheae. In addition, the branch 2D gives rise to two small branches (a and b) which innervate the lateral internal oblique and ventral external oblique muscles respectively. The third connective branch (3D) of the dorsal nerve again fuses with the trans- verse nerve. A side branch (a) from this nerve bifurcates to innervate the sternopleural muscles, tracheae and integument. The main dorsal nerve then passes towards the mid-dorsal region and extends the fourth branch (4D) which passes anteriorly and then curves sharply 174 New York Entomological Society [Vol. LXXIV Fig. 3. Diagram of the metathoracic nerve pattern of the larva of P. litura. postero-dorsally, giving origin to a number of minute branches at intervals. Its first minute branch (a) innervates the paratergal muscle, the second, third and fourth branches (b, c, d) innervate the lateral internal oblique muscles and the dorsal internal lateral longitudinal muscle. The other branches inner- vate the various median and lateral longitudinal muscles of the dorsal region. The fifth branch (5D) of the dorsal nerve is short and fuses with the longi- tudinal nerve of the dorsum (LND) at the point x. The main dorsal nerve by now becomes extremely thin and fine and bifurcates into two branches which innervate the integument, dorsal internal median and dorsal external muscles. the ventral nerve: This nerve (VN) leaves the ganglion from the middle of its lateral margin and passes obliquely posteriorly beneath the crossed diagonal muscles and over the ventral external oblique muscles. It is com- December, 1966] Bahadur and Srivastava: Lepidopteran Nerves (Prodenia) 175 paratively shorter and extends into the metathoracic leg to innervate its muscles. It, however, gives rise to a number of branches during its course. The first branch (IV) runs below the ventral internal lateral muscles and gives rise to a small branch which immediately subdivides into four minute branches. Among these, the first three branches (a, b, c) innervate the integument whereas the other branch (d) passes over the lateral longitudinal tracheal trunk and bi- furcates to innervate the sternopleural muscle and the integument of the dorsal region. The main branch (IV) extends further ahead over the lateral longi- tudinal tracheal trunk and gives rise to several small branches which innervate the lateral external oblique and tergosternal muscles and the integument. The second branch (2V) of the ventral nerve runs anteriorly and bifurcates. Whereas one branch (a) innervates the metathoracic leg, the other (b) inner- vates the integument. The third branch (3V) arises near the origin of the second branch and while it proceeds laterally, it gives rise to a number of minute branches which innervate the ventral external and internal oblique muscles, sternopleural and tergopleural muscles and the integument of that region. The fourth branch (4V) penetrates into the leg to innervate it whereas the fifth branch (5V) divides into three branches. The first branch (a) inner- vates the leg muscles, the second (b) innervates the ventral internal median muscle, the integument and tracheae and the last branch (c) runs posteriorly to innervate the two ventral external oblique muscles. the transverse nerve: A pair of transverse nerves (TN) arises by the bifurcation of a median nerve. Each transverse nerve passes over the dorsal and ventral internal longitudinal muscles and receives three connections from the dorsal nerve as already stated. During its course, it gives rise to two very minute branches (a and b) which innervate the tracheae. Before terminating, the transverse nerve divides into three minute branches. The first two branches (c and d) pass inwards to supply certain small muscles, integument and tracheae whereas the third branch (e) passes towards the mid-dorsal region to innervate the aorta. DISCUSSION In the Prodenia larva, the thorax bears three distinct ganglia. The dorsal nerve arises directly from the prothoracic ganglion but in the meso and meta- thoracic segments it arises from the interganglionic connectives. Nesbitt (1941) in Orthoptera described an anterior ganglionic connective extending from one ganglion to the other. He has shown that the anterior part of the nerve may adhere to or even become incorporated in the adjoining interganglionic con- nective. This nerve has been termed as intercalary nerve by Pipa and Cook (1959) and Matsuda (1956) whereas dorsal nerve connective or anterior gan- glionic connective by Schmitt (1959). In Dissosteira, Acheta, Periplaneta and Orchelimum, Schmitt found varying degrees of adherence to or fuse with the 176 New York Entomological Society [Vol. LXXIV adjoining interganglionic connective so that with the adherence of the anterior ganglionic connective too, the dorsal nerve seems to emerge from the connective. This condition is seen in the metathorax of Orchelimum. In Prodenia larva also the dorsal nerves in the meso and metathorax arise from the interganglionic connectives, as already stated and as such the case appears to be parallel with that of Orchelimum . In Chauliodes (Neuroptera) also Maki (1936) found a similar condition and Schmitt thinks that it is possible that the dorsal nerve in this insect is simply adhering to the nerve cord and does not lack an anterior connective but the resemblance with Orchelimum suggests that a dorsal nerve connective occurs in Chauliodes too. It must, therefore, be taken as fused. In the prothoracic segment of the Prodenia larva, however, the dorsal nerve arises directly from the ganglion. It appears that the proximal part of the dorsal nerve in this case has not fused with the interganglionic connective but the anterior ganglionic connective has fused with the interganglionic connective. Among other lepidopterans, Weber (1954) found an anterior connective of the dorsal nerve but Du Porte (1915) did not observe it in Sphida and has shown the dorsal nerve to arise from the interganglionic connective similar to the condition seen in Prodenia larva. The median or unpaired nerve in the larva is a short nerve which bifurcates to form two transverse nerves of a segment. Exceptions were noted in the larva of Papilio by Hillemann (1933) who figured a continuous median nerve between the second and third thoracic ganglia, and the same was observed by Marquardt (1939) in Carausius. The origin of the prothoracic median and transverse nerves is different in the larva so that the latter have been named as the subconnective nerves. Each nerve meets the dorsal nerve through a plexus and hence on this basis it can be conveniently presumed that these nerves are actually the transverse nerves. Peterson (1912) also reported the presence of subconnective nerves in the larva of tomato worm. In Prodenia larva, fusion of the transverse nerve from the prothoracic ganglion with meso- thoracic dorsal nerve and of the transverse nerve from the mesothoracic ganglion with the metathoracic dorsal nerve has been observed. Such fusions have also been reported in Chauliodes, Aqulla, Perla, Carausius, Blattella, Periplaneta, Telea, Dissosteira and Papilio. Most writers have designated the transverse nerve by that name but Pipa and Cook (1959) identified it simply as “nerve 8.” Whereas in the prothoracic segment, the subconnective nerve joins the dorsal nerve through a plexus, in the meso and metathoracic segments the connection between the transverse and dorsal nerves is maintained by three connectives. Du Porte (1915) also reported three such connections in Sphida larva but Swaine (1920) observed two or three in Sthenopis larva. Hillemann (1933) found two connections in the Papilio larva. Nothing appears to be known regarding the function of the axons which presumably pass from the transverse or subconnective nerve to the prothoracic dorsal nerve. Wittig (1955) found December, 1966] Bahadur and Srivastava: Lepidopteran Nerves (Prodenia) 177 that in Perla these transverse nerves pass to certain small dorsal longitudinal muscles and have no contact with the dorsal nerves but Schmitt (1959) re- ported that in Dissosteira the transverse nerves join the second cervical nerves and that somewhat distad of the junction, there is a connection from the second cervical nerve to the prothoracic dorsal nerve by means of which presumably axons from the transverse nerve could reach the same destination as in Carausius and Chauliodes and perhaps Perla also. In Prodenia larva the case too appears to be similar though the connection between the subconnective and prothoracic dorsal nerves is through a plexus. In the larva the transverse nerve actually terminates in the dorsal vessel and the same innervation was described by Libby (1961) in the abdomen of Hyalophora. In the abdomen of Orthoptera, how- ever, Alexandrovicz (1913), Nesbitt (1941) and Schmitt (1959) found the innervation of the dorsal vessel from the dorsal nerves. The question whether there is a real difference in the innervation of the dorsal vessel in Lepidoptera and Orthoptera or the same axons are involved but follow different nerve paths appear problematic. The fact that the transverse nerve directly innervates the dorsal vessel but receives three connective branches from the dorsal nerve suggests that the dorsal vessel is innervated not only by the axons of the trans- verse nerve exclusively but by those of the dorsal nerve, also. This interpreta- tion reconciles the views of orthopteran and lepidopteran workers. In the larva of Prodenia , there is only one pair of thoracic spiracles lying in the prothorax. Each is innervated by a branch from the second connective joining the transverse and dorsal nerves of the mesothoracic segment. Case (1957) has shown that in the cockroach the axons to the spiracular muscle actually issue from the transverse nerve and the same was demonstrated by Hoyle (1959) in Schistocerca gregaria. On the basis of their findings it can be presumed that here also the axons from the transverse nerve travel into the connective branch and then into the spiracular muscle through another minor branch. The spiracular muscle also receives axons from the dorsal nerve, travelling by the same path. It has already been pointed out that the thoracic spiracle is innervated by a branch from the connective 2D so that it may be considered to be homologous to the A-B connection present in the abdomen in Orthoptera, Plecoptera (Schmitt 1954, 1962, 1963), Lepidoptera (Libby 1959), and Neuroptera (Maki 1936). The ventral nerve usually innervates the leg muscles, the various ventral oblique muscles and the integument. In the meso and metathorax of Prodenia larva there is a pair of ventral nerves in each segment. In other lepidopterous larvae also the same arrangement and number has been observed (Swaine, 1920; Hillemann, 1933; Du Porte, 1915 and Peterson, 1912). In the pro- thorax of the larva, the ventral nerve not only innervates the leg muscles and oblique muscles but also the muscles lying at the base of the head. In Dis- 178 New York Entomological Society [Vol. LXXIV sostera, Schmitt found a prothoracic nerve to join one of the cervical or ventral nerves. He further considered this prothoracic nerve to be the counter part of the dorsal nerve of the meso and metathorax. In the present case though there is no connection between the prothoracic nerve and the ventral nerve, yet there exists a connection (6V) between the dorsal and the ventral nerves. On the basis of Schmitt’s interpretation, it may be concluded that it is through this connective that the axons from the dorsal nerve travel to the muscles at the base of the head. The concept that in ancestral insect, there was a common ancestral pattern of musculature as well as innervation in each segment of the body, gets support in the present study that the nerve patterns in thorax as well as in abdomen (unpublished) of Prodenia larva are practically identical especially with refer- ence to the 2D or A-B connection. This connection has been described in the abdomen of widely separated orders of insects like Orthoptera, Plecoptera (Schmitt 1954, 1962, 1963), Neuroptera (Maki, 1936), and Lepidoptera (Libby, 1959). The presence of this connection in different insect groups suggests the existence of a basic segmental nerve plan. SUMMARY The nerves of the thoracic segments of the larva of Prodenia litura have been described in detail. The thorax bears three distinct ganglia, each giving rise to three pairs of nerves which are dorsal, ventral and transverse. The dorsal nerve mainly innervates the dorsal muscles whereas the ventral nerve innervates the leg and ventral muscles. The transverse nerve mainly supplies the dorsal vessel. The dorsal nerve of the prothoracic ganglion remains con- nected by a plexus to the subconnective nerve which has been considered to be a transverse nerve. In the other two segments, the dorsal nerve fuses with the transverse nerve at three points by means of three connectives. In con- trast to this, the ventral nerve does not fuse with the transverse nerve. The spiracular muscles are innervated from the connective lying in between the dorsal and transverse nerves. The pattern of the nerves is the same as found in other lepidopterous and orthopterous insects and that supports the concept that a basic segmental nerve pattern exists within the insects. Acknowledgement The authors thank Professor H. Swarup, Head of the School of Studies in Zoology, Vikram University, Ujjain for providing all necessary facilities during the course of this work. Literature Cited Alexandrovicz, J. S. 1913. The innervation of the heart of the cockroach. J. Comp. Neurol, 41: 291-309. Case, J. F. 1957. The median nerves and cockroach spiracular function. J. Insect Physiol., 1: 85-94. December, 1966] Bahadur and Srivastava: Lepidopteran Nerves (Prodenia) 179 Du Porte, E. M. 1915. On the nervous system of the larva of Sphida obliqua. Trans. R. Soc. Canada, 8: 225-253. Hillemann, H. H. 1933. Contributions to the morphology of the nervous system of the mature larva of Papilio polyxenes. Ann. Ent. Soc. Amer., 26: 575-583. Hoyle, G. 1959. The neuromuscular mechanism of an insect spiracular muscle. J. Insect. Physiol., 3: 378-394. Libby, J. L. 1959. The nervous system of certain abdominal segments of the Cecropia larva. Ann. Ent. Soc. Amer., 52: 469-480. . 1961. The nervous system of certain abdominal segments and the male re- productive system and genitalia of Hyalophora cecropia. Ann. Ent. Soc. Amer., 54: 887-896. Lyonet, P. 1762. Traite anatomique de la chenille qui ronge le bois de saule (La Haye, Amsterdam), 616 pp. Maki, T. 1936. Studies on the skeletal structure, muscuLature and nervous system of the alderfly, Chauliodes formosanus. Mem. Fac. Sci. & Agric., Taihoku Imp. Univ., 16(3) : 117-243. Marquardt, F. 1939. Beitrage zur Anatomie der Muskulatur and periphern Nerven von Carausius ( Dixippus ) morosus. Br. Zool. Jahr. Anat., 66: 63-128. Matsuda, R. 1956. The comparative morphology of the thorax of two species of insects. Microentomology, 21 : 1-63. Nesbitt, H. H. J. 1941. A comparative morphological study of the nervous system of the Orthoptera and related orders. Ann. Ent. Soc. Amer., 34: 51-81. Peterson, A. 1912. Anatomy of the tomato worm larva, Protoparce Carolina. Ann. Ent. Soc. Amer., 5: 246-269. Pipa, R. L. and Cook, E. F. 1959. Studies on the hexapod nervous system. I. The pe- ripheral distribution of the thoracic nerves of the adult cockroach, Periplaneta ameri- cana. Ann. Ent. Soc. Amer., 52: 695-710 Ruckes, H. 1919. Notes on the male genital system in certain Lepidoptera. Ann. Ent. Soc. Amer., 12: 192-209. Schmitt, J. B. 1954. The nervous system of the pregenital abdominal segments of some Orthoptera. Ann. Ent. Soc. Amer., 47: 677-682. . 1959. The cervicothoracic nervous system of a grasshopper. Smithsonian Inst. Pubis. Misc. Collections, 137: 307-329. Swaine, J. M. 1920. The nervous system of the larva of Sthenopis thule. Can. Entomol- ogist, 52: 275-283. Weber, H. 1954. Grundriss der Insektenkunde, Stuttgart, 428 pp. Wittig, G. 1955. Untersuchungen am thorax von Perla abdominalis (Larve und Imago). Zool. Jahrb. Anat., 74: 491-570. Received for Publication August 10, 1966 KEY TO ABBREVIATIONS DN = Dorsal nerve, LND = Longitudinal nerve of the dorsum, MN = Median nerve, PMN = Post median nerve, px = plexus, SG = Suboesophageal ganglion, SN = Subcon- nective nerve, TN = Transverse nerve, Th.G = Thoracic ganglion, VN = Ventral nerve. 180 New York Entomological Society [Vol. LXXIV Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XIII1 Charles P. Alexander Amherst, Massachusetts Abstract: Six new species of Eriopterine crane flies are described, these being Neolimnophila citribasis n. sp., from Assam; N. daedalea n. sp., Sikkim; Lipsothrix decurvata n. sp., Sik- kim; Styringomyia subobseura n. sp., Assam; S. tarsatra n. sp., Nepal; and Toxorhina ( Ceratocheilus ) tuberifera n. sp., Sikkim. Part XII of this series of papers was published in the Journal of the New York Entomological Society, 74: 66-71, 1966. As before, the materials dis- cussed were collected by Dr. Fernand Schmid and Dr. Edward I. Coher, to whom my sincere thanks are extended. Neolimnophila citribasis n. sp. General coloration of body brown, the praescutum with four darker brown stripes, the intermediate pair narrowly separated; antennae 16-segmented ; wings brownish yellow, the basal third, including the veins, clear orange-yellow, narrow brown seams over cord and outer veins; R2 about twice its length before fork of RM. female: Length about 7 mm; wing 7.8 mm; antenna about 1.4 mm. Rostrum dark brown; palpi black. Antennae 16-segmented, black, the scape more pruinose; proximal two flagellar segments barely connate, the separating suture narrow but complete, outer segments progressively more slender, the outer pair shorter; longest verticils subequal to the segments. Head brownish gray; vestiture black, from small dark punctures. Pronotum dark brown. Mesonotal praescutum yellowish brown with four dark brown stripes, the intermediate pair separated only on posterior two-thirds; pseudosutural foveae and tuberculate pits black; posterior sclerites of notum brownish black. Pleura dark gray, dorsopleural membrane brown. Halteres elongate, clear light yellow. Legs with coxae black, gray pruinose; trochanters obscure yellow; femora brownish black, bases yellowed, broadly on the posterior pair, remainder of legs brownish black; tibial spur of fore leg lacking, present on hind pair (middle legs lacking). Wings brownish yellow, the basal fifth clear orange yellow, including the veins; narrow brown seams over cord, outer end of cell 1st M2, origin of Rs, and the outer forks, more diffuse and paler on Cu and the outer veins; veins light brown, the yellow bases extended outwardly to include all of Sc and less evidently on other primary veins. Venation: Sci ending opposite fork of Rs; R» about twice its length before the fork of R:i+i; origin of vein Rt angulated and short-spurred; cell M\ subequal to its petiole; m-cu just beyond the fork of M. Abdomen black, the genital segment intensely so ; valves of ovipositor horn yellow. Holotype $, Jhum La, Kameng, North East Frontier Agency, Assam, 7,800 feet, May 13, 1961 (Schmid). In its general appearance Neolimnophila citribasis is most similar to N. 1 Contribution from the Entomological Laboratory, University of Massachusetts. December, 1966] Alexander: Himalayan Crane Flies, XIII 181 daedalea n. sp., of Sikkim, being readily told by the extensive brightening of the wing base and the more restricted darkened pattern of the disk. The lack of a distinct flagellar fusion-segment is especially noteworthy. Both N . daedalea and N . fuscinervis Edwards, of Yunnan, have the basal seg- ment of the flagellum elongate, resulting from four fused segments, with ten free segments beyond. Neolimnophila daedalea n. sp. General coloration of thorax blackened, the praescutum with four darker stripes, the intermediate pair vaguely separated; wings light yellow, most of the veins heavily seamed with brown, cells C and Sc conspicuously brownish yellow; a darkened cloud in outer half of cell R behind Rs. male: Length about 5.5 mm; wing 8 mm; antenna about 1.2 mm. female: Length about 6-6.5 mm; wing 8-8.2 mm; antenna about 1.4 mm. Rostrum and palpi black. Antennae black, scape pruinose; fusion-segment of flagellum involving four segments, with ten free segments beyond. Head brownish gray; anterior vertex broad. Pronotum blackish gray. Mesonotal praescutum with four blackened stripes, the inter- mediate pair only vaguely separated, the lateral stripes poorly indicated, lateral margins of segment light gray ; posterior sclerites of notum black, very sparsely pruinose. Pleura gray. Halteres light yellow. Legs with coxae black, pruinose; trochanters brown; remainder of legs brownish black, femoral bases very narrowly paler. Wings strongly light yellow, the prearcular field bright yellow; a heavy brown pattern over the cord, outer end of cell 1st M-2 and vein Cu, narrower but still conspicuous on veins beyond cord with the excep- tion of M i+2, Mi and Ms+i; no darkenings on M or 1st A; a conspicuous marking in outer half of cell R behind Rs; cells C and Sc brownish yellow. Venation: R2 some distance before fork of Rs+i, subequal to R 3; position of r-m slightly variable, in cases at or just before the fork of Rs. Abdomen dull black. Valves of ovipositor long and straight, tips of cerci with coarse yellow setae. Holotype 2, Kalep, Sikkim, in Rhododendron association, 12,100 feet, June 18, 1959 (Schmid). Allotype, 8, Yumtang, Sikkim, in Rhododendron associa- tion, 12,140 feet, June 27, 1959. Paratypes, 38 8, with the allotype; 1 $, Chachu, Sikkim, 11,500 feet, June 29, 1959 (Schmid). Other Himalayan species include Neolimnophila genitalis (Brunetti), with unpatterned wings, together with N. bijusca Alexander and N . citribasis n. sp., previously described in the present report. In the higher mountains of western China still other species are found, including N. fuscinervis Edwards, N. perreducta Alexander and N. picturata Alexander, all with the details of wing pattern and venation distinct. Lipsothrix decurvata n. sp. Pronotum and anterior end of praescutum brownish black, the remainder of the prae- scutal stripes paler, pleura light yellow ; femora yellow, tips conspicuously brownish black ; wings faintly darkened, prearcular and costal fields more brownish yellow; Rs relatively long, nearly twice R2+ 3+4, vein R± very strongly decurved outwardly, its tip at or beyond the wing apex, cell 1st M2 short-rectangular. 182 New York Entomological Society [Vol. LXXIV male: Length about 7-7.2 mm; wing 7. 5-7. 8 mm; antenna about 2. 7-2. 8 mm. Rostrum brownish yellow; palpi black. Antennae relatively long, as shown by the measurements; scape and pedicel brownish yellow, flagellum brownish black; segments long- subcylindrical, verticils short and sparse. Head light brown. Cervical region and pronotum brownish black. Mesonotal praescutum extensively obscure yellow, with a light brown central stripe, anterior half more brownish black, this being a continuation of the pronotal darkening, region of the suture yellowed; scutal lobes and posterior sclerites darkened, including the pleurotergite. Pleura light yellow. Halteres with stem pale, knob darkened. Legs with coxae and trochanters yellow; femora yellow, tips conspicuously brownish black; tibiae obscure yellow, tips more narrowly darkened; tarsi obscure yellow; claws long, with a major subbasal spine and two or three smaller more proximal denticles. Wings faintly darkened, prearcular and costal regions more brownish yellow, stigma still darker; veins brown, more yellowish brown in the brightened fields. Macrotrichia of veins very long. Venation: Sc i ending just beyond fork of the long Rs, Sc2 near its tip; R2 faint, subequal to or shorter than R1+2) vein Ri very strongly decurved outwardly, ending at or beyond the wing tip, cell at margin slightly more extensive than cell R>; cell 1st M* short-rectangular, less than one-half the veins beyond it; m-cu about one-third its length beyond the fork of M, in cases close to the fork. Abdomen including hypopygium, dark brown. Male hypopygium with the interbase slender; phallosome strongly developed, about as in malla. holotype S , Chateng, Sikkim, 8,700 feet, June 12, 1959 (Schmidt). Paratopo- types, 4 8 S , on two pins. Lipsothrix decurvata is close to L. malla Alexander, of Nepal, differing in details of body coloration and in the venation, especially cell 1st Mo and the radial field, including the more decurved vein R±. Styringomyia subobscura n. sp. Allied to obscurer, general coloration of body black; legs blackened, middle and hind femora each with a narrow yellow subterminal ring, posterior tarsi whitened; wings slightly suffused, virtually unpatterned; male hypopygium with a long sinuous rodlike spine near base of inner arm of dististyle; apex of phallosome bilobed. male: Length about 6.5 mm; wing 4.6. female: Length about 6 mm; wing 4.5. Rostrum and palpi black. Antennae with proximal five or six segments black, the outer ones brownish yellow; flagellar segments oval. Head brownish black. Thorax black, sparsely gray pruinose to appear dull; central region of sternum some- what paler. Halteres with stem dark brown, knob brownish black. Legs with coxae light brown; trochanters brownish yellow; remainder of fore legs uniformly blackened; middle femora restrictedly obscure yellow at base, remainder brownish black with a narrow obscure yellow ring some distance before tip, tibiae brownish black, the extreme base pale, tarsi brownish black, the extreme bases vaguely paler ; posterior legs chiefly black, femur with a conspicuous pale yellow subterminal ring, the darkened tip nearly three times as extensive, tibiae brownish black, tarsi whitened, the extreme tips of the individual segments pale brown, terminal segment uniformly darkened. Wings beyond cord with a slight darkened suffusion, basal cells more whitened; a dusky seam along vein Cu involving both cells; veins dark brown. Venation: Anterior branch of Rs more nearly erect than in obscura. Abdomen black. Male hypopygium generally as in obscura, differing in all details, especially of the dististyle and phallosome. Outer rod of dististyle without basal setae; outer arm large, its surface with numerous scattered setae; margin with an unbroken December, 1966] Alexander: Himalayan Crane Flies, XIII 183 comb of strong spinoid setae, those at either end of row slightly longer; inner arm of style more slender, with two terminal combs, two strong similar spines near base, and a long sinuous rodlike spine on outer margin near base ; origin of dististyle with a blackened rod, its apex dilated into a head. Phallosome on either side with a recurved or pendant lobe, the obtuse apex blackened; in obscura this represented by a single powerful terminal spine. holotype 8, Chapai, Kameng, North East Frontier Agency, Assam, 700 feet, February 26, 1961 (Schmid). Allotype 2, Bhairabkunda, Kameng, 700-1,000 feet, March 5, 1961 (Schmid). The closest relatives of the present fly are Styringomyia obscura Brunetti and S. schmidiana Alexander, both with the hypopygial structure quite distinct. The male hypopygium of obscura has been described and figured by the writer (Philippine Jour. Sei. 86: 447-448, pi. 4, fig. 56; 1957). Styringomyia tarsatra n. sp. Size small (wing 4.5 mm or less) ; general coloration of mesonotum dark gray and black, ventral half of thoracic pleura abruptly yellow ; halteres black ; femora black, bases and a narrow subterminal ring yellow, all tibiae and tarsi black ; wings with a weak brownish tinge, the basal third more whitened; abdomen black; male hypopygium with the modified sternal setae apical in position ; outer lobe of basistyle with a single modified seta ; inter- mediate and inner arms of dististyle with rows of blackened pegs; phallosome unusually small and inconspicuous, the blackened apex rounded. male: Length about 6. 2-6. 5 mm; wing 4-4.5 mm. female: Length about 6 mm; wing 4.2 mm. Rostrum and palpi black. Antennae with scape, pedicel and proximal flagellar segments black, intermediate segments paler, the outer ones again blackened; pedicel enlarged, flagellar segments oval. Head dark brown to brownish black, sparsely pruinose. Pronotum dark brown, obscure yellow medially. Mesonotum gray, patterned with black including sublateral praescutal stripes and margins to the scutal lobes; scutellum with a central pale yellow spot. Pleura conspicuously blackened above, including the dorsopleural membrane, lower half abruptly yellow, including also the coxae of all legs. Halteres black. Legs with trochanters yellow, femora black, the bases restrictedly paler, with a narrow obscure yellow subterminal ring at about twice its length from the tip; tibiae and tarsi of all legs black. Wings with a weak brownish tinge, the basal third or more whitened; veins brown. Venation: Anterior branch of Rs oblique; cell 2nd M 2 narrowly to more broadly sessile. Abdomen black, hypopygium brownish black. Male hypopygium with the tergite nar- rowed outwardly, apical lobe provided with dense retrorse setae ; sternite long and narrow, the two modified setae terminal, placed at outer apical angles of sternal lobe, surface microscopically setulose. Basistyle with a single modified seta, subequal in length to its basal tubercle. Dististyle with outer arm bearing a single weak seta at near one-third the length; intermediate and inner arms provided with blackened pegs; inner arm with a slender pale rod on outer margin, the inner edge near base with a group of about 10 to 12 very long setae. Phallosome unusually small and inconspicuous, the outer end rounded and blackened. holotype 8, Parewavir, Nepal, March 28, 1957 (Coher). Allotype 2, Am- lekhgang, Nepal, 520 meters, July 26, 1957 (Coher). Paratopotypes, 6 8 8, with the type, March 15-28, 1957 (Coher). 184 New York Entomological Society [Vol. LXXIV Other somewhat similar regional species include Styringomyia obscura Bru- netti, S. schmidiana Alexander, and S. subobscura n. sp., from which the present fly differs in the small size, details of coloration, including the uni- formly blackened tarsi of all legs, and in the details of hypopygial structure, including particularly the dististyle and phallosome. Toxorhina ( Ceratocheilus ) tuberifera n. sp. General coloration of head and thorax gray, the praescutum with three virtually con- fluent brown stripes; halteres and legs black, the femoral bases restrictedly yellow; wings subhayaline, base more yellowed, anterior branch of Rs sinuous, cell Rx narrowed at margin ; abdomen brownish black ; male hypopygium with a strong tubercle near proximal end of basistyle ; dististyle terminal, the marginal tubercle small ; arms of aedeagus very short. male: Length, excluding rostrum, about 5 mm; wing 5 mm; rostrum alone about 3 mm. female: Length, excluding rostrum, about 5.5 mm; wing 5 mm; rostrum about 3 mm. Rostrum black, more than one-half the length of wing. Antennae black. Head black, sparsely pruinose, without a corniculus ; anterior vertex relatively narrow, slightly wider than the diameter of the scape. Cervical region and pronotum blackened. Mesonotal praescutum with three virtually confluent brown stripes, the median extension darker in front, lateral praescutal borders gray ; posterior sclerites of notum black, gray pruinose, scutal lobes more infuscated. Pleura black, sparsely pruinose to appear plumbeous. Halteres black throughout. Legs with coxae brownish black, trochanters brownish yellow; remainder of legs black, femoral bases re- strictedly yellowed. Wings subhyaline, the base more yellowed; veins brown, those at wing base more brownish yellow. Certain veins beyond cord with sparse trichia, including both sections of R5 and distal section of M i+2; a single trichium near outer end of vein Ms. Venation: Sci ending just beyond origin of Rs, Sc» before the origin; anterior branch of Rs sinuous but more erect than in mesorhyncha, cell R\ narrowed at margin; Rs nearly as long as basal section of R5; m-cu before fork of M. In the allotype female both wings have cell Ms open by the atrophy of m. Abdomen, including hypopygium, brownish black. Male hypopygium with a strong tubercle on mesal face of basistyle near proximal end, provided with several strong black setae. Dististyle single, terminal in position, extended into a long slender beak bearing a low lateral flange, outer margin shortly before midlength with a small tubercle. Interbases appearing as narrow blades. Arms of aedeagus unusually short, less than the distance separating them at bases. holotype $ , Lathong, Sikkim, 6,560 feet, July 26, 1959 (Schmid). Allotopo- type, 9. The closest relative is Toxorhina ( Ceratocheilus ) mesorhyncha Alexander which differs in the venation of the radial field and in the hypopygial struc- ture, especially the dististyle and aedeagus. Received for Publication August 5, 1966 December, 1966] Klots: Amblyscirtes samoset Larva 185 The Larva of Amblyscirtes samoset (Scudder) (Lepidoptera: Hesperiidae) Alexander B. Klots1 Abstract: The mature larva is described, figured and compared with that of the largely sympatric A. vialis (W. H. Edwards). Some larval characters of the genus Amblyscirtes Scudder are discussed. A female Amblyscirtes samoset (Scudder) was observed, by Cyril dos Passos and the writer, ovipositing on Poa pratensis L. near West Bridgewater, Vermont, on 9 June 1956. A single egg was found, from which the larva was reared to maturity by Dr. dos Passos. The larva was then photographed, studied and preserved by the writer. It is in the American Museum of Natural History. The larva of this species is relatively unknown, almost the only published data on it being those of Scudder (1889, p. 1589-1592, pi. 77, fig. 29). Scudder, however, merely copied one of Abbot’s pictures; and his description and figure are quite inadequate. DESCRIPTION OE MATURE LARVA length at rest: 22.5 mm. Head rounded and only very slightly emarginate dorsally at epicranial suture (Fig. 3), from anterior aspect as wide as high; and with very little taper dorsad; covered with fine, ridgelike reticulations; very sparsely and finely setose. Face (including the central triangular sclerite, the narrow sclerites bordering this laterally, and the anterior edges of the epicrania2) dark brown, forming a triangle that narrows toward vertex, and behind vertex joins the dark posterior region of the head; laterad of this on either side a broad, very pale brown band running dorsad almost to vertex (Figs. 1 & 2) ; posterad of this on either side a dark brown band, ventrally including the anterior 4 stemmata, running dorsad to vertex; posterad of this on either side a broad, pale band running dorsad almost to vertex; vertex and posterior region of head dark brown. Labrum shallowly emarginate. A strong, projecting, slightly recurved spine (here called the paraclypeal spine ) arising laterad of each ventro-lateral angle of clypeus, protruding forward and ventrad. Stemmata: Nos. 1-4 forming an anterior curving group; of these, 4 is the largest, 3 is slightly smaller than 4, 1 is slightly smaller than 3, and 2 is slightly smaller than 1. No. 6 is almost directly caudad of 4 and about as far from it as 1 is from 3. 1 Professor of Biology, The City College of New York; and Research Associate, American Museum of Natural History. 2 The nomenclature of the anterior surface of the head of lepidopterous larvae is some- what confused. Most recent systematists call the large, triangular, central sclerite the frons , the narrow sclerites along each side of this the adfrontals, and the transverse area ventrad of the so-called frons the clypeus. However, as shown by Snodgrass (1935, p. 121, fig. 64) the triangular central sclerite is really the clypeus; most of the true frons is invaginated within the so-called epicranial suture dorsad of the true clypeus; and the narrow, lateral sclerites are the ventral remnants of the true frons, separated by the dorsad extension be- tween them of the true clypeus. Scudder (loc. cit ., I, p. 8) calls the whole complex the “facial triangle, or clypeus.” 186 New York Entomological Society [Vol. LXXIV Figs. 1 & 2. Mature larva, Amblyscirtes samoset (Scudder), from life. Fig. 1: lateral aspect. Fig. 2: dorsal aspect. No. 5 is ventrad and slightly cauaad of 6, just above and slightly caudad of base of antenna and directed ventrad. Prothoracic shield heavily sclerotized, black, shining, its ventral margins somewhat undulate. Prothoracic spiracle very large, broadly and sym- metrically oval. Ground color of body very pale whitish green with darker dull green markings (Figs. 1 & 2). Meso- and metathorax fairly thickly covered with short setae arising from circular, well-sclerotized bases ; remainder of body with shorter, much sparser setae, nearly all of which arise from almost unsclerotized bases. A distinct narrow, mid-dorsal, dark line from anterior end of mesothorax to posterior end of abdomen, weakening posteriorly; a more diffuse, dark, lateral, supraspiracular line from thorax to posterior end of abdomen, weakening posteriorly. A pale whitish, subspiracular line along the edge of a distinct, folded, lateral ridge from anterior edge of prothorax to posterior end of abdomen. Meso- thorax almost completely dark, metathorax lighter, abdominal segments progressively lighter. On the posterior part (somewhat more than half) of each abdominal segment are 4 or 5 very narrow, somewhat irregular transverse dark lines between which are transverse rows December, 1966] Klots: Amblyscirtes samoset Larva 187 Figs. 3 & 4. Head of mature larva, Amblyscirtes samoset (Scudder). Fig. 3: anterior aspect; the shading shows pigmentation, not contour. Fig. 4: lateral aspect, showing also prothoracic shield and spiracle. of dark dots; and on the anterior part (somewhat less than half) of each segment a number of dark dots, sometimes more or less in transverse rows, sometimes irregularly located. DISCUSSION Scudder’s description and figures of the larvae of A. vialis (loc. cit., p. 157 5— 1588, PI. 77, fig. 24 and PI. 80, figs. 46-50) show it differing from that of A. samoset in a number of features. A. vialis has the head narrower and more emarginate and tapering dorsally; the frontal triangle is more largely pale; on either side of it is a narrow, vertical dark stripe that does not run dorsad to join the other dark areas; and the body is paler and more thickly setose and lacks the dark middorsal line and most of the other dark markings of samoset. Scudder cites the dorsally tapering, emarginate head and the pro- truding paraclypeal spines as generic characters for Amblyscirtes. Heitzman (“1964” 1 1965] and 1965) has described and figured in detail the larvae of A. nysa W. H. Edwards and A. belli A. Freeman. Each has a dark middorsal line, a dorsally tapering, emarginate head, and a distinctive dark and light banded head pattern generically like those of A. vialis and samoset. Paraclypeal spines are figured for the larva of A. belli but not mentioned; but are not men- tioned or figured for the larva of A. nysa. In the latter case they may have been overlooked. The stemmata and prothoracic spiracle are not shown in detail. It seems probable that the paraclypeal spines and the banded head pattern may be regarded as characters for Amblyscirtes, but that the dorsally broader and non-emarginate head is peculiar to A. samoset. Details of the body pattern 188 New York Entomological Society [Vol. LXXIV and the surface sculpturing of the head may well prove to be characters of specific value when the larvae of more species are known. Literature Cited Heitzman, J. R. “1964” [1965]. The habits and life history of Amblyscirtes nysa (Hes- periidae) in Missouri. Jour. Res. Lep., 3: 154-156. . 1965. The life history of Amblyscirtes belli in Missouri. Jour. Res. Lep., 4: 75-78. Scudder, S. H. 1889. The butterflies of the eastern United States and Canada with special reference to New England. Cambridge, Massachusetts. Snodgrass, R. E. 1935. The principles of insect morphology. McGraw-Hill, New York. Received for Publication August 10, 1966 The Discovery of Additional Journals of Frank E. Watson In an obituary of Frank Edward Watson, 1877-1947, published in the Journal of the New York Entomological Society (1958, 66: 1-6) the finding of some of his loose-leaf journals covering the years 1904 in part, 1906-1910, 1911 in part, 1912-1913, 1915 and 1923-1925 was reported. During Watson’s last years he made his home with William Friedle of Ozone Park, Long Island, New York. With the death of William Friedle a few months ago his step nephew, Mr. Bruce Friedle, discovered a number of additional volumes of Watson’s journals that had not been delivered to the undersigned when he purchased Watson’s butterfly collection and library from Friedle after the former’s death. These additional journals have been kindly given by Mr. Friedle to the Department of Entomology of the American Museum of Natural History and are as follows: 1896-1905, 1914-1922, 1926-1931, 1934-1947. Thus the American Museum now has all of Watson’s diaries in the Department of Entomology with the exception of those covering the years 1932 and 1933. These must be assumed to have been lost. The Watson journals, as before observed, are extremely interesting and important as showing his activities in the field from day to day and in rearing Lepidoptera. Further details of these matters will be found on page 3 of the aforementioned paper. They also fix definitely his collecting localities which are only indicated on his specimens by code letters. Cyril F. dos Passos December, 19661 Manischewitz: Parasitic Mites of New Jersey 189 Studies on Parasitic Mites of New Jersey1 Jack R. Manischewitz Rutgers — The State University, New Brunswick, N.J. Abstract: A study of mites of the Trombiculidae, Myobiidae, Pyemotidae, Tetranychidae, and Acarinae collected from mammals in New Jersey included 26 recognized species and 3 probable new species. New records for the state and host and date-locality records are included. INTRODUCTION In view of the scarcity of New Jersey ectoparasite records, a survey was undertaken from 1951 to 1953 by the New Jersey Agricultural Experiment Station with cooperation from the New Jersey Department of Agriculture, and the New Jersey Division of Fish and Game. During this survey, about 4,000 mammals of twenty-nine species were collected. This paper summarizes information on the Trombiculidae, Myobiidae, Pyemotidae, Tetranychidae, and Acarinae which were taken from mammals and are new collection and/or host records for New Jersey. Most of the small mammals other than rats were collected with Sherman live traps or snap mousetraps. Rats were usually collected from municipal dumps by use of cyanide gas. All mites were mounted in Hoyer’s medium. RESULTS New records for the state are listed below together with information on hosts and comments on species where warranted. In the records abbreviations are as follows: L indicates larva, N indicates nymph, T indicates tritonymph, F indicates female, and M indicates male. Specimens without letter designations are adults. Numbers appearing after the word “plus” indicate specimens not mounted and not identified by the author, but which were thought at the time of mounting to be identical with those mounted. All mites of the same species found on the same day on the same host species in the same locality are dealt with as one record. TROMBICULIDAE Wharton and Fuller (1952) summarize much general information pertaining to the biology and ecology of chiggers. They also present keys to genera, and list all species and all references. Brennan and Jones (1959) present keys in- cluding all North American species of chiggers. 1 Paper of the Journal Series, New Jersey Agricultural Experiment Station. From a thesis submitted to the Graduate Faculty, Rutgers — The State University in partial fulfillment of the requirements for the M.S. degree. 190 New York Entomological Society [Vol. LXXIV Table 1. Mites and hosts found in the present study. <0 8 8 53 Co 8 q 8 *8 8 Co • 8 • V #co » CO Co V o CO 8 • 5S* <8 Co 8 CO • 5S* &0 CO p s rs* « 53 8 53 8 53 8 •p* -+8i Co 8 •^1 8 ►si O § 5S> *>>* ... / r iA //. K ( .As. /• A Os , • ' if- - ' . > \ / ^ JOURNAL of the NEW YORK ENTOMOLOGICAL SOCIETY The JOURNAL of the NEW YORK ENTOMOLOGICAL SOCIETY is de- voted to the advancement and dissemination of knowledge pertaining to insects and their related forms. INSTRUCTION TO AUTHORS CORRESPONDENCE Submit manuscript in duplicate (original and one car- bon) to Dr. L. W. Clausen, Editor, 115 West 68th Street, New York, N. Y. 10023. Send material by registered mail in flat form. 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Single copies, as issued — $2.00 each. ^ : ' Please make all checks, money-orders or drafts payable to the NEW YORK ENTOMOLOGICAL SOCIETY. ■\i : 7 r T „r / 4/11 / V. i/zO \ . rx x /\ kt’t i !1' in’ ■ y >6 1 \ Journal of the New York ENTOMOLOGICAL SOCIETY Devoted to Entomology in General VOLUME LXXV Published by the Society New York, N. Y. ALLEN PRESS, INC Lawrence, Kansas INDEX OF AUTHORS ALEXANDER, CHARLES P. Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XIV 24 ALEXANDER, CHARLES P. Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XV 183 BENTON, ALLEN H. A Case of Teratology in Monopsyllus vison (Baker) 31 BENTON, ALLEN H. Peromyscopsylla hamifer hamijer (Rothschild): an Addition to the Entomological Fauna of New York State 159 BERTHOLD, ROBERT, Jr. Behavior of the German Cockroach, Blatella germanica (L.), in Response to Surface Textures 148 DAWSON, R. W. New and Little Known Species of Serica (Coleoptera: Scarabaeidae) X 161 FORBES, JAMES The Male Genitalia and Terminal Gastral Segments of Two Species of the Primitive Ant Genus Myrmecia (Hymenoptera: Formicidae) 35 GRAY, P. H. H. Some Biometrics in Pieris and Colias (Lepidoptera: Pieridae) in Quebec and Nova Scotia 12 GERTSCH, WILLIS J. A New Liphistiid Spider from China (Araneae: Liphistiidae) 114 GUPTA, A. P. Further Studies on the Internal Anatomy of the Meloidae. III. The Digestive and Reproductive Systems as Bases for Tribal Designation of Pseudomeloe miniaceoniacidata (Blanchard) (Coleoptera: Meloidae) 93 HOFFMAN, RICHARD L. and LINDA S. KNIGHT A New Genus and Species of Spirostreptoid Millipeds from the Pacaraima Mountains, British Guiana 56 IVIE, WILTON Some Synonyms in American Spiders i 126 KELLY, ROBERT P. and DANIEL LUDWIG Distribution of Nitrogen During the Embryonic Development of the Mealworm, Tenebrio molitor Linnaeus 45 KELLY, RONALD J., DENNIS M. O’BRIAN, and FRANK F. KATZ The Incidence and Burden of Hymenolepis diminuta Cysticercoids as a Function of the Age of the Intermediate Host, Tribolium confusum 19 KISTNER, DAVID H. A Revision of the Termitophilous Tribe Termitodiscini (Coleop- tera: Staphylinidae) Part I. The Genus Termitodiscus Wasmann; its Systematics, Phyolgeny, and Behavior 204 KLOTS, ALEXANDER B. A Note on the Flight of Acrolophus morns (Grote) (Lepidoptera: Acrolophidae) 18 KLOTS, ALEXANDER B. The Adaptive Feeding Habit of a Pine Caterpillar 43 KLOTS, ALEXANDER B. Larval Dimorphism and Other Characters of Heterocam pa pulverea (Grote & Robinson) (Lepidoptera: Notodontidae) 62 KLOTS, ALEXANDER B. Two New Species of Crambus Fabricius from Western North America (Lepidoptera: Pyralididae) _ 154 iii LEONARD, MORTIMER D. Further Records of New Jersey Aphids (Homoptera: Aphididae) 77 MULLER, JOSEPH Melanism in New Jersey Cat ocala Schrank (Lepidoptera: Noctuidae) 195 OBRAZTSOV, NICHOLAS S. Genera Tortricoidarum Check List of Genera and Sub- genera Belonging to the Families Tortricidae (Ceracidae, Chlidanotidae, Schoenotenidae and Olethreutidae Included) and Phaloniidae 2 OBRAZTSOV, NICHOLAS S. Some Apocryphal Species of the Tortricinae (Lepidop- tera: Tortricidae) 34 PECHUMAN, L. L. Observations on the Behavior of the Bee Anthidium manicatum (L.) 68 POWELL, JERRY A. Apomyelois bistriatella : A Moth Which Feeds in an Ascomycete Fungus (Lepidoptera: Pyralidae) 190 RINDGE, FREDERICK H. A New Species of Nepytia from the Southern Rocky Mountains (Lepidoptera: Geometridae) 74 ROUSELL, P. G. Activities of Respiratory Enzymes During the Metamorphosis of the Face Fly, Musca autumnalis De Geer 119 ROZEN, JEROME G., Jr. The Immature Instars of the Cleptoparasitic Genus Dioxys (Hymenoptera: Megachilidae) 236 ROZEN, JEROME G., Jr. and MARJORIE S. FAVREAU Biological Notes on Dioxys pomonae pomonae and on its Host, Osmia nigrobarbata (Hymenoptera: Megachilidae) 197 TORCHIO, PHILIP F., JEROME G. ROZEN, Jr., GEORGE E. BOHART, and MAR- JORIE S. FAVREAU Biology of Dufourea and of its Cleptoparasite, Neopasites (Hymenoptera: Apoidea) 132 YOUNG, ALLEN M. Observations of Epicordidia princeps (Hagen) (Odonata: Corduliidae) at a Light 179 BOOK REVIEWS BATRA, SUZANNE W. T. Insect Behaviour. Symposium No. 3, Royal Entomological Society (P. T. Haskell, ed.) 100 HAGMANN, LYLE E. Handbook of the Mosquitoes of North America by Robert Matheson 147 TREAT, A. E. The New Field Book of Freshwater Life by Elsie B. Klots 29 WYGODZINSKY, PEDRO Monograph of Cimicidae by Robert L. Usinger 30 PROCEEDINGS of the NEW YORK ENTOMOLOGICAL SOCIETY 101, 249 NEW MEMBERS 110 IV ' 7 Vol. LXXV 'Y IK I MARCH 1967 No. 1 T { m n ' , . | . ,.„ V- Vy> ;;- > W.& * : mmP, h Devoted to Entomology in General U ■ l,i 7/ ky- ;>v :£t'S\ •7; JS--- ' 'b.vS \ - 5KV-A,> r~r 'rC ; '• Kb^- v>cK 1 -Y '/a ’T v..; m '{ s 3 -■ •A ■YKi,7^ k.‘ ■/' y m/y W- X'isvl v,v ■ m > ‘.4= 'Art V ■ \ XX\ V'\$ m A(1 . ; ' K\: X 'O \f > Vi;. ');Mi ' uvl . : %V irff- m 3X ‘ u ' X'M XXXX . X ■ ' i' ' ' '' 1 V Officers for the Year 1967 ' SA Ay. A. Y; Yj v * President, Dr. Richard Fredrickson College of the City of New York 10031 ■VjYjYi 'V- / ' fl , Ay Y^YYa'^Y1'' Y I”;/ Y'.ft; •< i?'& W! M Nf l ■' M i . ^ \\ Y-J A ill 7 7,AA)'.,r A7 Vice-President, Dr. David Miller .1 (V m j .( x ftp, / vN. I I I : College of the City of New York 10031 i> ./?,]>■ /', ' ' - : i i / y / < 5 v; ■ Secretary, Mr. Howard Topoff American Museum of Natural History, New York 10024 A f Assistant Secretary, Mr. Albert Poelzl * ff ffik: \X/ff 1 / A Treasurer, Mr. Raymond Brush f \C r s YY XXX, I A i YL Y Y Y 7 A fY ■ Y \XX Y 230 E. 78th Street, New York 10021 m X ■ 7 A fN \// 1 'Ak;", I V ■, W i. H, ' , , , , P7\l American Museum of Natural History, New York 10024 Assistant Treasurer, Mrs. Patricia Vaurie T \ _!. +'/ _ y\ / f ~ ( . American Museum of Natural History, New York 10024 \X IV ' Y Lfk ; V 1-4' \- ' V\ -rM i w \< i \f x' \ i-i Trustees A « A A p-' .dfr’. v. ■ i- V I'ffi'Ar; x\ , 1 .1 I ; /7 v X AS Class of 1967 ' v7‘ Dr. Jerome Rozen, Jr. Classy of 1968 Dr. Elsie Klots ■k \ 7 ' / 1 /,. /! r, ‘ p i ly ' New York Entomological Society A; CJ Id1 >, . V'l y,i,: ssmi : n, v y\%H ■\ h Organized June 29, 1892 — Incorporated February 25, 1893 Md sG l : v \ jIM 1 v) c'Xxx'- Reincorporated February 17, 1943 f- fe a? T'- » V ;pdX :r A y' d 4 'X Si/, -A • . 1 l i rS V ' I- v'XX d XY..X m ' jj ,, v/ , t / m fy /• 1 1/ .- • ' /I'--. \ • V»H'T/ /’- \ K • ■■ n,i-/Tv - ( ..,/ -. 1 , ./ .// , 14 ) The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural -*TV I SV X •ft ' History, 79th St., & Central Park W., New York 24, N. Y. 'Ty ■ i rf M-' /_ dX Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. I’ y. , y - . \> , yyH c u x • xycyvxXxxxX' Members of the Society will please remit' their annual dues, payable in January, to the At Treasurer. I r t . >1* V ■>( v •A 1/ & V vv T YdAXdUv) \XX- V-r'- ;V4^ >1 : V ' V J I, X' i ii I I A. it 1 ' l W . v, i . fyM'ti ' 1 Officers for ^ihe Year 1967 n ; \ 1 President , Dr. Richard Fredrickson : 8 v 1 I V /f Xi iWh io ' < ' ' I % ' X; X / 'X • C A / L ' College of the City of New York 10031 y\j Vice-President , Dr. David Miller ; ; |i 1, ; A ; 7 t ’Hx ' ' I \i\i ' V . ' C it l n \A A. ( m -i College of the , City of New York 10031 li/f : ' ,N Vv i rVV' Secretary , Mr. Howard Topoff . < MS A' r\‘ A , X /. American Museum of Natural History, New York 10024 rt'.'Vt v ( . ' Assistant Secretary , Mr. Albert Poelzl >tr U’-'A j 230 E. 78th Street, New York 10021 Treasurer , Mr. Raymond Brush Ml ■■ > i American Museum of Natural History, New York 10024 ( r ! 1 y !i .v , j 1 ../ Assistant Treasurer, Mrs. Patricia Vaurie XXV'7 im i y . \v ' 14 miyt-y ’ Av' A> A American Museum of Natural History, New York 10024 r> . m-, /..A. Ayy\ X;'kt fbd y\\ ■ g"7\ J / A 1 1 1 1 ' i / NV > Yr^i / ■ • | 1 1' s : 1 / - '.A',' dyy t SC id t f '\ A A Ad \l V < .h V , \ \ ' A * V ,.x- A' V Jf • M . \ i V A . 1 - l I I . \ --Vt Trustees IV\ ivi 'I 1 ■!) ' ii - V'X' Class of 1967 ■h , A'it ui r > , Dr. Jerome Rozen, Jr. Class of 1968 Dr. Elsie Klots vf ,'rU r ^ A1' k vp> ■ u ■/ N .\.^A iVi Vi yVLv-'A - -Vi • M) y-. ; i 1 J ' ' N 1 , : / Mr. Robert Buckbee ^ \ K " ' iVA V' Vv.l, 'v I k ty ■; A ,IM\ / \ ", k V t \ X ' Ii r. \j « ■/ J 1 'y\ lAXit' ' y' 1 tl , f\ ' < ! a . \ - tX 'W'^: \ y f Xf Air. Bernard Heineman — u, r- Mailed June 29, 1967 \ . A )• \ ^ l \ -1 Vy. y \ The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press Inc., 1041 /New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas. . . , , i} i - Mw t ■ ' ' . ’ ■ kV .. Vk- h k d v 1 /*. if i it Yd Xy - w : '~Y> f- :if\r A ha ; v ' / v A W 4. kk P" ;v u ' 7. Y w- ■’ % X >- To ® i .A1 I . / V ' V I>,d A X ' ^ > d r -i. / 7 : „ \ .. TV I k , . k:\vX AryHVA \ i h. '! \ ■ ■< \ y A/- Journal of the New York Entomological Society Volume LXXV June 29, 1967 No. 2 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen College of Pharmaceutical Sciences, Columbia LTniversity 115 West 68th Street, N. Y. 10023 Associate Editor James Forbes Fordham University, N. Y. 10458 Publication Committee Dr. Kumar Krishna Dr. Asher Treat Dr. Pedro Wvgodzinsky CONTENTS Larval Dimorphism and Other Characters of Heterocam pa pulverea (Grote and Robinson) (Lepidoptera : Notodontidae) Alexander B. Klots 62 Observations on the Behavior of the Bee Anthidium manicatum (L.) L. L. Pechuman 68 A New Species of ISepytia from the Southern Rocky Mountains (Lepidoptera: Geometridae) Frederick H. Rindge 74 Further Records of New Jersey Aphids (Homoptera: Aphididae) Mortimer D. Leonard 77 Further Studies on the Internal Anatomy of the Meloidae. III. The Digestive and Reproductive Systems as Bases for Tribal Designation of Pseudomeloe miniaceomaculata (Blanchard) (Coleoptera: Meloidae) A. P. Gupta 93 Book Review 100 Proceedings 101 New Members 110 Invitation to Membership 111 Larval Dimorphism and Other Characters of Heterocampa pulverea (Grote & Rohinson) (Lepidoptera: Notodontidae) Alexander B. Klots American Museum of Natural History and City College of New York Abstract: A group of sibling larvae of Heterocampa pulverea (Grote & Robinson) from Connecticut showed a very distinct dimorphism of color and pattern with no appreciable intergradation. Of 66 larvae reared to maturity 30 were green, 36 were brown. The dimorphism was apparently not linked with rate of development, sex or any discernible adult characteristic. The larvae of both morphs were highly, but differently, cryptic. Possible adaptive advantages of the morphs are discussed. Dorsal thoracic tubercles in the last larval instar, characteristic of this nominal species, are visible as vestiges in the pupa. On August 1966 a batch of eggs was obtained from a 9 Heterocampa pulverea at Putnam, Windham Co., Connecticut. The larvae from these were reared on Quercus coccinea. Ten were given to another Lepidopterist, but 56 were reared to maturity by the writer, emerging 8-26 October 1966, indoors. Tt was not until the larvae were in the 4th (penultimate) instar that it was realized that a distinct color and pattern dimorphism existed, approximately half being green and half brown. The two groups were then segregated and reared separately. Records of both types in the last two instars were made by color photography. Table 1 shows the record of the adults that emerged, grouped by larval morph, sex and the dates of emergence. The adults differ from each other in only very minor details, well within the limits of variation of any series from the region. These data show that the morphs, which must be genetically con- trolled, are not linked with either sex or rate of development. The larva of this species was first described by French (1880, p. 83) from an Illinois specimen. Packard (1895, p. 249-250 & 282, PI. 33, fig. 8-8a) re- printed French’s description, described a preserved specimen from Massachu- setts, and gave 2 small outline drawings copied from figures of Doubleday of a supposed synonym. Packard also refers to an unpublished colored sketch of the larva by Abbot. The French and Packard descriptions are of green larvae with a pattern not unlike the green morph described and figured here, but differing greatly in some respects. Apparently the white dorsal areas char- acteristic of both the green and the brown morphs of the present paper, and the lateral white areas of the green one, were not present in the French and Packard specimens, since French refers to these areas as “orange” or “purple,” and Packard either does not state what their colors were or else refers to them as “reddish.” Neither author mentions a brown larva. The Doubleday figures are too small and simple to be of much value. It is very likely that the larvae of pulverea show a considerable amount of variation, predictably much more than would be expected in a sibling group 62 June, 1967] Klots: Heterocampa Larval Dimorphism 63 Table 1. Sibling H. pulverea grouped by larval morph, sex and date of adult emergence. Green Brown 3 $ 3 $ October 8 2 _ 9 - - - 1 10 - 3 2 3 11 2 1 - - 12 3 - 1 4 13 - - - 3 15 2 3 2 3 16 - 2 - 4 17 1 1 1 - 18 - 1 1 1 19 1 - 4 - 22 1 - - - 23 — 2 - - 26 - - - 1 Totals 12 13 11 20 such as that described here. The extent of this in local populations, the amount it is subject to regional variation, and the genetic factors responsible, will all have to be worked out by many rearings of sibling groups and by genetic crosses. At present H. pulverea (type locality, Pennsylvania) is considered a northern subspecies of H. umbrata Walker (type locality St. John’s Bluff, East Florida). It is more than likely that the relationship is a clinal one. DESCRIPTIONS OF MATURE LARVAE Green Morph (Fig. 1). Body bright green speckled with small, dark, purplish fuscous dots which remain separate from each other, not coalescing to form lines or scrawls. A distinct white spot around the base of each primary seta. A white patch on either side of metathorax and 1st abdominal segment, running dorso-caudad diagonally from leg base, sometimes barely reaching spiracle, sometimes enclosing it and extending about one or two spiracle’s lengths above it. On abdominal segment 3 a broad, white patch running dorsad from the proleg base to join the white dorsal markings, occupying nearly all of the lateral area of the segment. On abdominal segment 6 a similar white patch running dorsad from the proleg base; this may join the white dorsal area or may fail to do so, extending no more than about two spiracle’s lengths dorsad of the spiracle. All three of these lateral white patches are very irregularly crenately edged, and contain curved, red-brown dashes and scrawls which differ greatly in extent in different individuals. Rarely there is a small, double patch above each metathoracic leg, and another on the posterior part of ab- dominal segment 7, largely ventrad of the line of the spiracle. Dorsally the markings are complex and differ greatly from one individual to another. The fundamental marking is a white dorsal stripe along the entire 64 New York Entomological Society [Vol. LXXV Figs. 1-2. Mature larvae, Heterocampa pulverea, lateral and slightly ventral aspect, drawn from projections of 35 mm. photographs. The setae of both larvae are incompletely shown. Fig. 1, green morph. Fig. 2, brown morph. length of the body, which is more or less margined and marked internally by dark red-brown scrawls, and differs greatly in width on different segments. Prothorax: stripe unmarked, anteriorly as wide as space between prothoracic tubercles, tapering posteriorly to half as wide, black-edged. Mesothorax and metathorax: stripe narrow anteriorly, widening greatly posteriorly, usually considerably marked internally, and sometimes nearly obliterated, by dark scrawls. Abdomen, segment 1: stripe widening greatly posteriorly to slightly more than half the width of the segment; rarely with any included dark mark- ings, but often pale green mid-dorsally, the green area narrow anteriorly but widening greatly posteriorly so as to leave only narrow, white, tapering edges laterally which in extreme individuals may not reach the posterior edge of the segment. Segment 2: white stripe becoming very broad posteriorly, containing more or less green mid-dorsally. Segment 3: white stripe very broad, laterally confluent with lateral white stripe, from dorsal view occupying all or nearly all of the segment; subdorsally a few small, dark, paired dots and scrawls, especially posteriorly. Segments 4 & 5: white stripe very broad anteriorly, narrowing greatly in segment 4 and still more in segment 5 ; within it a broad, dark scrawled, X-shaped saddle, centering about anterior edge of segment 5, that may obliterate much of the white. Segments 6 & 7 : rarely almost solid green June, 1967] Klots: Heterocampa Larval Dimorphism 65 mid-dorsally with only indications of the white stripe laterally; sometimes with only central portions green, and white stripe on either side of this broad and confluent with lateral white stripe on segment 6. Segments 8, 9 & 10: mid- dorsal area green, white stripe on either side of this broadest at anterior edge of segment 8, narrowing to segment 9, broader at anterior edge of segment 9, narrowing posteriorly; sometimes the green areas of the sides and the mid-dorsal green are confluent along the anterior edge of segment 9, breaking the white stripe. Brown Morph (Fig. 2). Head, prothoracic tubercles, legs and seta bases as in green morph. Body brown with only a faint greenish cast in recently enclosed individuals. Laterally with no white bands or areas other than a few small areas enclosed by dark scrawls. All brown areas with many irregular, dark brown curved lines and scrawls and smaller, orange-brown dots and curved lines. Dark scrawled markings heavier and coalescing to form a diagonal line running dorso-caudad from base of 3d leg across metathorax and abdominal segment 1 to join dark-scrawled border of dorsal markings. A similar, but less complete, line of markings running dorso-cephalad from base of proleg on ab- dominal segment 3. A similar, also less complete, diagonal line of dark markings running dorso-cephalad from base of proleg on abdominal segment 6 to spiracle on abdominal segment 5, and more or less continued cephalad across abdominal segment 4. Abdominal segment 7 with dark-scrawled patch caudad and mostly ventrad of spiracle, dorsally more or less joining lateral dark edging of dorsal markings. Dorsally, fundamental pattern like that of green morph, but with some dif- ferent distribution of white. Prothorax: as in green morph. Mesothorax & metathorax: also much as in green morph, but with less white, the dorsal areas largely filled in with brown scrawled marks as in the most heavily marked green individuals. A large, irregularly edged, diamond-shaped white area from pos- terior part of metathorax back to about middle of abdominal segment 4, widest in posterior part of abdominal segment 2 ; within this for most of its length is a pair of narrow, irregular, closely subdorsal, dark lines. An almost solidly brown saddle (in the same position as the dark-scrawled, X-shaped saddle of the green morph), continuous with brown sides, on posterior half of 4th and anterior half of 5th abdominal segments. A large, posterior white patch, be- ginning narrowly at about middle of 5th abdominal segment and extending to posterior end; on 8—1 0th abdominal segments this is more or less filled in dorsally with brown scrawls and lines; within it, as in the anterior white patch, is a pair of irregular, thin, dark, closely subdorsal lines for most of its length. Despite the considerable amount of individual variation, the two morphs in this group of siblings were very distinct, with no intermediate individuals. The nearest to anything of the sort was in a few larvae of the brown morph that had a greenish tone during the early last instar; and one individual of the 66 New York Entomological Society [Vol. LXXV green morph that had the green areas much paler than usual and slightly brownish tinged, but had the green morph pattern. 4th instar larvae The larvae of this instar are easily recognizable by the ends of the prothoracic tubercles, which have two distinct small, setiferous tubercles at the tips, in- stead of being terminally smooth as in the 5th instar. On the face these larvae have two thin fuscous lines on either side of the median light area instead of the single line of the 5th instar. The white lateral patches, and to a lesser degree the white dorsal patches, of the green larvae tend to be more obscured by dark scrawls. The brown larvae frequently had considerable of a greenish tinge, although their patterns were definitely of the brown morph. PRE-PUPAL LARVAE As the larvae stopped eating and entered the ground for pupation, drastic color changes ensued. All fine details of the pattern disappeared. The brown larvae turned a brilliant pink overall, the dark markings of the saddle on ab- dominal segments 4 & 5 showing slightly darker. The green larvae, on the other hand, changed to a darker green with the white areas of both the sides and the dorsum very bright pink, making them very conspicuous looking objects. All larvae then became pale and almost colorless just before eclosion to the pupa. The pink larvae that had been brown did this at a uniform rate overall. In the green larvae, however, the pink areas were the first to become color- less, so that for a short time these larvae were green with pale, colorless areas. Doubtless these color changes have some physiological significance, but they can hardly have any protective value (as is the case in some other pre-pupal color changes) since they normally take place underground. DISCUSSION The patterns of both of the larval morphs are decidedly, but differently, procryptic, the brown larvae resembling crumpled, dead leaves with shadow or edge patterns, and the green larvae resembling green leaf areas with pieces missing. The larvae of both types are highly disruptive from the dorsal aspect, and the green larvae are disruptive from lateral aspect as well. The white lateral patches are so shaded as to appear almost protuberant and three dimen- sional. A predator that had learned to recognize the appearance of one of the morphs would be very unlikely, because of this, to react to the appearance of the other one and might very well, in fact, be more likely to ignore the other one if the two were close together. The dimorphism must function in this way as a protective device per se, most valuable when the two morphs are com- pletely different from each other, and still more valuable when each morph is highly cryptic. June, 1967] Klots: Heterocampa Larval Dimorphism 67 The proportions of the morphs in this sibling group and their distinctness from each other strongly suggest a single controlling genetic factor. The evi- dence of French’s and Packard’s larval descriptions shows that there is much more larval variation than this sibling group showed, and suggests that the morphs may not always be as distinct from each other. For the time being we suggest that the morphs have evolved, and are maintained, by visual predator selection, but that this may well be strongly affected by all sorts of pleiotropic effects of which nothing is known. Much further work is certainly called for to determine the genetic status, possible pleiotropy and extent within both H . pulverea and H . umbrata of larval dimorphism. The pupae all showed vestiges of the prothoracic tubercles. Since these tubercles appear to be present in the 5th instar of only the larvae of H . pulverea and H. umbrata , their presence in the pupa can be used for identification, at least of H. pulverea. Identification of the material as H . pulverea was by comparison with the 2 type in the American Museum of Natural History. The material here reported upon has been placed in the collection of this museum. Literature Cited French, G. H. 1880. Canadian Ent. 12: 83. Packard, A. L. 1895. Mem. Nat. Acad. Sci. 8: 249-250 & 282, PI. 33, fig. 8-8a). Received for publication March 16, 1967 Observations on the Behavior of the Bee Anthidium manicatum (L. ) L. L. Pechuman Cornell University, Ithaca, N.Y. Abstract: Collection records of the Palaearctic bee Anthidium manicatum (L.), reported by Jaycox in 1967 as being adventive in the United States, are brought up to date. New flower host records are included. European literature on the aggressive behavior of the male is briefly summarized. Observations on the behavior of A. manicatum in 1965 and 1966 show the male to be territorial and aggressive. The female works without hindrance while other species of bees are struck and driven from the territory being patrolled by the male. No bees showed any inclination to defend themselves against the attacking male of A. manicatum. It is believed that A. manicatum is a rather unique subject for further study, including distribution, behavior, nest building, flower preferences and genetics. Jaycox ( 1967) reports the presence in the United States of the Old World bee Anithidium manicatum (L.) (Megachilidae) based on specimens collected by Dr. Roger A. Morse and the writer in 1963, 1964, and 1965. A. manicatum is found throughout Europe, part of Asia, and North Africa. It is the only species of Anthidium found in England. As mentioned by Jaycox, A. manicatum has recently been found in the Canary Islands and in South America. The specimens seen in 1963 were reared by Dr. Morse from a five inch deep, one quarter inch diameter trap nest in a white pine block, placed in the field early in 1963 near Ithaca, N.Y. The wooden block containing the nest was removed from the field on 27 June 1963; on 20 August 1963, adults, 2 3 3 and 8 9$, emerged from the nest. All specimens collected by the writer in 1964 and 1965 were taken, as reported by Jaycox, from the flowers of Caryopteris X clandonensis at Ludlowville, N.Y. In 1966, A. manicatum was again found at Ludlowville, N.Y. visiting the flowers of Caryopteris. Specimens were observed between August 28 and Oc- tober 3 with peak abundance during the second week of September. It was noted in 1964 and 1965 and again in 1966 that A. manicatum visited only the flowers of Caryopteris although Chrysanthemum and Potentilla were interplanted with the Caryopteris and were in bloom during the flight period of the bee. Two species of Mentha in bloom nearby were attractive to other species of wild bees but were not seen to be visited by A. manicatum. Also in 1966, a total of 13$ 9 and 15 3 3 were taken on the Cornell Campus at Ithaca on various dates between August 23 and September 2 by Jan Nowakowski, Paul Minacci and George Strang from a bed solidly planted to blue flowering salvia ( Salvia jarinacea ) . Dr. Nowakowski informs me that none were taken from adjoining beds planted to white salvia (S. jarinacea) and red salvia (S. splendens) . Also on the Cornell Campus, Dr. Nowakowski took 3 9 9 and 1 3 from Lythrum salicaria on August 1 6 and a single 9 from 68 June, 1967] Pechuman: Observations on Anttiidium manicatum 69 Solidago on September 12. Dr. Nowakowski noted aggressive actions against other bees by the males of A. manicatum he collected from salvia. It is of interest, although possibly of little significance, that during a three year period all but one specimen of A. manicatum were collected on blue or purple flowers and all but five specimens from the rather closely related families Labiatae ( Salvia ) and Verbenaceae (Caryopteris) . Friese (1898) says A. manicatum prefers Labiatae in Europe but there is no general agreement by other workers on this. It also raises the question of why plants of Mentha (Labiatae) in full bloom were ignored at Ludlowville. Unfortunately no notes were made on the structure of the nest from which specimens were reared by Dr. Morse in 1963. None have been found in trap nests in subsequent years. Very likely the nest is made from soft flocculent material scraped from plants as reported in Europe. Fabre refers to nests of the group to which A. manicatum belongs as “ — quite the most elegant speci- men of entomological nest building” and Friese calls them “wunderbaren Nestbau.” In 1965 the writer observed a female stripping the pubescence from the flower stem of a potted geranium (Pelargonium) , probably with the intent of using it as nesting material. No collections of adults have been made in New York before August. How- ever, the specimens reared in August 1963 by Dr. Morse came from a trap nest placed in the field early in 1963 and completed by June 27. This may indicate that A. manicatum has two broods. Green (1921) in England seems to have been the first to note the aggressive habits of A. manicatum males when he reported it attacking Bombus. Ward (1928), also in England, published detailed observations on attacks by males of A. manicatum on bumble bees ( Bombus ) and hive bees (Apis). He indicates that definite territories were marked out when he states, “ — males patrolling patches of Red Dead Nettle at two spots and having the effect of keeping other insects away; but a few yards away Bumble Bees feeding fairly regularly at the Dead Nettle with little or no molestation.” He noted that aggression declined when the sun was obscured by clouds. Ward also found that some individuals of bumble bees and honey bees had their wings damaged so they could not fly when struck by A. manicatum. In spite of Ward’s detailed notes, Perkins (1928) regarded the attacks on other bees as “an accidental occurrence.” Sitowski (1947) in Poland reports that the male of A. manicatum , “ — hovers in an area, or patrols where the female is working and kills or drives out all competing intruders with ferocious attacks.” He states that not only is the competing bee knocked to the ground but that the male A. manicatum may continue its attack on the ground using its mandibles, and abdominal spines on the last two abdominal segments, to disable or kill honey bees and bumble bees. 70 New York Entomological Society [Vol. LXXV The observations of Haas ( 1960) in Germany are similar to those of Ward. He regards the territory established by the male as part of a behavior pattern which involves swarming. The territory itself he believes to be sort of an exclusive swarming area in which the male as Haas puts it, “swarms alone.” The writer first observed the male of A. manicatum attacking other bees on 14 September 1965. An abstract of notes taken on that day follows: 14 September 1965 In addition to honey bees, bumble bees and a few other native bees, two female Anthidium manicatum were present most of the day on Caryopteris flowers. The females were distinguished by their very fast flight and by being easily disturbed and alarmed; when disturbed by anything other than another bee they would leave the area and not return for some time. The females were far from aggressive. If one started to land on a flower and found it occupied by another bee, it would go to another flower. A bumble bee once pushed a female from a flower; the female flew to a leaf where it remained motionless for almost three minutes, then preened its legs and antennae for half a minute and then flew to another Caryopteris flower on a different plant. The male A. manicatum moved very rapidly. It would work a flower for a second or two but it spent most of its time patrolling the largest Caryopteris plant. It was very aggressive and would strike honey and bumble bees which were working flowers, knocking them from the flowers. The male frequently would strike two or three bees in as many seconds. On one occasion the writer frightened the male and it flew away for several minutes. In its absence, two bumble bees and three honey bees moved to the Caryopteris plant which had been patrolled by the male A. manicatum. On its return, the male im- mediately struck all five bees almost faster than the eye could record, the whole episode being over in five seconds or less with all five bees in flight. Observations were made on two successive days in 1966. The area under observation involved one large (56 in. high, covering an area of 18 sq. ft.) Caryopteris and a group of smaller (42 in. high, covering an area of 14 sq. ft.) Caryopteris plants separated by a pink flowering Chrysanthemum plant 23 in. high, covering an area of 3.5 sq. ft. The notes made on these two days follow: 10 September 1966 One male and one female appeared at approximately 9 A.M. During the day only one female was observed at any one time and apparently only one specimen was involved. The first male to appear was very dark and is referred to as No. 1. A second male with more extensive yellow markings appeared shortly on the smaller Caryopteris and is referred to as No. 2. Male No. 1 spent most of its time patrolling the large plant. Occasionally male No. 2 would extend his patrol of the smaller plant into the patrol area of male No. 1. Male No. 1 would immediately drive No. 2 away. On one occasion when No. 1 had pursued No. 2 to the outer side of the smaller plant, No. 2 turned and faced No. 1. Both males hovered about two inches apart, gradually descending toward the ground; at about four inches from the ground hovering continued at essentially one place for about half a minute; then No. 1 struck No. 2 head on knocking it to the ground beneath the plant where it remained with wings partly outstretched and with the apical third of the abdomen vibrating. Male No. 2 remained on the ground about three minutes and then it flew away. It was not seen again. Following this episode, male No. 1 rarely left the large Caryopteris all day. Occasionally it would make a quick patrol of the group of small plants formerly patrolled by No. 2. It had a regular route around and through the large plant and conducted its patrol by hovering a second or two and then flying four to six inches. All bees except female A. June, 1967] Pechuman: Observations on Antiiidium manicatum 71 manicatum were driven away. Usually it would strike the center of the thorax, possibly because this was the usual aspect exposed; it was seen once to strike a bumble bee head on and once struck a bumble bee from below. Rarely a very small bee would manage to visit a flower and be overlooked by the male but usually it would be struck as soon as it tried to move to another flower. All bees, including the largest bumble bees, appeared to be panic stricken when struck by the male A. manicatum ; none made any attempt to fight back and only one, a large Xylocopa, was noted to require two strikes. Bumble bees slowly flying by the plant were sometimes struck and immediately put on an amazing burst of speed. The male rarely bothered the female. Several times it landed on the dorsum of the female giving the impression it was trying to mate. It could not be determined if mating took place but the contact would sometimes last for eight to ten seconds. During contact the female would keep working the flower but once the pair fell from the plant, separating before they reached the ground. A bumble bee was killed with cyanide and immediately pinned to a flower in a natural position. The male A. manicatum did not strike it but circled it twice about one half inch away ; from then on it was ignored by the male on his patrol except at rare intervals when it would fly very close to the pinned bee. When the bumble bee was moved to another flower, it continued to be ignored. A bumble bee was quieted with DDVP and tied to a blossom while still moving its wings. It was struck by the male as it was being tied but was ignored from that time on except for a rare quick investigation. At the same time the male was striking all intruding bees. It was noted that during the heat of the day the male was extremely aggressive and spent very little time on flowers and none resting. After 5 P.M. it made many stops probing flowers although each stop was only of a few seconds duration. It also would rest for five to eight seconds on foliage. At this time of day it was not quick to strike intruders but it did strike them eventually. This may have been due to lower tempera- ture, wearyness, or the need to secure some nectar to sustain itself. 11 September 1966. The activities of male No. 1 were about the same as noted on the previous day. It now took over the smaller plants patrolled by No. 2 the day before but about 75 percent of its time was still devoted to the large bush. Two females were present most of the time. A second male appeared but was driven off and did not return. When the male would land on the dorsum of the female, its behavior was quite different than when striking an intruding bee. As it approached the female it would stretch out its legs as for grasping and the female would be seized by them. When striking another bee, the legs were kept tightly under the body and the approach was much faster. Live bumble bees were attached by a long thread to the end of a stick. To the observer they looked and behaved quite naturally but only occasionally would there be a glancing strike by the male Anthidium whether the bumble bees were on a flower or flying. How- ever, if a tethered live bumble bee was dangled two to four inches directly in front of the hovering male, the male could be led for a foot or two but it would not strike. One live bumble bee tied to a flower was closely investigated several times but not struck; most of the time it was ignored. The male would investigate anything that moved including dangling portions of old flowers but did not strike such objects. It showed only slight aggression against flies and butterflies and these did not show the fear of the male exhibited by the other bee species. The strike against flies and butterflies was usually glancing rather than direct and these insects would usually return to the same or a neighboring flower immediately. By 6 P.M. the male was spending most of its time visiting flowers. As it approached the flower it would drop its hind legs as does the female. 72 New York Entomological Society [Vol. LXXV Observations after September 1 1 were mostly a repetition of previous ob- servations. The Caryopteris bloom was almost gone by the end of September. The last A. manicatum noted was seen for a few moments on October 3 about 3 P.M. It was a male and appeared to be the same specimen observed on September 10 and 11. Observations made in 1965 and 1966 seem to indicate that the male of A. manicatum is aggressively territorial. Possibly the easily disturbed timid female needs protection when there is competition for pollen and nectar. Other bees seem to fear the male of A. manicatum and never were observed to attempt to defend themselves. Flies and butterflies, although occasionally knocked from flowers, showed no such fear and usually returned to the same or a nearby flower. The male was noted to be most aggressive in bright sunshine during the heat of the day; it is less quick to respond to invasions of its territory as the temperature drops later in the day. Although the male investigates all movement within its territory it does not strike dangling leaves or flowers or bees which are dead or whose movements are inhibited in any way. It is suggested that further studies of Anthidium manicatum in New York are likely to be rewarding. Currently it is not known outside of a limited range in the towns of Ithaca and Lansing in Tompkins County and its pattern of distribution as it spreads will be of interest. No native Anthidium is known from New York and one wonders if A. manicatum will fit in some unoccupied ecological niche or whether one or more of our native bees may be displaced by this aggressive species. The present population of the species is probably the result of the introduction of a limited number of individuals, possibly of a single nest, so a study of the genetics of the population might be in order. It is of interest in this connection that the color pattern of the males collected in New York run the complete gamut of patterns described from Europe — from mostly yellow with a few black markings to almost completely black. This variation in color pattern is very convenient for the observation of specific individuals. The writer wishes to thank Dr. Roger A. Morse for providing information on the specimens he reared from a trap nest. Acknowledgement is also due Dr. Jan Nowakowski for information on the specimens collected by him and by Mr. George Strang and Mr. Paul Minacci and additionally for translating the paper by Sitowski. The writer also wishes to thank Dr. Morse and Dr. Elbert Jaycox for reading the manuscript. Literature Cited Fabre, J. Henri. 1920. Bramble-bees and others. Dodd, Mead and Co., New York. 456 p. Friese, Heinrich. 1898. Die Bienen Europa’s (Apidae europaeae). IV. Solitare Apiden. C. Lampe, Innsbruck. 304 p. Green, E. E. 1922. Note on the habits of the bee, Anthidium manicatum. Proc. Ent. Soc. London 1921: lxxii— lxxiii. June, 1967 I Pechuman: Observations on Antitidium manicatum 73 Haas, Adolph. 1960. Vergleichende verhaltensstudien zum paarungsschwarm solitarer Apiden. Zeit. Tierpsychol. 17(4): 402-416. Jaycox, Elbert R. 1967. An adventive Anthidium in New York State (Hymenoptera: Megachilidae) . J. Kansas Ent. Soc. 40(1): 124-126. Perkins, R. C. L. 1928. A note on Mr. Ward’s observation on Anthidium manicatum. Entomologist 61(787): 273. Sitowski, Ludwik. 1947. [Anthidium, as an exterminator of bees and bumble-bees gathering honey.] Roczn. Nauk. Roln. Lesn. 49: 434-437. In Polish with an English summary. Ward, J. Davis. 1928. An unrecorded habit of the male of the bee Anthidium manicatum L. Entomologist 61(787): 267-272. Received for publication March 27, 1967 A New Species of Nepytia from the Southern Rocky Mountains (Lepidoptera: Geometridae) Frederick H. Rindge Department of Entomology, the American Museum or Natural History, New York Abstract : Nepytia janetae, new species, is described from material collected in New Mexico and eastern Arizona. The genitalia of both sexes are illustrated. Recent collecting trips to the higher mountains of New Mexico by the author and his family produced a nice series of a heretofore undescribed species of the genus Nepytia Hulst. One additional specimen was found in the collec- tion of the American Museum of Natural History, being from the White Moun- tains of Arizona, ex collection of G. H. and J. L. Sperry. These moths are now being described in order to make this name available. The material was collected under the auspices of National Science Founda- tion Grant numbers G-9037, G-25134, and GB-3856. This assistance is grate- fully acknowledged. Nepytia janetae, new species Figures 1, 2 This species is allied to regulata Barnes and McDunnough, and may be distinguished from it by its smaller size, paler color, and by the large discal spot filled with ground color on each forewing. male: Head with vertex and front creamy white, with variable number of yellow scales; palpi slender, grayish brown; antennae with very long pectinations, up to 1.6 mm in length. Thorax pale gray above, with elongate hair-like scales, and with grayish black scaling anteriorly; beneath white. Abdomen pale gray, with a few scattered pale brown scales above. upper surface of wings: All wings rather thinly scaled; forewings with ground color pale gray, with scattered black and yellowish scales, the latter concentrated along upper portion of t. p. line and inner margin; t. a. and t. p. lines broad, black or grayish black, and tending to be somewhat diffuse; t. a. line arising on costa one-third of distance from base, outwardly dentate on veins and in cell, inwardly oblique from anal angle to inner margin ; discal spot large, occupying most of width of cell, roughly triangular, filled with yellowish ground color; t. p. line strongly inwardly dentate on veins, connected with discal spot anteriorly along vein R5, and with broadening of t. p. line at junction of base of discal spot, in some specimens with small spot of ground color at origin of vein Ms; subterminal area with nebulous yellowish band distal of t. p. line in upper portion of wing, and with weakly defined s. t. line, shaded distally by ground color; fringe con- colorous with wing, with blackish gray spots at ends of veins. Hind wings white, with scattered brownish black scales; extradiscal line weakly indicated, extending straight across wing; discal dot weakly represented in some specimens; fringe like that of forewings. under surface of wings: Forewings pale grayish white, with maculation of upper sur- face weakly indicated; hind wings white, with faint extradiscal line. length of fore wing : 15 to 18 mm; holotype, 17.5 mm. female: Similar to male, but with maculation tending to be slightly heavier. length of forewing: 15 to 18 mm; allotype, 17 mm. 74 June, 1967] Rindge: New Species of Nepytia 75 Figs. 1 and 2. Genitalia of Nepytia janetae, new species. Fig. 1. Female, allotype. Fig. 2. Male, paratype from type locality. male genitalia: Gnathos with sides very slender, median spinose enlargement triangular in outline; valves with apex of costa protruding from end of valve, and with outer margin of valvula rounded; furca angled to right side, short, not attaining posterior margin of transtilla, broad, with inner margin straight and outer margin rounded; aedeagus with ventrolateral, sclerotized, posteriorly and asymmetrically bidentate area, and with slender, elongate, posterior, sclerotized process. female genitalia: Sterigma very broad, posterior margin evenly rounded, and with V-shaped anterior process ventrad of posterior one-half of short ductus bursae; corpus bursae with narrow posterior neck and anteriorly rather short and globular, with stellate signum. types: Holotype, male, Bursum Camp, 18 miles east of Alma, Catron County, New Mexico, elevation 9000 feet, July 9, 1961 (F., P., and J. Rindge); genitalia mounted on slide no. F.H.R. 10,650. Allotype, female, 76 New York Entomological Society [Vol. LXXV same data, July 15, 1961; genitalia mounted on slide no. F.H.R. 13,774. Paratypes: same data as types, various dates between July 7-16, 1961, 26 males and 21 females; Pine Camp, 2 miles northeast of Cloudcroft, Otero County, New Mexico, elevation 8000 feet, July 3-5, 1964 (F., P., and M. Rindge), five males; Bear Trap Camp, 28 miles southwest of Magdalena, Socorro County, New Mexico, elevation 8500 feet, July 1-11, 1965 (F., P., and M. Rindge), seven males and five females; Alpine, Apache County, Ari- zona, June 18, 1936 (G. FI. and J. L. Sperry), one male. All the type material is in the collection of the American Museum of Natural History. remarks: This species flies with its close ally, regulata , at all of the known localities for janetae. The new species can be separated from regulata by its yellowish vertex, the much longer antennal pectinations in the male, by the paler wing color, and by the very large discal spot of each forewing being filled in with yellowish ground color. The genitalia of the new species are similar to those of regulata. The males of janetae can be recognized by the distinctive gnathos, the apex of the costa extending above the surface of the valve, and by the straighter and broader furca. The females structures are characterized by the broader, semicircular sterigma, and by the narrower posterior portion of the corpus bursae. This species is named for Janet, my oldest daughter, who helped collect the topotypical series. Received for publication March 17, 1967 Further Records of New Jersey Aphids (Homoptera: Aphididae) Mortimer D. Leonard Collaborator, Entomology Research Division, Agricultural Research Service, U. S. Department of Agriculture, Washington, D. C. Abstract: Listed are 93 aphids arranged alphabetically by genera and by species under each genus. Detailed records of the localities, dates, food plants and collectors are given for each species and a list of 101 food plants on which the aphids have been collected is included. Of the aphids 20 species and of the food plants 26 have not previously been recorded from New Jersey. At present 227 aphids on 267 plants are known to occur in New Jersey. This is a third paper on the distribution and food plants of New Jersey aphids. The previous paper entitled “Additional records of New Jersey aphids” was published in the Jour. N. Y. Ent. Soc. 72: 79-101, 1964. It increased the number of aphids known to occur in New Jersey to a total of 207 on 241 food plants. The present paper, based largely on collections made during the past three years, 1963-1965, records 93 aphids on 101 plants of which 20 aphids and 26 plants were not in the previous papers. At present 227 aphids on 267 plants are known to occur in New Jersey. During visits to Haddon field I have continued to operate a yellow water-pan or Moericke Trap (in text as MT) in the back yard garden at 217 Rhoads Ave. Starting with 1963 I used the inverted top of an old ash can about 22 inches in diameter placed on a standard which raised it about two and one-half feet above the ground. This pan was exposed continuously in 1963 from 23 July to 30 November during which period about 2500 winged aphids were taken from it. Nearly 25% of this total was collected during November and a little over 40% during October, both of which months were unusually mild. Because of the difficulty of identifying so many free-flying aphids only a few of the records of these are here included. It is hoped at some later time more complete records from the yellow water-pan can be published. LIST OF APHIDS* Acyrthosiphon dirhodum (Wlk.) — see Metopolophium. * A cyrtho siphon pelargonii (Kaltenbach ) , Geranium Aphid. Maywood (Hoffman, Florist), 5 Aug. 1965, a general heavy infestation of all plants of Salmon Irene geraniums ( Pelargonium sp.) on stock in a greenhouse which is open in the summer (Conlon coll.) Acyrthosiphon pisum (Harris), Pea Aphid. In November, 1961, L. W. * Names preceded by an asterisk (*) are in addition to those in the previous two papers. 77 78 New York Entomological Society I V ol. LXXV Coles of the Japanese Beetle Laboratory, USDA, Moorestown, wrote me about the status of parasites of this aphid in New Jersey. This was omitted from “Additional Records.” He says: “The parasites of the pea aphid in New Jersey that we are familiar with are Aphidius pisivorus C. F. Smith and Praon simulans Provancher. We have observed them in all areas of New Jersey every season for the past six years. A. pisivorus is very common and very effective we feel. Praon can be found commonly but not nearly so as A. pisivorus.'1' Also omitted from “Additional Records” were any notes on the pea aphid although it is given in the Plant List under alfalfa and red clover. The data is as follows: “The pea aphid caused less damage than usual [to alfalfa]. In some areas of southern counties populations reached 200-300 per sweep during May but, in general, populations were much lighter.” (Summary of Insect Conditions — 1957 in New Jersey in CEIR 8(1): 6, Jan. 3, 1958.) “Was far less damaging than usual.” (Summary of Insect Conditions — 1958 in New Jersey in CEIR 9(7): 189-190, 1958.) New Brunswick, 16, 20 May and Beemerville, 6 May 1960 on alfalfa (Wave coll.), Middlebush, 8 June 1960 on red clover (Wave coll.). Acyrthosiphon porosum (Sanderson), Yellow Rose Aphid. McGuire Air Force Base, 3 alatae, 2 “pupae,” 2 mature and 3 immature apterae, collected from the buds of cultivated rose in mid-May 1965 (Quinden coll.). Second record for New Jersey. * Acyrthosiphon sibericum (Mordvilko). Haddonfield, 2 Sept. 1965 on Urtica sp. (MDL and DLW coll. — ATO det.). Although this aphid is recorded as fairly common in the Rocky Mountain Region it is known elsewhere in the USA only by one collection in N. Y. and 2 in Pa. Acyrthosiphon solani (Kaltenbach) (placed by some in Aulacorthum) , Fox- glove Aphid. New Brunswick, 23 June 1960, 1 mature aptera on Taraxacum officinale (Wave coll.). Omitted from the previous paper. Anoecia corni Fabricius. Haddonfield, 16-30 Sept., 9 alatae; 1-15 Oct., 2 alatae; 16-31, 4 alatae; — 1963 and 1-5 Nov. 1965, 2 alatae — all in MT (MDL coll.). Anuraphis viburnicola (Gillette) — see Ceruraphis viburnicola (Gillette). Aphis coreopsidis Thomas. Whitesbog, 13 July 1961 on Nyssa sylvatica (Marucci coll.) . Aphis crataegifoliae Fitch — see Brachycaudus crataegifoliae Fitch. Aphis fabae Scopoli, Bean Aphid. Moorestown, 12 April 1963 on Euonymus europaeus (EAR coll.); 21 May 1963 on rhubarb plants in a garden heavily infested, with leaves curled and crinkled (HWA coll.); 5 Sept. 1963 on cult, nasturtiums (EAR coll.). Bordentown, 24 May 1963, abundant on Philadel- phus sp. (Webber coll.). Ridgewood, 10, 18 June 1963 a few on Arctium June, 1967] Leonard: New Jersey Apeiids 79 minus (MDL coll.). Columbus, 1 Oct. 1963, 20 alatae from Arctium sp. (LW Coles coll.). Haddonfield, 27 Aug. 1963, scarce on nasturtiums (MDL coll.). Dr. Allen’s heavily infested rhubarb made me realize that I had seldom seen records of this aphid on rhubarb. A search reveals it appears there are not many in the United States. The files of Survey and Detection Operations, Plant Pest Control Div., USDA have only the following: USDA Yearbook for 1908, p. 570 “caused serious injury to rhubarb in New Jersey”; Manhattan, Kans., 1 July 1948 taken on rhubarb (R. C. Smith coll. — LMR det.); “Serious damage caused to rhubarb on Mar. 20, 1948 in Arcadia.” (Calif. Truck Crop Emergency Survey); Palmer in Aphids of the Rocky Mt. Region states that it occurs on rhubarb. For New York there are only two records: Lockport, 1959 and Orient, L.Id, 1962. Amherst, Mass., 6 June 1960 on rhubarb. Aphis gossypii Glover, Cotton or Melon Aphid. Whitesbog (Pemberton), 13 July 1961, 3 alatae, 3 apterae on Leucothoe racemosa and 9 apterae on Rhododendron ( Azalea ) viscosa (Married coll. — JOP det. with query). In litt. from Marucci — “ A phis gossypii apparently can live on ericaceous plants. On 1 Sept. 1947 we found it colonizing blueberries in our screen-house at Pemberton. The aphids were being attended by ants. We used these aphids to try to transmit blueberry stunt disease and they lived quite well on blue- berries. USNM made the determination.” Moorestown, 15 May 1963, abun- dant on shoots of rose-of-sharon (MDL coll, in EAR’s garden); 13 June 1963, several mature apterae and some younger ones on Sophora japonica (EAR coll, in his garden); 21 May 1965, 9 alatae on Aguilegia longissima (HWA coll.) — these may be “drifts” since this aphid has been recorded from Aguilegia only by Hall in Egypt; 21 May 1965, 8 alatae on tips of several shoots of Forsythia sp. (HWA coll.) — these may be “drifts” since this aphid has been recorded from Forsythia only in Japan; 9 alatae on buds of peony (HWA coll.) — probably “drifts” since aphids were stuck to the buds and pre- sumably no aphid has been recorded from peony. Haddonfield, 15 May 1963 a very few apterae on Campsis radicans (MDS); 19-26 Sept. 1964, an occa- sional leaf on two rose-of-sharon shrubs with a single alate, one of these near several very small pale young (MDL); mid-May 1965, a very small immature on a tender tip of rose-of-sharon (MDL & DLW coll.). Trenton, 2 Sept. 1964, the “small form” heavy on leaves of Catalpa sp. (Stinson coll.). Princeton, 17 Aug. 1964, heavy infestation on small twigs of several 4-5 foot trees of Sophora japonica (Stinson coll.). *1 Aphis incognita Hottes & Frison. Pemberton, 1948, an alate on sticky board trap in blueberry field (Marucci coll. — LMR det. as “near incognita''1) . This species has been recorded from Utah, Colorado, and Illinois from Sym- phoricarpos. Aphis oestlundi Gillette. Mt. Laurel, 25 May 1963 on Oenothera sp. (HWA 80 New York Entomological Society LVol. LXXV coll.). Indian Mills, 26 May 1963 on Oenothera sp. (HWA coll.). Moorestown, 17 Oct. 1963 on O. biennis (T. L. Ladd coll.). Aphis pomi DeGeer, Apple Aphid. Bordentown, 27 May 1963 on crabapple (Weber coll.). Somerville, 19 June 1963 on flowering crab (Stinson coll.). Bridgeton, 7 Aug. 1964 on Jap flowering quince (W. Junghans coll.). Moores- town, 21 May 1965 on Jap flowering crab, heavily infested (HWA coll.). Aphis pseudohederae Theobald, Ivy Aphid. Haddonfield, 1963 — none could be found during the season on the English ivy at 217 Rhoads Ave., until about Sept. 1 when three or four occurred on as many tender tips; on 31 Oct. 4 small colonies ( 1 or 2 alatae in each) on vines in another similar situation; 2 Dec. a few were found including several alatae from which several atypical Lysephlebius testae eipes (Cresson), det. Muesebeck, were reared; on 25 Sept. 1964 a few, including 3 or 4 alatae, on the tender tips of the English ivy on the house at 217 Rhoads Ave. (MDL); 6 Nov. 1965 a very small colony on the same vines attended by the ant, det. D. R. Smith, as Prenolepis imparis (Say). Ridgewood, 28 Oct. 1965, a fair sized colony on a tender shoot of an English ivy vine on a tree trunk, attended by the ant, det. by D. R. Smith as Prenolepis imparis (Say), (MDL). Aphis rumicis Linnaeus, Dock Aphid. Rancocus, 13 May 1963, a heavy infestation on Rumex ads pus (B Puttier coll.). Moorestown, 17 May 1963 on R. crispus (EAR coll.). Deerfield, 20 May 1963 on R. crispus (Buck coll.). Mt. Laurel, 25 May 1963 on R. crispus (HWA coll.). Aphis spiraecola Patch, Spiraea Aphid. Whitesbog (Pemberton), 13 July 1961, alatae (possibly “drifts”) on Aronia atropurpurea (Marucci coll. — JOP det.) and 5 apterae on Lyonia ( Pieris ) mariana (Marucci coll.— JOP det.). Haddonfield, 1963 — several plantings of Spiraea sp. in a garden only slightly infested throughout the season. Shiloh (Perkins-deWilde Nursery), 15 May 1963, many on Pyracantha coccinea var. lalandi (Pope coll.). Moorestown, 21 May 1965, terminal growth of Spiraea prunifolia and of Pyracantha sp. mod- erately infested (HWA coll.). Aulacorthum solani (Kalt.)- — see Acyrthosiphon solani (Kalt.). Brachycaudus crataegijoliae (Fitch), formerly Aphis. Old Bridge (Helka Bros.), 12 Aug. 1964, heavy on leaves of Crataegus sp. (Driver coll.). Brevicoryne brassicae (Linnaeus), Cabbage Aphid. “Observed generally throughout the State on cabbage, broccoli, and other cole crops (Ins.-Dis. Newsltr. in CEIR 14(33): 941, 14 Aug. 1964). Calaphis betulaecolens (Fitch) group. Cherry Hill and Haddonfield, 1963 cn Betula lenta (MDL coll. — Richards det.). Calaphis betulella Walsh. Haddonfield, 1961, 1 alate in MT (Gladys Tester- man coll.); 1-15 Aug. 1963, 3 alatae in MT; 16-22 May 1965, 6 alatae in MT; 24 Aug. 2 Sept. 1965, 1 alate in MT (all MDL). Calaphis castaneae (Fitch). Medford Lakes, 6 June 1965, 1 alate and June, 1967] Leonard: New Jersey Aphids 81 several small apterae, the latter whitish with antennae black, on chestnut (HWA coll.). Capitophorus elaeagni (Del Guercio), Oleaster Thistle Aphid. Haddonfield, all 1961, alatae in MT — May 15-25, 3; 1-15 June, 3 and Oct. 4 (Gladys Tester- man coll.) ; 26-30 May 1961, 3 (MDL coll.) ; 1963 — 1-15 Oct., 4, 16-31, 3, Nov. 1-15, 32, 16-30, 21, all males (MDL coll.); 1-15 Nov. 1965, 3 alatae in MT (MDL coll.). Riverton, 14 Oct. 1963 on Elaeagnus umbellata (EAR coll.). Capitophorus glandulosus (Kaltenbach) . Haddonfield, 1963 — The small patches of mugwort reported on during the past several seasons were abun- dantly infested on 26 June but were only very slightly infested during July and early August and none could be found from then on. Nov. 15 one ovipara containing a single egg was blown into the MT and on Nov. 17 several eggs were found on the underside of several lower leaves. None could be found during 1964 nor in 1965 although during this latter year almost all of the mugwort had been pulled out of the garden. Capitophorus hippophaes (Walker), Polygonum Aphid. Wycoff, 14 Oct. 1960, very abundant on a small patch of Polygonum caespitosum var. longi- setum (det. E. C. Leonard, USNM), (MDL & DDL coll.). Moorestown, 1 Aug. 1962, 1 alate in MT (EAR coll.). Haddonfield, 14 Sept. 1963, scarce on Polygonum caespitosum (det. Shetler), (MDL coll.); by daily collecting in MT throughout Oct. and Nov. 1963 at least 100 alatae were obtained, all males (MDL coll.); 29 Aug. and 2 Sept. 1965, scarce on a large patch of P. pennsyl- vanicum (det. Shetler), (MDL & DLW coll.). Capitophorus ribis (L.) — see Cryptomyzus ribis (L.). Cepegillettea myricae Patch. Medford Lakes, 27 Oct. 1963, several plants of Comptonia ( Myrica ) peregrina var. asplenifolia growing in a woods lightly infested (G. G. Rohwer coll.). Chaitophorus sp. Haddonfield, 16-31 Nov. 1963, 1 alate in MT (MDL coll. —ANT det.). Chaitophorus populicola Thomas, Cloudy-winged Cottonwood Leaf Aphid. Haddonfield, 1-15 June 1961, 1 alate in MT (Gladys Testerman coll.). Pitts- grove, 11 Sept. 1963, heavily infested, scattered small aspens, Populus grandi- dentata (C. W. Holsworth, Senior Forester, Parvin State Park coll.). Medford Lakes, 3 June 1965, 1 “stray” alate on laurel (Quinden coll.). Chaitophorus viminicola Hille Ris Lambers. Indian Mills, 20 May 1963 on Salix sp. (HWA coll. — MacGillivray det.). Recorded elsewhere only from Iowa, Illinois, and Pennsylvania. *Chromaphis jugandicola (Kaltenbach), Walnut Aphid. Moorestown, 28 Aug. 1965, fairly common on a large English walnut (MDL & EAR coll.). Cryptomyzus ribis (Linnaeus), (formerly in Capitophorus) . Currant aphid. Moorestown, 26 May 1963 on currant (HWA coll.). Dactynotus spp. The following collections were examined by Dr. Olive 82 New York Entomological Society [Vol. LXXV who was unable to determine them specifically; Ridgewood, July 1963 on Rudbeckia hirta (DDL coll.). Haddonfield, 1, 16 Oct. 1963 on Aster simplex (Shetler det.), (MDL coll.); 1-15 Sept. 1963 in MT (MDL coll.); a number of alatae from mid-Sept. to mid-Oct. 1963 in MT (MDL coll.). Moorestown, 26 Aug. 1963 on Rudbeckia sp. and many specimens on Rudbeckia sp., 30 June 1965 on Solid a go sp. (HWA coll.). Dactynotus ( Dactynotus ) ambrosiae (Thomas), Brown Ambrosia Aphid. Haddonfield, 20 Sept. 1963 and Ridgewood, 24 Oct. 1963 on Ambrosia trifida (MDL coll.— ATO det.). Dactynotus ( Lambersius ) anomalae (Hottes & Frison). The small patch of hardy purple asters in the garden at 217 Rhoads Ave., Haddonfield was moderately infested several times during the 1963 season but most of the colonies dried up. Predators were often present but no parasites were observed. The last collection was made 18-22 Oct. and soon after the plants were mostly dead. No aphids were observed on these plants in 1964 and fairly early in 1965 all the plants were dug out. Dactynotus ( Dactynotus ) chrysanthemi (Oestlund). Medford, 11 Sept. 1963 on Bidens coronata var. trichosperma (EAR coll.). * Dactynotus ( Uromelan ) eupatorijoliae Tissot. Haddonfield, 27, 30 Sept. 1963, fairly common on a small patch of Eupatorium rugosum (Shetler det.), (MDL coll.— ATO det.). * Dactynotus ( Lambersius ) gravicornis (Patch). Haddonfield, 27, 30 Sept. 1963 on Solidago rugosa (MDL coll.— ATO det.). * Dactynotus ( Dactynotus ) leonardi Olive. Ridgewood, July 1963 on Rud- beckia hirta (paratypes) and Aug. 1964 on R. hirta (DDL coll. — ATO det.). Dactynotus ( Dactynotus ) sonchellus (Monell). Indian Mills, 26 May 1963 on Lactuca sp. (HWA coll. — ATO det.). * Dactynotus ( Uromelan ) taraxaci (Kaltenbach), Dark Dandelion Aphid. Cherry Hill, 5 Nov. 1963, a number of dandelion plants, on leaves and some of the stems, heavily infested with apterae in a back yard lawn and on Nov. 11 many more found, including two alates (DLW coll. — ATO det.). Dactynotus ( Lambersius ) tissoti (Boudreaux). Haddonfield, 27, 30 Sept, and 15 Oct. 1963 on Solidago rugosa (Shetler det.), (MDL coll. — ATO det.). Dactynotus ( Uromelan ) tuataiae Olive — Correction to records in “Addi- tional Records” — the data for this species should read as follows: Medford Lakes, 2 Aug. 1962 (G. G. Rohwer coll.) and Moorestown 1 Aug. 1962 (HWA coll.), both on Ambrosia art emisii folia. Drepanaphis sp. Haddonfield, 16-31 Oct. 1963, 26 males in MT (MDL coll. — CFS det. who writes “I cannot identify these at the present time.”). Drepanaphis acerifolii Thomas, Painted Maple Aphid. Haddonfield, 15 Sept. 1963 on Acer rubrum var. trilobum (MDL coll. — CFS det.); 25 Sept. 1963 a large trilobum maple very heavily infested (MDL coll. — CFS det.); 4 June, 1967] Leonard: New Jersey Aphids 83 Oct. many males, oviparae, nymphs and 16 Oct. 1963 males and oviparae on trilobum maple; in MT, 1963 — Aug., 1 alate, 1-15 Sept., 1 alate, 15-30 Sept., 5 alatae, 16-31 Oct., 26 alatae. Cherry Hill, 30 Sept. 1963 apterae on a trilobum maple (MDL & DLW coll.— CFS coll.). *Drepanaphis carolinensis Smith. Haddonfield, 15 Sept. 1963, 1 alate on trilobum maple (MDL coll. — CFS det . ) ; 16-30 Sept. 1963, 4 alatae in MT (MDS coll.— CFS det.). Dr e panaphis parvus Smith. Haddonfield, 16-31 Oct., 1963, 2 alatae in MT (MDL coll. — CFS det.). Second record for New Jersey. *Drepanaphis simpsoni Smith. Haddonfield, 16-31 Oct., 2 alatae in MT (MDL coll.— CFS det.). Eriosoma crataegi Oestlund. Princeton Nurseries, Allentown Farm, 3 Aug. 1964 on Crataegus mollis (Stinson coll.). Dunellen, 18 Aug. 1964, heavy in- festation on Crataegus sp. (Stinson coll.). * Euceraphis lineata Baker. Ridgewood, 29 Oct. 1962 at Duck Pond, oviparae on Betula alba (MDL & DDL coll. — Richards det.). *Eulachnus rileyi (Williams). Haddonfield, 1963 in MT — 1-15 Oct., 1 alate, 2 Oct., 15 alatae, 3-15 Oct., 1 alate (MDL coll. — ANT det.). *Georgiaphis ulmi (Wilson). Bound Brook, 31 May 1963 on leaves and bark of Ulmus sp. (Weber coll. — CFS det.). Hamamelistes spinosus Shimer, Spiny Bud-gall of Witchhazel. Terns River, H. B. Scammell & Son, 10 June 1964 in corrugated leaves of white birch (Pope coll.). Lachnus salignus (Gmelin), Giant willow Aphid. Freehold, 5 Aug. 1963 on Salix sp. (Pope coll.). *Macrosiphonieila millejolii (deGeer). Indian Mills, 26 May 1963, several on Achillea millefolium (HWA coll.). Macro si phoniella sanborni (Gillette), Chrysanthemum Aphid. Haddonfield, 1963 — a small patch of chrysanthemums in a garden was uninfested until about mid-Oct. when some colonies began to appear. Moorestown, 21 May 1965, “hardy'’ mums lightly infested (HWA coll.). Macrosiphum spp. Haddonfield, 21 Oct. 1963 on Mentha spicata (MDL coll. — ATO det.); alatae in MT — 1-15, 16-30 Sept, and 1-15 Oct. 1963 (MDL coll. — ATO det.); a number of specimens were collected. Macrosiphum dirhodum (Wlk.) — see Metopolophium dirhodum (Wlk.). Macrosiphum euphorbiae (Thomas), Potato Aphid. Cherry Hill, 6 May 1962, 1 alate, several young on Euonymus europaeus (DLW coll.). Moores- town, 2 Aug. 1962 on tomato in a garden (HWA coll.); 21 May 1965, a single mature aptera among many Aphis fabae on rhubarb (HWA coll.). Medford Lakes, 28 May 1963, heavy infestation on cult, roses (G. G. Rohwer coll.). Medford, 31 May 1963 and 17 June 1964, light on tomato (Quinden coll.). Mt. Laurel, 25 May and Indian Mills, 26 May 1963 on Apocynum cannabinum 84 New York Entomological Society [Vol. LXXV (HWA coll.). Linwood, 29 April 1963 light on Tulipa sp. and Shiloh, 15 May 1963, heavy on cult, roses (Buck coll.). Haddonfield, 16-31 Aug., 1 alate and 1-15 Nov., 6 alatae 1963 in MT (MDL coll. — ATO det. ) ; 9-22 May 1965, many alatae in MT (MDL coll. — JOP det.). McGuire Air Force Base, mid- May 1965, several apterae on cult, rose (Quinden coll.). POTATO APPIID ( Macrosiphum euphorbiae ) — NEW JERSEY — Survey at 25 sites in Cumberland, Salem, Gloucester, Burlington, Mercer, Monmouth, and Middlesex counties revealed smaller number of eggs than in 1964; however, percentage of viable eggs higher. Counts higher in Mercer, Monmouth, and Cumberland counties. Table below gives total number of eggs found and per- centage which were viable at time of survey, for last 9 years. (Ins.-Dis. Newsltr.). (CEIR 15( 19): 447, 1965). Comparison of Total Number of Eggs and Percentage of Viable Eggs Year 1957 1958 1959 1960 1961 1962 1963 1964 1965 Total No. Eggs 427 226 1522 178 713 411 745 1192 774 Percent Viable 65.6 53.1 54.7 74.7 25.2 74.2 78.3 45.6 73.6 Note: It should be pointed out that the egg surveys are made on plants of the swamp rose (Rosa palustris). Macrosiphum liriodendri Monell, Tuliptree Aphid. The following collections were inadvertently omitted from “Additional Records”: Montclair, 27 May 1954 (Bartlett Tree Research Laboratories). New Brunswick, 10, 26 June 1962 (Wave coll. — CFS det.). Moorestown, 19 July 1962 (HWA coll.) and 29 July 1962 (det. W. Jones coll.). Oldwick, 8 Aug. 1960 (Wave coll.). West- mont, 28 June 1962 (J. J. Earley of PPCD, USDA coll.). The following collections were made 1963-1965, all on tuliptree unless other- wise specified: Waterford, 26 May 1963, immatures on Magnolia virginiana (HWA coll.— ATO det.). Summit, 22 June 1963, common on a very large tree (MDL coll.). Moorestown, scarce, 31 July 1963 (EAR coll.) and 21 May 1965 (HWA coll.). Haddonfield, 24, 31 July, 19 Aug., 13 Sept., and 27 Aug. 1965, a small street shade tree lightly infested when examined on each of these dates (MDL & DLW coll.), Medford Lakes, 6 June 1965, about 35 apterae of various sizes on Magnolia virginiana (HWA coll.). Macrosiphum rosae (Linnaeus), Rose Aphid. Haddonfield, no aphids could be found on roses at 217 Rhoads Ave., except a few in mid-May, until late Sept. 1963 when the tender shoots on one large bush became heavily infested; several alatae in MT, 16-30 Sept. 1963 (MDL coll. — ATO det.); during May, Aug., and in late Dec. 1965 no aphids could be found. Moorestown, 27 Dec. 1965 EAR collected 3 mature apterae and several young on a rose cutting which had been taken indoors — the weather had been unseasonably mild. 'l'Masonaphis sp. Cooper Creek, Haddonfield, 2 Sept. 1965 alatae on Boehmeria cylindrica (MDL & DLW coll. — MacGillivray det. who states “I cannot place these in any species known to me.”). June, 1967] Leonard: New Jersey Aphids 85 Masonaphis ( Ericobium ) azaleae (Mason). Philip E. Marucci, Cranberry and Blueberry Research Laboratory, N. J. Ag. Exp. Sta., New Lisbon wrote me on 8 Oct. 1965 that Leon Coles’ statement in my “Additional Records” in regard to light parasitism by Aphelinus sp. needs correction. Marucci says “ Aphelinus is a very effective parasite of M. azaleae in the field. Last year the ratio of mummified aphids to live aphids was 50 to 1 and this year it is about 71 to 1.” Ben Puttier writes me, 11 Jan. 1966 that it was he who originally identified this parasite as undoubtedly Aphelinus semijlavus How. but that no specimens were preserved. A hyperparasite reared from this aphid from Lebanon State Forest by Marucci was determined by Paul M. Marsh, USNM, as Logocerus niger (How.). It has been recorded as parasitizing a species of Aphidius. Melanocallis caryaejoliae (Davis), Black Pecan Aphid. (Richards, Mem. Ent. Soc. Can. 44: 102, 1965 places this in Tinocallis) . Moorestown, 1 Aug. 1962, 2 alatae in MT (EAR coll.). *Metopolophium dirhodum (Walker), Rose Grass Aphid. (Has also been placed in Acyrthosiphon and Macrosiphum) . Ridgewood, 28 Oct. (MDL coll.) and Summit (MDL & DDL coll.), 29 Oct. 1965, rose bushes in the garden had a number of leaves, each bearing on the underside a single (occasionally two) alate, each with a number of newly born young nearby. This is the first time these fall migrants have been noticed on roses in New Jersey (MacGilli- vray det.). Monellia caryae (Monell), American Walnut Aphid. (Richards, Mem. Ent. Soc. Can. 44: 99, 1965, places this in Monelliopsis) . Ft. Lee, 2 July 1909 on Juglans nigra (Gillette in Jour. Econ. Ent. 3(4): 367, 1910). Moorestown, 19 July, 1 Aug. 1962, 10 alatae in MT (EAR coll. — Bissell det.). Haddonfield, 29 Aug. 1965, fairly common on several large black walnut trees (MDL & DLW coll. — Bissell det.). Monellia caryaella Fitch. Moorestown, 23 May 1962, very scarce on large Juglans nigra (MDL & EAR coll. — Richards det.). Monellia costalis (Fitch), Black-margined Aphid. Haddonfield, 30 May 1947 on Carya sp. (MDL coll. — Bissell det.); 1-15 Sept. 1963, 1 alate in MT (MDL coll. — Richards det.). Monellia nigropunctata Granovsky. Haddonfield, 30 May 1947 on Carya sp. (MDL coll. — Bissell det.). Myzocallis alhamhra Davidson, Western Dusky-winged Oak Aphid. (Rich- ards, Mem. Ent. Soc. Can. 44: 57, 1965 considers this as merely a melanistic form of M. punctata (Monell). (This species was in the Plant List but not in the Aphid List of “Additional Records”). Haddonfield, 30 May 1947, “drift” alatae on chestnut (MDL coll.); 23-31 July, 1 alate, 16-31 Aug., 6 alatae and 1-5 Sept., 6 alatae — all 1963 in MT (MDL coll.). New Brunswick, 15 July 1960, a “drift” alate on Ulmus americana (Wave coll.). Moorestown, 1 Aug. 1962, 1 alate in MT (EAR coll.) and 26 Aug. 1965, 1 alate in MT 86 New York Entomological Society I V ol. LXXV (EAR coll.). Ridgewood, 18-22 June 1963, 1 alate in MT (MDL coll.). Sum- mit, 22 June 1963 on Quercus rubra (MDL & DDL coll.). Myzocallis bella (Walsh), Haddonfield, Aug. 1963, 3 alatae in MT (MDL coll. — ANT det.). * Myzocallis exultans Boudreaux & Tissot. Haddonfield, 15-17 Sept. 1963, l alate mixed in with several M. jrisoni B & T on a small pin oak street tree (MDL coll. — ANT det.); Aug. 1963, 3 alatae in MT (MDL coll. — ANT det.). Medford Lakes, 3 June 1964, 1 “stray” alate on laurel (Quinden coll.). * Myzocallis jrisoni Boudreaux & Tissot. Haddonfield, 15, 27 Sept. 1963, alatae, nymphs, 3 oviparae from several small moderately infested pin oaks (MDL coll. — ANT det.); 16-31 Aug., 4 alatae and 1-15 Sept. 1963, 4 alatae and 2 alatae, 24 Aug.-2 Sept. 1965 in AIT (MDL coll. — ANT det.); 25-28 Aug. 1965 several large colonies on a pin oak (MDL coll.). Myzocallis melanocera Boudreaux & Tissot. Haddonfield, Aug. 1963, 2 alatae in MT (AIDL coll. — ANT det.). Myzocallis multisetis Boudreaux & Tissot. Haddonfield, 1-15 1963, 2 alatae in MT (MDL coll.— ANT det.). Myzocallis punctata (Monell), Clear-winged Oak Aphid. Haddonfield, 16- 30 Sept. 1963, 1 alate in MT (MDL coll.). Myzocallis tiliae (Linnaeus), Linden Aphid. Haddonfield, 16-31 Aug., 1 alate, 1-15 Sept., 2 alatae, and 1-15 Nov., 1 alate 1963 all in MT (MDL coll.). Aloorestown, 28 Aug. 1965 scarce in a T ilia europaea (MDL & EAR coll. ) . Myzocallis ulmifolii (Monell), Elm Leaf Aphid, (Richards, Mem. Ent. Soc. Can. 44: 104, 1965 places this in Tinocallis) . Princeton, 22 Sept. 1965 common on Ulmus sp. (Weber coll.). Myzocallis walshii (Monell). Cherry Hill, 30 Sept. 1963, a few leaves of a large Quercus velutina lightly infested, alatae and nymphs present (MDL & DLW coll. — ANT det.). Haddonfield, Aug. 1963, 2 alatae and 16-31 Oct. 1963, 2 alatae in AIT (MDL coll. — ANT det.). Myzus cerasi (Fabricius), Black Cherry Aphid. Helmetta, 5 Aug. 1964, light on leaves of Kwanzan cherry, Prunus sp. (Driver coll.). Moorestown, 21 Alay 1965, a cult, sour cherry, Prunus cerasus , lightly infested (HWA coll.). *Myzus dianthi Schrank, Carnation Aphid. In my “Additional Records” Myzus polaris Hille Ris Lambers is recorded from Weston, 5 April 1946 on carnation (F. S. Smith coll., 1 slide in USNM). It has since been found that this is the carnation aphid. This aphid and/or Myzus persicae presumably occurs on carnations in New Jersey but no collections (other than the above) have been made to substan- tiate the presence of either. However, on 31 Dec. 1965 I visited a florist in Barrington who had a large glass house of carnations. Unfortunately time did not permit me to examine any of the plants but I was told by the production June, 1967] Leonard: New Jersey Aphids 87 foreman that small infestations of a small greenish aphid occasionally appeared but were readily held in check by timely applications of an insecticide. Myzus persicae (Sulzer), Green Peach Aphid. Moorestown, 19 Sept. 1962, many on Cleome spinosa (MDL & EAR coll.); omitted from “Additional Rec- ords.” Linwood, 4 June 1963, a heavy infestation on Anthurium sp. (Sohl coll.). Mrs. Sohl writes that “the tips and flower stems of the new growth of many plants growing in a greenhouse were heavily infested and that the leaves and flowers were affected by slight crinkling and/or gnarling.” I can find no previous record of any aphid on this plant. Haddonfield, 1963, alatae in MT; 23-31 July, 2; 1-15 Aug., 5; 16-31 Aug., 1; 1-15 Sept., 7; 16-30 Sept., 1; 1-15 Oct., 3; (MDL coll.); 24 Aug.-2 Sept. 1965, 25 alatae in MT (MDL coll.). New Jersey — “Heavy flight noted throughout State during past week. Con- trol recommended for peppers and tomatoes.” (Ins.-Dis. Newsltr. in CEIR 15(32): 897, 6 Aug. 1965). New Jersey — “Increasingly important on broccoli in southern area; controls recommended.” (Ins.-Dis. Newsltr. in CEIR 15(34): 968, 20 Aug. 1965). Ridgewood, 27 Oct. 1965, the buds and stems moderately infested in a large house of ’mums (Schweinfurth’s Florists), (MDL & DDL coll.). Barrington, 31 Dec. 1965, a large greenhouse of ’mums very lightly infested. The propaga- tion foreman told me that occasional spraying readily held the aphids in check. Summit, 30 Oct. 1965, a very few apterae on an indoor plant of Jerusalem cherry (MDL coll.). P. E. Marucci wrote me on 8 Oct. 1965 “I am sure Myzus persicae often invades strawberries. Last year a very heavy infestation of peppers overflowed into adjacent strawberries and the population was so high that the grower found it necessary to spray for them.” On 20 May 1965 E. A. Richmond found a number of plants of Duranta repens moderately infested in the Mall, a large enclosed shopping center at Cherry Hill. The writer and Dr. Richmond examined these plants together on 28 Aug. At this time no aphids could be found but the leaves were rather heavily infested with a whitefly. I find only one previous record of the oc- currence in the USA of this aphid on this plant. In 1900 Gillette and Taylor published Colorado Agr. Exp. Sta. Bull. 133 entitled “A few orchard plant lice.” In the discussion of Myzus persicae a list of plants is given on which this aphid had been found establishing colonies in the greenhouses (presumably at Ft. Collins). One of the plants listed is Duranta plumieri (now re pens). It has been reported elsewhere from Egypt and Israel. N eoceruraphis viburnicola (Gillette), (formerly in Anuraphis) , Snowball Aphid. Haddonfield, 13 Nov. 1963, 1 viviparous alate and several oviparae on a large Viburnum opulus (MDL coll.). Moorestown, 21 May 1965, a Viburnum sp. heavily infested with heavily parasitized aphids (HWA coll.); many para- 88 New York Entomological Society [Vol. LXXV sites emerged within the next four days in a covered box and were identified by Paul Marsh, USNM as Lysephlebius testaceipes (Cresson). N eoprociphilus aceris (Monell). Chatham, 21 May 1963 on sugar maple, woolly aphids (Weber coll.). Ovatus crataegarius (Walker), Mint Aphid. Medford, 28 July 1963, 34-40 apterae; 18 May 1965, 3 alatae, several “pupae” and apterae; 6 June 1965, several alatae and apterae, 4 of the latter obviously parasitized; a small dip- terous larvae also present — all on mint and coll, by Quinden. Periphyllus calif or niensis Shinji. Haddonfield, 9-15 May 1965, 540 alatae in MT most of which came to the yellow pan in the first 4 days. The total number of aphids in the pan during the week was 1336 of which this aphid constituted about 40%. 16-22 May 1965, 15 alatae of this aphid in the MT out of a total of 647 aphids. This species was described from California and has been recorded elsewhere from Washington and Pennsylvania and once before, with a query, in New Jersey. It is recorded as feeding on Japanese maple. Periphyllus negundinis Thomas, Boxelder Aphid. Moorestown, 24 Sept. 1963, scarce on a large boxelder (MDL & HWA coll.); 21 May 1965, a box- elder heavily infested and leaves sticky with honeydew (HWA coll.). Phyllaphis fagi (Linnaeus). Bound Brook, 31 May 1963 on beech (Weber coll. — CFS det.). Haddonfield 1963 — the copper beech at 213 Rhoads Ave. only very slightly infested when first observed on 28 June and continued so until into Oct. at which time somewhat more were present and 9 alatae were obtained; 1965 — in mid-May this tree was heavily infested and sticky with honeydew; alatae scarce on leaves but infested leaves placed in a closed box soon produced many alatae. Rhopalosiphum maidis (Fitch), Corn Leaf Aphid. Bridgeton, 30 July 1963, a heavy infestation on the stalks of corn (Sohl coll.). Rhopalosiphum nymphaeae (Linnaeus), Waterlily Aphid. Saddle River, 17 June 1965, a heavy infestation on pond lilies in a greenhouse (Wm. Tricker, Inc.), (Condon coll.), Chatsworth, 26 Sept. 1965 on Nuphar advena , a num- ber of plants considerably infested (HWA coll.). Rhopalosiphum serotinae Oestlund. Waterford, 26 May 1963 on Solidago sp., 17 apterae (HWA coll.). *Schizolachnus piniradiatae (Fabricius). Boonton, 22 July 1964 on Pinus resinosa (Kegg coll.). *Therioaphis maculata (Buckton), Spotted Alfalfa Aphid. In regard to the first find of this aphid in New Jersey L. Donald DeBlois, Entomologist, Divi- sion of Plant Industry, N. J. Dept. Agr. wrote me on May 6, 1965 as follows: “The collections were made in the course of a survey for this insect during the fall of 1964 in 95 alfalfa fields throughout New Jersey. Rough sorting of the collections was done here and final identifications were made by Louise June, 1967 Leonard: New Jersey Aphids S9 Russell. The spotted alfalfa aphid collections were made by one of our in- spectors, G. Robert Glass. One alate and one apterous viviparous female was taken in Greenwich in Cumberland County on September 24, 1964. We will be making extensive surveys throughout the State to determine the extent of the infestation.’’ 1965 — Cape May County: Woodbine 29 Nov., 7 apterae. Cumberland County: Canton 23 Sept., 100 apterae; Greenwich 11 apterous viviparae, 1 ovipara; Jones Island 17 Nov., 4 apterae; Rhoadstown 23 Sept., 9 apterae; Shiloh 23 Nov., 5 apterae. Gloucester County: Jefferson 10 Dec., 1 aptera; Mullica Hill 23 Nov., 1 aptera; Pitman 23 Nov., 1 aptera. Salem County: Alloway 21 Sept., 1 aptera; Centerton 20 Sept., 4 apterae; Elmer 20 Sept., 10 apterae; Hancock’s Bridge 23 Sept., 8 apterae. All collections were made on alfalfa by G. R. Glass and submitted by L. D. DeBlois both of the N. J. Dept. Agr. Trenton, N. J. Determinations by Louise M. Russell, Ent. Res., USDA, Washington, D. C. ( Therioaphis trifolii (Monell), Yellow Clover Aphid. Ben Puttier wrote me on 11 Jan. 1966 that he has taken Aphelinus semiflavus Howard from this aphid in New Jersey.) FOOD PLANT LIST* Acer negundo (Boxelder) Periphyllus negundinis Acer rubrum var. trilobum Dr e panaphis acerifolii Dr e panaphis car olinensis Acer saccharum (Sugar or Hard Maple) Drepanaphis acerifolii N eoprociphilus aceris * Achillea millefolium (Common Yarrow) Macro sip honiella millefolii Alfalfa — see Medicago Ambrosia trifida (Giant Ragweed) Dactynotus ambrosiae * Anthurium sp. Myzus persicae Apocynum cannabinum (Dogbane) Macro sip hum euphorbiae *Aquilegia longissima (Longspur Colum- bine) Aphis gossypii Arctium sp. (Burdock) Aphis fabae Arctium minus (Common Burdock) Aphis fabae *Aronia atro purpurea Aphis spiraecola Artemisia vulgaris (Mugwort) Capitopho ru s gland ulo sus Aspen — see Poplus grandidentata Aster novae-angliae (New England or Hardy Purple Aster) Dactynotus anomalae ■'-Aster simplex Dactynotus sp. * Azalea viscosa (Swamp Azalea) ? Aphis gossypii Beech — see Fagus Betula alba (European White Birch) Eucer aphis lineata Hamamelistes spinosa Betula lenta (Black Birch) C ala phis betulaecolens group *Bidens coronata var. trichosperma Dactynotus chrysanthemi Birch — see Betula Blackcved Susan— -see Rudbeckia hirta Blueberry — see V accinium corymbosum *Boehmeria cylindrica Masonaphis sp. Boxelder — see Acer negundo Brassica oleracea var. botrytis (Broccoli) Brevicoryne brassicae * Plants marked with an asterisk (*) are additions to the two previous lists. 90 New York Entomological Society [ Vol. LXXV Myzus pevsicae Brassica oleracea var. capitata (Cabbage) Brevicoryne brassicae Broccoli — see Brassica oleracea var. botry- tis Burdock — see Arctium Cabbage — see Brassica oleracea var. capi- tata Camp sis ( Tecoma ) radicans (Trumpet Creeper) Aphis gossypii Capsicum jrutescens (Redpepper) Myzus persicae Carnation — see Dianthus Cary a sp. (Hickory) Monellia costalis Monellia nigro punctata Castanea dentata (American Chestnut) Calaphis castaneae Catalpa sp. A phis gossypii C haenomeles sp. (Flowering Quince) Aphis pomi Cherry, Sour — see Prunus cerasus Chestnut — see Castanea Chinese Scholar Tree— see Soph ora japo- nic a Chrysanthemum sp. Macro si phoniella sanborni Myzus persicae Cleome spinosa Myzus persicae Columbine — Aquilegia C omptonia ( Myrica ) peregrina var. aspleni- folia (Sweetfern) Cepegillettea myricae Corn — see Zea Cowlily — see Nuphar advena Crab, Flowering — Mains sp. Crataegus sp. (Hawthorn) Brachycaudus crataegif oliae Eriosoma crataegi * Crataegus mollis Eriosoma crataegi Currant — see Ribes Dandelion — see Taraxacum Dianthus caryophyllus (Carnation) Myzus diant hi Dock, Curled — see Rum ex c.rispus Dogbane — see A pocynum * Durant a repens (Golden Dewdrop) Myzus persicae Elaeagnus umbellata Capitophorus elaeagni Elm — see Ulmus English Ivy — Hedera Euonymus europaeus (European Spindle- tree) A phis jabae M aero si ph u m euphorbiae *Eupatorium rugosum (White Snakeroot) Dactynotus eupatorif oliae Evening Primrose — see Oenothera Fagus sp. ( Beech) Phyllaphis jagi Fagus sylvatica var. purpurea (Copper or Purple Beech) Phyllaphis jagi Firethorn — see Pyracantha Quince, Flowering — see Chaenomeles *F orsythia sp. Aphis gossypii Fragaria sp. (Strawberry) Myzus persicae Geranium — see Pelargonium Golden Dewdrop — see Duranta Goldenrod — see Solidago Hawthorn — see Crataegus Hedera helix (English Ivy) Aphis pseudohederae *H elianthus annuus (Common Sunflower) Aphis helianthi Hibiscus syriacus (Rose-of-Sharon) Aphis gossypii Hickory — see Carya Ipomoea batatas (Sweet Potato) Myzus persicae Jerusalem Cherry— see Solanum pseudo- capsicum Juglans nigra (Black Walnut) Monellia caryae Monellia caryaella * Juglans regia (English or Persian Walnut) C hromaphis juglandicola Lactuca sp. (Lettuce) Dactynotus sonchellus Lettuce— see Lactuca *Leucothoe racemosa Aphis gossipyi Linden — see Tilia Liriodendron tulipifera (Tuliptree) Macro sip h um lirio den dri June, 1967 I Leonard: New Jersey Aphids 91 Ly coper sicon esculentum (Tomato) Macro si phu m euphorbiae Myzus persicae Lyonia ( Pieris ) mariana (Stagger bush) Aphis spiraecola * Magnolia virginiana (Sweetbay) Macrosiphum liriodendri Mains sp. (Flowering Crab) Aphis pomi Maple, Hard or Sugar — see Acer saccharum Medicago sativa (Alfalfa) Acyrthosiphon pisum Therioaphis maculata Mentha sp. (Mint) Ovatus crataegarius Mentha spicata (Spearmint) Macrosiphum sp. Mint — see Mentha Mockorange — see Philadelphus Mugwort — see Artemisia vulgaris Nasturtium — see Tropaeolum *Nuphar advena (Cowlily) Rhopalosiphum nymphaeae Nyssa sylvatica (Tupelo) Aphis coreopsidis Oak — see Quercus Oenothera sp. (Evening Primrose) Aphis oestlundi Oenothera biennis (Common Evening Prim- rose) Aphis oestlundi * Pelargonium sp. (Geranium) Acyrthosiphon pelargonii Peonia sp. Aphis gossypii * Philadelphus sp. (Mockorange) Aphis jabae Pine — see Pinus Pinkweed — see Polyonum pennsylvanicum * Pinus resinosa (Red Pine) Schizolachnus piniradiatae * Polygonum cae spit o sum Capit op horns hippo phaes * Polygonum caespitosum var. longisetum Capit ophorus hippo phaes Polygonum pennsylvanicum (Pinkweed) Capit ophorus hippo phaes Populus grandidentata (Aspen) Chait ophorus populicola Primus sp. (Kwanzan Cherry) Myzus cerasi Primus cerasus (Sour Cherry) Myzus cerasi Pyracantha sp. (Firethorn) Aphis spiraecola Pyracantha coccinea var. lalandi (Laland Firethorn) Aphis spiraecola Quercus palustris (Pin Oak) Myzocallis exult ans Myzocallis jrisoni Quercus rubra (Red Oak) Myzocallis alhambra Quercus velutina (Black Oak) Myzocallis walshii Ragweed — see Ambrosia Red Clover — see Trifolium pratense Redpepper — see Capsicum Rheum rhaponticum (Rhubarb) Aphis fabae Macrosiphum euphorbiae Rhubarb — see Rheum Ribes sp. (Currant) Cry ptomyzus ribis Rosa sp. (Rose) Acyrthosiphon porosum Macrosiphum euphorbiae Macrosiphum rosae M eto polo p hiu m dir hod u m Rosa palustris (Swamp Rose) Macrosiphum euphorbiae Rose of Sharon — see Hibiscus Rudbeckia ( serotina ) hirta (Blackeyed Susan) Dactynotus sp. Dac.tynotus leonardi Rumex crispus (Curled Dock) Aphis rumicis Salix sp. (Willow) ? Chait ophor us viminicola Lachnus salignus Snakeroot — see Eupatorium rugosum ■'-Solanum pseudocapsicum (Jerusalem Cherry) Myzus persicae Solidago sp. (Goldenrod) Dactynotus sp. Rhopalosiphum serotinae Solidago rugosa Dactynotus gravicornis Dactynotus tissoti *Sophora japonica (Chinese Scholar Tree) 92 New York Entomological Society I Vol. LXXV Aphis gossypii Spiderf lower — see Cleome Spindle Tree, European — see Euonymus Spiraea sp. A phis spiraecola * Spiraea prunifolia (Bridalwreath Spiraea) Aphis spiraecola Staggerbush — see Lyonia Strawberry — see Fragaria Sunflower — see Helianthus Sweetbay — see Magnolia virginiana Sweetfern — see Comptonia Sweetpotato — see Ipomoea Taraxacum officinalis (Common Dandelion) Acyrt h o sip h o n sola n i Dactynotus taraxaci * Tilia europaea (European Linden) Myzocallis tiliae Tomato — see Ly coper sic on Trifolium pratense (Red Clover) Acyrt hosiphon pisum Acknowledgments As in the past a number of persons, in addition to the writer (MDL), made several to a number of collections each. These included again: Drs. Harry W. Allen (HWA) and E. Avery Richmond (EAR) of Moorestown, Gregory G. Rohwer of Medford, Marie C. Quinden of Medford Lakes, Donald D. Leonard (DDL) of Ridgewood and David L. Winters (DLW) of Haddonfield. Several members of the Staff of the Division of Plant Industry, New Jersey Department of Agriculture collected — -Wm. F. Condon, Addison Driver, Frank N. Pagliaro, Geo. L. Pope, Francis S. Stinson, W. A. Junghans, J. D. Kegg, and Paul V. V. Weber as well as B. K. Buck and Irene H. Sohl of the Plant Pest Control Division, A.R.S., U. S. Department of Agriculture. Determinations, other than those by the author, were made by: Miss Louise M. Russell (LMR), Ent. Res. Div., A.R.S., U.S.D.A.; Dr. A. Tom Olive (ATO), Wake Forest College, Winston-Salem, N. Car.; Dr. Clyde F. Smith (CFS), North Carolina State University, Raleigh, N. Car.; Prof. John O. Pepper (JOP), Pennsylvania State University, University Park, Pa.; Prof. Theo. L. Bissell, University of Maryland, College Park, Mary- land; and Dr. Archie N. Tissot (ANT), University of Florida, Gainesville, Florida. Dr. Stanwyn G. Shetler, Dept. Botany, U. S. National Museum, Washington, D. C. kindly made several determinations of plants. To all of the above — those who collected and those who determined — I extend my sincere thanks for their help. Dr. John B. Schmitt has been kind enough to oversee the preparation of the final typescript of this paper. Tropaeolum sp. (Nasturtium) Aphis fabae Trumpet creeper — see Campsis Tulipa sp. Macrosiphum euphorbiae Tulip Tree — see Liriodendron Tupelo — see Nyssa Ulmus sp. (Elm) Georgia phis ulmi Myzocallis ulmifolii Urtica sp. (Nettle) Acyrt ho sip h o n si be ric u m V accinium corymbosum (cult. Highbush Blueberry) Aphis gossypii Masonaphis azaleae Viburnum sp. N eoceruraphis viburnicola Willow — see Salix Yarrow — see Achillea Zea mays (Corn) Rhopalosi phum maidis Received for Publication February 1, 1966 Further Studies on the Internal Anatomy of the Meloidae. III. The Digestive and Reproduetive Systems as Bases for Tribal Designation of Pseudomeloe miniace omaculata (Blanchard)* (Coleoptera: Meloidae) A. P. Gupta Department oe Entomology and Economic Zoology Rutgers-The State University, New Brunswick, New Jersey Abstract: The digestive and reproductive systems of Pseudomeloe miniace omaculata (Blanchard) has been described. On the basis of such internal anatomical features as V- shaped folds in the stomodaeal intima, absence of a basal spermathecal diverticulum, a tubular female accessory gland, an irregularly convoluted first pair and a recurved or bent second pair of male accessory glands, this genus is placed in the tribe Eupomphini of the subfamily Meloinae. The inclusion of Pseudomeloe in Eupomphini now extends the distribution of this tribe to South America as well. In 1928, Van Dyke defined the tribe Calospastini (= Eupomphini) and stated that “the tribe is restricted to North America.” Gupta (1965) showed that all the members of this tribe shared several internal anatomical features. On examination, the South American blister beetle, P. miniace omaculata was found to possess all the characteristic tribal features of Eupomphini, as defined by the present writer (1965). The purpose of the present paper is to describe the internal anatomy of this beetle, and to establish its inclusion in the tribe Eupomphini. The beetles were collected and identified by Dr. Antonio Martinez, Buenos Aires, Argentina, and were kindly made available to the author by him. MATERIALS AND METHODS For technical details, the reader is referred to the earlier work (Gupta, 1965). In the present paper, descriptions have been kept to the minimum, and are meant to supplement the diagrams, and point out important features. In the drawings of the reproductive systems, only the organs of one side have been shown. In the drawing of the male reproductive system, the second pair of accessory glands has been stippled to distinguish it from others. Phase con- trast photomicrographs of the stomodaeal intima are included for the first time in this series of papers. All photomicrographs were taken by Leitz dark phase microscope at magnifications of 250X and 400X. For this purpose, the intima was lightly stained in azocarmine. DESCRIPTIONS DIGESTIVE SYSTEM: EXTERNAL (Fig 1): Esophagus much broadened posteriorly ; ventriculus with few remnants of transverse wrinkles; lobes of pyloric valve barely visible externally; six malpighian tubules arising * Paper of the Journal Series, Agricultural Experiment Station, Rutgers-The State Uni- versity, New Brunswick, New Jersey, U.S.A. 93 94 New York Entomological Society [Vol. LXXV POFL CO SPDU Fig. 1. Lateral view of alimentary canal. Fig. 3. Female reproductive system, dorsal view. Fig. 2. Internal view of stomodaeum. Fig. 4. Male reproductive system, ventral view. Abbreviations Used in Figures ACF accessory folds CO colon EJDU ejaculatory duct FAG female accessory gland IL ileum 1MAG . . first pair of male accessory gland 2MAG . second pair of male accessory gland 3MAG . third pair of male accessory gland MAL malpighian tubules OF esophagus OV ovary PFL lateral primary fold PFMD .... median dorsal primary fold PFMV .... median ventral primary fold POFL posterior flexure POIN . . . posterior intestine or rectum PROV proventriculus PY pylorus PYL lobes of pyloric valve SCLC sclerotized channel SFDL dorsolateral secondary fold SFVL .... ventrolateral secondary fold SPCA spermathecal capsule SPDU spermathecal duct TE testis TF tertiary fold VA vagina VD vas deferens VF V-shaped fold VS vesicula seminalis June, 1967 Gupta: Meloidae Internal Anatomy 95 Fig. 5. Magnified view of stomodaeal intima showing emarginate thickenings provided with microscopic spines (arrows). Fig. 6. Magnified view of portion of median ventral primary fold showing stout spines. 96 New York Entomological Society [Vol. LXXV Fig. 7. Magnified view of portion of median primary fold and transverse corrugations (arrows) . Fig. 8. Magnified view of portion of sclerotized channel showing irregular rectangular and polygonal patterns. June, 1967] Gupta: Meloidae Internal Anatomy 97 separately, their posterior attachment at inner bend of posterior flexure, basal swelling absent. INTERNAL (Figs. 2, 5-10) : Stomodaeal intima with 4 primary, 4 V-shaped, 4 secondary and 8 tertiary folds, several irregularly arranged accessory folds present in regions of esophagus and proventriculus ; transverse corrugations discontinuous; V-shaped folds continued posteriorly into primary stomodaeal lobes and flanking sclerotized channels, latter more sclerotized than those flanked by secondary and tertiary folds, latter flanking sclerotized channels between secondary and V-shaped folds in proventricular region, surface of stomodaeal intima with emarginate thickenings provided with microscopic spines, spines on primary, V-shaped and secondary folds stout, spines also present on apices of stomodaeal lobes, surface of sclerotized channels with irregular rectangular and polygonal pattern without spines. Stomodaeal valve with 4 primary lobes, secondary and tertiary lobes poorly developed. REPRODUCTIVE SYSTEM: FEMALE (Fig 3): Spermathecal capsule robust, constricted near base, portion beyond constriction broadened, rather wrinkled, tapering distally, portion below constriction rounded and smooth, spermathecal duct short and curved; accessory gland tubular, elongate, tapering distally, and with a short duct; vagina very short. MALE (Fig. 4): Testes rather large, spherical, vas deferens narrow near testis, vesicula seminalis rather narrow; first pair of accessory gland ovally or spherically coiled, second pair smallest and recurved distally, recurved portion shorter than basal portion, third pair larger than second and convoluted; ejaculatory duct slightly broader beyond middle, very strongly bowed and bent distally. material examined i 7 specimens (in 8% formaldehyde), Pcia. de Buenos Aires, Partido de Puan, Estacion Felipe Sola, 1-31-1966 (A. Martinez). tribal designation: Fairmaire and Germain first established the genus Pseudomeloe in 1863 (Borchmann, 1917). Beauregard (1890) grouped this genus, among others, with Meloe, Megetra and Cysteodemus in the category of “Meloites.” Later, Borchmann (1917) and Blackwelder (1945) also grouped Pseudomeloe with several presently recognized eupomphine genera in the tribe Meloini. Denier’s (1935) tribe Lyttini also consisted of Pseudomeloe and such genera as Tetraonyx, Pyrota, Lytta, Meloe and several of the current eupomphine genera. As far as is known, there is no mention of the inclusion of Pseudomeloe in the tribe Calospastini (= Eupomphini), after this tribe was first established by Van Dyke in 1928. He included Calospasta ( — Eupompha ), Tegrodera, Gynae- comeloe, Cysteodemus, Megetra, Pleurospasta, Phodaga, Negalius, Cordylospasta and Brachyspasta in this tribe. Gupta (1965) demonstrated that members of this tribe, as constituted by Van Dyke, show such common features as V- shaped folds in the stomodaeal intima, a spermathecal capsule without a basal diverticulum, a tubular female accessory gland, an irregularly convoluted first pair of male accessory glands, and a recurved or bent second pair. He further stated that on the basis of the number of V-shaped folds, and tertiary intimal folds, the tribe can be divided into 2 groups: one group with 3 V-shaped folds and 6 tertiary folds ( Phodaga and Negalius ), and the other with 4 V-shaped folds and 8 tertiary folds ( Eupompha , Tegrodera, Gynaecomeloe, Cysteodemus, Megetra and Pleurospasta) . He did not study Cordylospasta and Brachyspasta. 98 New York Entomological Society [Vol. LXXV Fig. 10 Fig. 9. Magnified view of portion of V-shaped fold showing spines. Magnified view of tip of one of the primary stomodaeal lobes showing spines. June, 1967] Gupta: Meloidae Internal Anatomy 99 Examination of the internal anatomy of Pseudomeloe revealed that it possesses all the characteristic features of Eupoinphini, as defined by Gupta, and belongs to the group with 4 V-shaped and 8 tertiary folds. Its inclusion is the tribe Meloini cannot be justified since it does not possess a well-developed vesicular spermathecal diverticulum, and a reduced 1st pair of male accessory glands, features which are characteristic of the tribe Meloini. Similarly, the presence of V-shaped folds and the absence of a well-developed spermathecal diverticulum precludes its inclusion in Lyttini. The placement of Pseudomeloe in Epicautini, Tetraonycini, and Pyrotini on the basis of V-shaped folds alone cannot be justified inasmuch as it does not possess several of the important features of these three tribes. That Pseudomeloe appropriately belongs to the Eupomphini seems certain, and its inclusion in this tribe thus extends the latter’s distribu- tion to South America was well. Literature Cited Beauregard, H. 1890. Les insectes vesicants. F. Alcan, Paris. Blackwelder, R. E. 1945. Checklist of the Coleopterous insects of Mexico, Central America, the West Indies, and South America. U. S. Nat. Mus. Bull. No. 185, pp. 481-488. Borchmann, F. 1917. Meloidae, Cephaloidae. In Coleopterorum Catalogus. 69: 1-208, W. Junk, Berlin. Denier, P. C. L. 1935. Coleopterorum americanorum femiliae Meloidarum enumeratio synonymica. Rev. Soc. Entomol. Argentina 7: 139-176. Gupta, A. P. 1965. The digestive and reproductive systems of the Meloidae (Coleoptera) and their significance in the classification of the family. Ann. Entomol. Soc. Amer. 58(4) : 442-474. Van Dyke, E. C. 1928. A reclassification of the genera of North American Meloidae (Coleoptera). Univ. Calif. Pub. Entomol. 4: 395-474. Received for publication April 10, 1967 BOOK REVIEW Insect Behaviour. Symposium No. 3, Royal Entomological Society (P. T. Haskell, ed.). Bartholomew Press, Dorking, 1966. 113 p., £2.50. In this book are published the papers presented at the Third Symposium of the Royal Entomological Society, held September 23-24, 1965 in London. The papers are: (1) Orientation behaviour in insects and factors which influence it, by G. Birukow ; (2) The role of rhythms in insect behaviour, by P. S. Corbet; (3) Flight behaviour, by P. T. Haskell; (4) Feeding behaviour, by V. G. Dethier; (5) Sexual behaviour, by A. Manning; (6) Insect communication, by J. D. Carthv ; (7) Behaviour of social insects, by E. O. Wilson; (8) Some outstanding questions in insect behaviour, by J. S. Kennedy. The discussion that took place at the symposium is published at the end of each paper. These relatively brief, illustrated papers review much of the pertinent literature appear- ing for the most part since 1955. They are of somewhat uneven quality, some papers being better organized and better written than others. Some papers deal with their sub- jects only on a relatively broad, elementary level but others present data and interpretations not as well summarized elsewhere. The final paper, by J. S. Kennedy, is especially valuable to the general reader because, in a few pages, it discusses in an interesting way the salient problems in insect behavior. A thought-stimulating discussion follows this paper. This hardcover book is aesthetically printed, with few typographical errors. It is recommended for all persons interested in animal physiology, behavior, and ecology. Suzanne W. T. Batra Department of Entomology The University of Kansas, Lawrence 100 Proceedings of the New York Entomological Society (Meetings held in Room 129 of the American Museum of Natural History unless otherwise indicated.) Meeting of October 4, 1966 President Richard Fredrickson presided; 19 members and 3 guests were present. Dr. Fredrickson reported on the status of the proposed merger with the Brooklyn Society. At the special meeting, held on June 14th, 12 members were present and 68 affirmative proxies had been received; thus, our Society has approved the merger. He was authorized to proceed with the negotiations. The following were proposed for student membership: Richard Arnold of Hinsdale, Illinois, and Mrs. Winifred B. Trakimas, Francis C. Ford, and Dominick J. Pirone, three graduate students at Fordham University. program. Summer Activities of Members. Richard Fredrickson described a short field trip he had made to Blue Ridge, Va. Lucy Clausen spoke of the great increase in earwigs in the Bronx. This was corroborated by Jacob Huberman and Edwin W. Teale. Dominick Pirone reared some 2000 walking stick insects, and reported that from a 3 inch walking stick a 13 inch gordian worm emerged. He also drew attention to the Britten Sanctuary near Croton, N.Y. which has 127 acres available for collecting. David Kander described the ravages of cherry tree borers, and Ann Birdsey of a web worm invasion in Brooklyn. Aaron Nadler, a lawyer by profession and an active amateur entomologist, told of collecting psocids and curating his own collection at the Museum. Edwin Teale made some brief remarks about his 11,000 mile trip through England, Betty White about her trip to the Grand Teton Mountains, and David Miller about his trip to Jamaica, W.I. Patricia Vaurie commented on the effects of the severe drought around Easton, Pa. Excellent slides of a variety of insects were shown by Albert Poelzl and on the emergence of a dragon fly by Robert Buckbee. Mr. and Mrs. Sidney Hessel and Mr. and Mrs. Bernard Heineman attended the meeting of the Lepidopterists’ Society in Ottawa, Canada in early June. Lucy Heineman, Sec. Meeting of October 18, 1966 The meeting was called to order by President Fredrickson in Room 319; 24 members and 30 guests were present. The four student members proposed at the last meeting, Mrs. Winifred B. Trakimas and Messrs. Richard Arnold, Francis C. Ford, and Dominick J. Pirone, were elected to membership. Dr. L. L. Pechuman of the Department of Entomology, Cornell University, Ithaca, N. Y. was proposed for life membership and Sergio Orminati of City College was proposed for student membership. program. Army Ants — A Study in Social Behavior. Dr. T. C. Schneirla of the Depart- ment of Animal Behavior, American Museum of Natural History gave a brief resume of his studies of the army ants made at the laboratory on Barro Colorado Island in the Panama Canal over the past number of years. He told of the arrangements to have the activities of these ants filmed by the Encyclopedia Britannica Films during March 1966. This film, which has the same title as our program, is in color and has a running time of 20 minutes. It is equipped with a sound track with an explanatory narrative and such appropriate forest background sounds as those of ant birds. The main subjects of the film are the bivouacs or temporary nests, the mass raids, and the emigrations of the swarm-raider, Eciton burchelli, with supplementary scenes of emigrating and responding 101 102 New York Entomological Society [Vol. LXXV to the queen involving the related species, E. harnatum. Reactions of the workers to chemical trails and to the odor of their queen are shown both in field behavior and in terms of simple laboratory and field tests. Mr. John Walker of E. B. F., the photographer answered many questions about the technical problems in the forest filming. Lucy Heineman, Sec. Meeting of November 1, 1966 The meeting was called to order in Room 319 by President Fredrickson; 28 members and 8 guests were present. Dr. L. L. Pechuman was elected a life member and Mr. Sergio Orminati a student member in the Society. Miss Alice Gray informed the group that the Junior Society now has fourteen members and there are two applicants for the 1 5th and final place. The Junior Society had a successful summer which included field trips and a spelonking trip on which the expedition captured some cave insects. program. The Insects of the Galapagos Islands. Dr. Robert L. Usinger, President of the Entomological Society of America illustrated his talk with a map of the islands and slides. (An abstract follows.) Lucy Heineman, Sec. THE INSECTS OF THE GALAPAGOS ISLANDS The Galapagos International Scientific Project was organized by the Extension Division of the University of California at Berkeley under a grant from the National Science Founda- tion. Transportation was arranged through the LInited States Maritime Commission, using their ship, the “Golden Bear.” Financial assistance was provided by the National Science Foundation and also by the Belvedere Foundation of San Francisco. Logistical support was supplied by the United States Navy, Army and Air Force. The Associates in Tropical Biogeography of the University of California provided funds for some personnel and equipment. The Shell Oil Company provided funds for extra fuel for the “Golden Bear.” The expedition was in the field for about two months — January, February and March of 1964. Sixty scientists participated and another seventy or eighty persons visited the islands soon after we arrived for the dedication of the Darwin Memorial Research Laboratory. Entomological work spanned all of the life zones, from the strand through the lowland cactus forests which are very arid up to the moist middle elevations and then to the Miconia forest of the highlands and finally to the grass and fern zone at the top. This whole span of zones was represented back of the laboratory on a trail that was used intensively by the expedition. Other trips were taken to the other islands in the archipelago, either by ship or by helicopter. The composition of the insect fauna is characteristic of oceanic islands and in marked contrast to a continental archipelago such as the British Isles. The British Isles have about three times as many families, ten times as many genera and thirty-two times as many species as the Galapagos, and it is significant that endemism in the Galapagos, although high, consists mostly of single species in each genus. In contrast to this, the much older Hawaiian fauna commonly has many species in each of the endemic genera showing adaptive radiation and subspeciation on the various islands. In the Galapagos there is only in- cipient subspeciation, a few groups such as the grasshoppers showing size and color differences in the populations on each of the different islands. By comparison, other more rapidly evolving groups such as the cacti and composite plants ( Scalesia ) and the iguanas, tortoises and birds show clear-cut differences at the subspecies and even at the species level between the various islands. June, 1967] Proceedings 103 A few of the special characteristics of Galapagos insects are the concealing coloration of many of the Cerambycids and moths that rest on the lichen-covered rocks and tree trunks. Lichens are a characteristic feature of the Galapagos landscape. Also there is a great scarcity of aquatic insects because standing or running water is extremely rare. Only the dragonflies have flourished with endemic as well as introduced species. Other interesting aquatics include a few water beetles and an endemic mosquito in fresh water in the epiphytic plants in the forest, related to the lowland salt marsh mosquito, and various insects associated with the salt water lagoons. Pollination was a subject of special interest to the entomologists on the expedition. Only one bee, the Darwin carpenter bee, is found in the islands and evidence was obtained by screening flowers of many native and introduced plants to indicate that the old endemic Galapagos plants are mostly self-pollinated. It is only the introduced and more recent plants that seem to require insect pollination and this coincides with the idea that the carpenter bee was introduced after some of the early plants. Of course, sphinx moths and some other insects and some of the birds no doubt play a role in pollination as well. Interestingly enough, the carpenter bee brought with it its Meloid parasite, Cissites, and this, too, has evolved into an endemic species. In general, and despite Darwin’s observations made in September and October of 1835 when the dry season made a veritable desert of the islands, we found the insect fauna to be relatively rich. February is the height of the rainy season and many of the islands were green. Light collecting was especially productive with moths and Cerambycid beetles comparing in numbers, though not in species, to light trap catches in mainland areas. Darwin said that “Excepting Tierra del Fuego, I never saw so poor a country” and G. R. Waterhouse, upon examining the insects which Darwin collected, reported that there was nothing in their appearance which would have led him to imagine that they had come from under the Equator. Beebe reported that his field work was the most arduous and uncomfortable of any that he had experienced and Melville described the islands as vast cinder heaps. Fortunately, due to the favorable season, we encountered none of these difficulties and had a very productive entomological experience on the islands. Robert L. Usinger Meeting of November 15, 1966 Dr. Richard Fredrickson presided; 32 members and 9 guests were present. Mr. Orville Steward of the Bayard Cutting Arboretum, Oakdale, Long Island was proposed for regular membership and his son, Roger Steward, for student membership. Mrs. John Buck, the wife of the speaker of the evening, was introduced. She has assisted her husband in much of his scientific work, and she has accompanied him on expeditions. program. Synchronous Flashing of Fireflies. Dr. John B. Buck, Chief of the Laboratory of Physical Biology of the National Institutes of Health, Bethesda, Maryland illustrated his talk with interesting slides and diagrams. Although his talk referred largely to fireflies in the Orient, he drew many interesting comparisons between the oriental fireflies and those of the United States. (An abstract follows.) Lucy Heineman, Sec. SYNCHRONOUS FLASHING OF FIREFLIES Many observers have described long-lasting synchronous rhythmic flashing by huge swarms of fireflies in riverbank trees in the tropical Orient, but neither mechanism nor meaning have been explained. From observations of such trees in Sarawak and Thailand 104 New York Entomological Society [Vol. LXXV in October, 1965*, photometric and cinematographic recordings of the flashing of indi- viduals and populations, and study of captive specimens in the darkroom we established that: (1) The tree fireflies all belong to undescribed species of Pteroptyx. (2) Synchrony in Sarawak was disturbed by the concurrent presence of three species: In Thailand there were two but one was in great excess, permitting impressive displays of synchrony. Only males participate. (3) The period of the rhythm of flashing is about 560 msec, 95% of the cycles falling within ± 5 msec of this figure. (4) Analysis of cinematographic records of mass flashing indicates that the synchronizing individuals flash within less than ± 16 msec of each other. (5) In the buildup of synchrony in darkroom populations the co- ordination between two individuals was shown to depend on visual feedback, to operate over a range of less than 6 feet and to involve a progressive approach of the individual flash times until coincidence occurred, after which the two rhythms were locked together. (6) Since the coincidence is far closer than the minimal eye-lantern “reaction time” the synchrony must depend on a regulating mechanism controlled by the results of the preceding mass flash, rather than a direct individual-to-individual response. (7) The firefly trees represent quasi-permanent congregations in which fireflies remain in the tree by day, and are joined nightly by recruits from the surrounding swampland. (8) The tree congregations are viewed as a mass-mating substitute for the pair courtships which are usual in roving- type fireflies. Such congregations are made necessary by the impossibility of line-of-sight signaling in the impenetrable Nypa-mangrove vegetation. Presumably the mated females disperse back over the land for egg-laying. The synchronous flashing enhances the effec- tiveness of the trees as mating beacons. John B. Buck Meeting of December 6, 1966 Dr. Richard Fredrickson presided; 25 members were present as were 13 guests. The president appointed two committees: the auditing Committee consisting of Messrs. Albert Poelzl, Kumar Krishna, and A. B. Klots; and the Nominating Committee consisting of Messrs. David Miller, Robert L. Buckbee, and Bernard Heineman. Mr. Orville Steward and Roger Steward were elected regular and student members respectively in the Society. Dr. A. B. Klots told of an article by Miss Miriam Rothschild, the British entomologist, that will appear in an early issue of Natural History. Miss Rothschild has been working with larvae of the Monarch butterflies. They give off a volatile substance which, if sniffed a great deal, puts one in the conscious state of feeling you have already experienced this situation before and therefore you appear to be predicting the future, perhaps some- what like L. S. D. program. Sensory Codes and Feeding Behavior. Dr. Vincent Dethier, Professor of Zoology at the University of Pennsylvania, described work done largely in Holland using the tobacco horn-worm as the experimental animal. Excellent slides and diagrams were used to illustrate the talk. (An abstract follows.) Lucy Heineman, Sec. SENSORY CODES AND FEEDING BEHAVIOR The principal chemoreceptors of lepidopterous larvae are located on the maxillae and antennae. The maxilla bears, in addition to numerous mechanoreceptors and some olfactory * The American Philosophical Society and the National Geographic Society provided travel grants for this investigation. June, 1967] Proceedings 105 receptors, two sensilla styloconica that are organs of taste. Each sensillum styloconicum contains five bipolar neurons. The dendrites of four of these have been traced to the tip of the sensillum where they are exposed to the air. There are indications that they may subdivide and branch apically. The sensilla styloconica respond to a wide variety of solutions, but the responses of the two are not identical. Differences can be expressed in number of cells firing, fre- quency of impulses per cell, and/or total frequency of all impulses per sensillum. In Pro- toparce sexta the medial sensillum contains cells sensitive as follows: one to water and salt, one to sucrose and glucose, one to acid. The lateral sensillum contains cells sensitive as follows: one to water, one to salt, one to glucose, and possibly one to inositol. In Gal- leria mellonella the medial sensillum contains a cell sensitive to water, one to salt, and one to sucrose; the lateral sensillum has a water cell and a salt cell. In Philosamia cynthia the medial sensillum has two cells sensitive to salt and one to glucose; the lateral sensillum has one cell sensitive to water, one to salt, one to sucrose, and one to glucose. In Protoparce the sap of plants fires a number of cells in each sensillum. Sap of accept- able food plants appears to cause a higher frequency of firing in the cells of the medial sensillum than in those of the lateral sensillum while the sap of unacceptable plants, in general, causes a higher frequency of firing in the lateral sensillum. Some plants are ex- ceptions to this rule. There is evidence from these findings that both “feeding stimulants” and “deterrents” play a role in food-plant discrimination. The detailed information that the caterpillar receives from its maxillary gustatory receptors allows for participation by nutrients as well as token stimuli. The third segment of the maxillary palpus bears olfactory receptors. In Hyalophora gloveri complex responses were obtained to odors of wild cherry, potato, tomato, parsley, cabbage, privet, and willow. Responses to benzaldehyde and salicylaldehyde showed a long and pronounced after effect. Geraniol stimulated some cells while citronellal did not. The three large sensilla basiconica on the antennae of caterpillars are olfactory organs. One contains four bipolar neurons, one has five, and the other has seven. The dendrites of these cells break up into fine arborizations upon entering the cuticular peg and are in direct communication with the outside via a multitude of minute pores. Records obtained with micro metal electrodes reveal a background activity in these cells. This activity is depressed by air and may be either depressed or enhanced by odors. Each cell responds to more than one odor but not in the same manner. Furthermore, not all cells exhibit identical response patterns although there is some overlap. For the caterpillar plant odors are obviously coded as complex patterns. Food-plant discrimination cannot be explained solely in terms of acceptance or rejection via the maxillary taste receptors but must also involve the wealth of olfactory information provided by the antennae and maxillae. Vincent Dethier Meeting of December 20, 1966 President Fredrickson presided; 22 members and 13 guests were present. The president announced that on the following afternoon there would be a meeting at the office of the Society’s attorney at which time he, the secretary or assistant secretary, and the president and secretary of the Brooklyn Society would sign the agreement merging the two societies. The merger will have to be reviewed by the courts. Mr. Anthony J. W. Owston was pro- posed as a regular member and Mr. Michael Boshes of City College as a student member. Dr. Schmidt asked if anyone could advise him as to where he could get information about 106 New York Entomological Society [Vol. LXXV a flea trap. Dr. Klots said that there was a picture of a medieval one in the article by Miss Miriam Rothschild in a recent issue of The Scientific American. Dr. Edwin W. Teale remarked that in the recent warm spell there were myriads of snow fleas at his home in Connecticut. program. Naturalists in South America. Mr. Heineman showed colored slides of a recent trip to South America, and Mrs. Heineman described and commented on them. Lucy Heineman, Sec. Meeting of January 3, 1967 — The Annual Meeting The meeting was called to order by Dr. David Miller in place of President Fredrickson who was ill. The Vice-president was not able to be present because of a class. Dr. Miller asked for nominations to elect a chairman for the evening; he was duly elected to conduct the meeting for the evening. Nineteen members and seven guests were present. Mr. Raymond Brush, the Treasurer, reported a favorable balance for the fiscal year, 1966. In the absence of the Editor, the Associate Editor, Dr. Forbes, announced that the De- cember 1966 issue of the Journal is expected very soon. He said more manuscripts would be welcomed. The Nominating Committee, consisting of Messrs. Miller, Heineman, and Buckbee, chairman, submitted the following slate for the coming year: President — Dr. Richard Fredrickson Vice-president — Dr. David Miller Treasurer — Mr. Raymond Brush Assistant Treasurer — Mrs. Patricia Vaurie Secretary — Assistant Secretary — Mr. Albert Poelzl Trustees (to serve two years) — Dr. Elsie Klots, Mr. Bernard Heineman Publication Com- mittee— Drs. Kumar Krishna, Asher Treat, Pedro Wygodzinsky. The chairman called for nominations for the office of Secretary from the floor. He stated that Mrs. Heineman has consented to continue for the month of January. No nominations were forthcoming and the slate was elected as presented. Dr. Miller continued to chair the meeting as the newly elected vice-president. Dr. Wygodzinsky announced that there are two vacancies in the Entomology Department of the Museum. One is for a scientific assistant and the other is for a technical artist. Both are for two years, and they are covered by grants. Dr. Forbes exhibited a copy of the December 9 issue of Medical World News. The cover is a picture of our member, Dr. Roman Vishniac, and the feature article is on Dr. Vishniac’s remarkable photography. Many magazines have carried articles on Dr. Vishniac’s photography, but this is the first time his photograph has appeared on a cover. Mr. Anthony J. S. Owston and Mr. Michael Boshes were elected regular and student members, respectively. Miss Ann Young of the City College of New York who has worked with Mr. Topoff, our speaker of the evening, at the Southwest Research Station was intro- duced. Mr. Nicholas Shoumatoff, a former president of the Society, was introduced. He has now returned to the United States from a period of employment in England. program. Behavioral and Physiological Studies in the Army Ant, ISeivamyrmex. Mr. Howard Topoff, a student at City University and the Department of Animal Be- havior of the Museum illustrated his interesting talk with charts and slides. (An abstract follows.) Lucy Heineman, Sec. June, 1967] Proceedings 107 BEHAVIORAL AND PHYSIOLOGICAL STUDIES IN THE ARMY ANT, NEIVAMYRMEX Social organization and behavior in the phyletic level that is characteristic of insects is influenced predominantly by the intensity of stimuli originating from reproductive, feeding, and reciprocal stimulative processes. In the army ant genus Neivamyrmex qualitative differences in the intensity of raiding during the nomadic and statary phases are also reflected quantitatively in the sharp in- crease in oxygen consumption at the onset of the nomadic phase, followed by a marked decrease as the statary phase is initiated. Thresholds of responses to a given intensity of light as well as to the changing olfactory stimuli which emanate from the brood, queen and workers, increase during the nomadic phase and decrease during the statary. The populational characteristics of three genera of doryline ants ( Neivamyrmex , Eciton, and Aenictus) were compared with a discussion of the trophic factors that influence caste determination in the army ants in particular, and in the social insects in general. Howard R. Topoff Meeting of January 17, 1967 President Richard Fredrickson presided; 22 members and 7 guests were present. Mr. Howard R. Topoff of the City University of New York was proposed and duly elected as Secretary of the Society, succeeding Mrs. Lucy Heineman. Dr. Jerry Vanderberg of the Department of Preventive Medicine, New York University Medical School was proposed for regular membership. Miss Betty White described a rare but delightful occasion she had comparing a photograph of a parasitic wasp from the book “Living Insects Of the World” with one that flew into her kitchen; they were identical. program. Dr. Robert Traub, Research Professor at the University of Maryland School of Medicine, presented two talks: Ecology of Scrub Typhus in Unusual Habitats in Paki- stan and Examples of Convergent Evolution in Fleas. In the first talk Dr. Traub dis- cussed the tremendous increase in interest in scrub typhus, especially in relation to suc- cessful military efforts in tropical habitats. He reported on his interesting and perplexing findings that this disease, which predominates in ecologically disturbed tropical environ- ments, has recently been found infecting the small mammal populations of primary forests, xerophytic forests, subalpine habitats in the Himalayas, and even in true alpine meadows as high as 11,000 feet. In the second talk Dr. Traub discussed the convergence of adapta- tions possessed by fleas, for attaching to their hosts. Particular mention was made of the fact that fleas associated with birds and arboreal mammals usually possess longer and more sharply pointed comb spines than fleas which parasitize ground-dwelling mammals. Howard R. Topoff, Sec. Meeting of February 7, 1967 was cancelled because of a heavy snowfall. Meeting of February 21, 1967 Dr. Richard Fredrickson presided; 19 members and 2 guests were present. Dr. Jerry Vanderberg was elected to regular membership. Dr. J. G. Butte of the State University of New York at Farmingdale was proposed for regular membership. Dr. Asher Treat 108 New York Entomological Society [Vol. LXXV called attention to the paper of Dr. Carol Williams, in the February 3 edition of Science, noting that the female of the polvphemus moth will not produce a sex attractant pheromone until she is stimulated bv an extract of oak leaves. program. Trap-Nesting Wasps and Bees and Their Associates. Dr. Karl Krombein, chairman, Department of Entomology of the Smithsonian Institute illustrated his talk with slides. (An abstract follows.) [ Editor’s note : This whole project is reported in detail in a book by Dr. Krombein, “Trap-Nesting Wasps and Bees: Life Histories, Nests, and Associates,” Smithsonian Press, 579 pp., 29 pis., 1967.1 TRAP-NESTING WASPS AND BEES AND THEIR ASSOCIATES The speakers discussed the field project he carried on from 1953 to 1964 investigating the biology of solitary wasps and bees which can be induced to nest in wooden traps. The traps were made from straight-grained pieces of white pine, each containing a boring 6" long and %, Vt or 1/9" in diameter. The traps were made into bundles containing one or two traps of each diameter. The bundles were placed in the field in situations where populations of solitary wasps and bees were nesting in abandoned borings of other insects in wood such as on dead branches and tree trunks, on sound oak branches bearing insect galls, and on structural lumber. Nests were obtained from trap settings in western New York, the area around Washington, D. C., coastal North Carolina, Archbold Biological Station in Florida, and the Southwestern Research Station in Arizona. The nests were opened in the laboratory to record the details of the nest architecture and to preserve samples of the food stored for the larvae; periodic reexamination of the nests provided information on the developmental stages of the wasps and bees, and their associated preda- tors, parasites and symbionts. Nine new species and subspecies of wasps and bees were described from these nests, as well as three new species of chalcid parasites, and two new genera and 17 new species of parasitic mites. Life history data were obtained for 75 pre- daceous wasps and 43 non-parasitic bees, and 83 associated parasites and predators (28 of them parasitic wasps or bees). Dr. Krombein illustrated his talk with a number of Kodachrome and black and white transparencies showing the nest architecture of a number of species, the life history of a typical vespid wasp, nesting behavior of the bee Osmia lignaria, certain aspects of the competition between three species of Trypargilum for nesting sites and spider prey, and examples of some of the mite, beetle, fly and wasp parasites associated with the host wasps and bees. Karl Krombein Meeting of March 7, 1967 Dr. Fredrickson presided; 14 members and 7 guests were present. Dr. J. G. Butte of the State University of New York at Farmingdale was elected a regular member, and Miss Ann Young, a graduate student at the City University of New York, was proposed for student membership. Miss Alice Gray of the Department of Entomology at the Museum displayed toy insects made in Hong Kong. program. Ecology of the Cave-Entrance Fauna. Professor Richard Graham of the Department of Physiology of Rutgers University discussed ecological zonation in caves with particular reference to the cave entrance as a persistent community. Howard R. Topoff, Sec. June, 1967] Proceedings 109 Meeting of March 21, 1967 President Richard Fredrickson called the meeting to order; 23 members and guests were present. Miss Ann Young of the City University of New York was elected to student membership. program. Mimicry in Butterflies. Dr. Michael G. Emsley, Assistant Curator of Insects of the Philadelphia Academy of Natural Sciences was the speaker of the evening. (An ab- stract follows.) Howard R. Topoff, Sec. MIMICRY IN BUTTERFLIES “At the close of the last century a confusingly large number of named forms of Heli- conious erato and Heliconious melpomene were described, many of them as descrete species. We now know that these two species show pronounce! geographic variation with mono- morphic forms occupying Central America, South America west of the Andes, northern South America, the valley systems of the eastern Andes, the Amazon Basin and south- eastern Brazil. Where the monomorphic populations meet there is a high degree of poly- morphism which has led to the large number of described forms. The most remarkable feature of this situation is that erato and melpomene vary so greatly over their range they maintain a mutually similar appearance everywhere they occur. The closeness of their similarity makes a convincing case that mimicry in butterflies is a real phenomenon. Unfortunately, though Dr. Brower and his co-workers have tried extremely hard to obtain convincing, experimental proof of the values of what we call warning coloration and its imitation by palatable mimics, the evidence is still far from complete. Any theory con- cerned with the explanation of the color pattern of butterflies in relation to their predators must also take into account that color is probably the prime factor in species recognition and in the releasing of courtship behavior.” Michael G. Emsley NEW MEMBERS The following persons have been elected to the Society since the membership list was published in the June 1966 issue (vol. 74, pp. 112-115). The class of membership other than regular member is designated by the letter in parentheses: L — Life, St. — Student. (St) Arnold, Richard, 735 McKinley Lane, Hinsdale, Illinois 60521 Bartolone, Pat J., 1661 East 172nd Street, Bronx, N.Y. 10472 Benton, Allen, State LTniversity College, Fredonia, N.Y. 14063 (St) Boshes, Michael, Department of Animal Behavior, American Museum of Natural History, 77th Street and Central Park West, New York, N.Y. 10024 Butte, J. G., State University of New York, Farmingdale, N.Y. 11735 Durden, Beatrice V., Carnegie Museum, Pittsburgh, Penna. 15213 Emsley, Michael G., Academy of Natural Sciences, Philadelphia, Penna. 19103 (St) Ford, Francis C., 650 Yonkers Avenue, Yonkers, N.Y. 10704 (St) Friedman, Kenneth, 33-05 90th Street, Jackson Heights, N.Y. 11372 (St) Kanter, David F., 154-04 25th Avenue, Flushing, N.Y. 11354 (St) Mesibov, Robert, 1905 Birge Terrace, Madison, Wisconsin 53705 Nadler, Aaron M., 101 Ocean Parkway, Brooklyn, N.Y. 11218 (St) Novak, John A., Kent State University, Kent, Ohio 44240 (St) Orminati, Sergio, 200 Eighth Avenue, New York, N.Y. 10011 Owston, Anthony J. W., 345 East 56th Street, New York, N.Y. 10022 (L) Pechuman, L. L., Department of Entomology, Cornel University, Ithaca, N.Y. 14850 (St) Pirone, Dominick J., 120 Esplanade, Mt. Vernon, N.Y. 10553 Pogany, Margaret, Simon & Schuster, Inc., 630 Fifth Avenue, New York, N.Y. 10020 Ruckes, Herbert, Jr., Biology Department, Manhattan Community College, New York, N.Y. Spear, Philip, National Pest Control Association, 250 West Jersey Street, Elizabeth, N.J. 07202 Steward, Orville, c/o Bayard Cutting Arboretum, P.O. Box 66, Oakdale, N.Y. 11769 (St) Steward, Roger, c/o Bayard Cutting Arboretum, P.O. Box 66, Oakdale, N.Y. 11769 Stibick, J. N. L., Department of Entomology, Purdue University, Lafayette, Indiana 47907 Stien, Harry, 12 Highland Drive, Ardsley, N.Y. (St) Topoff, Howard R., Department of Animal Behavior, American Museum of Natural History, 77th Street and Central Park West, New York, N.Y. 10024 (St) Trakimas, Winifred B., 137 Stratford Street, Roslyn Heights, N.Y. 11577 Vanderberg, Jerry, 333 East 14th Street, New York, N.Y. 10003 Watson, Kennith, 53 Kenefick Avenue, Buffalo, N.Y. 14220 (St) Young, Ann, 512 East 79th Street, New York, N.Y. 10021 110 INVITATION TO MEMBERSHIP The New York Entomological Society was founded in 1892 and incorporated the following year. It holds a distinguished position among scientific and cultural organizations. The Society’s Journal is one of the oldest of the leading entomological periodicals in the United States. Members and subscribers are drawn from all parts of the world, and they include distinguished professional naturalists, enthusiastic amateurs, and laymen for whom insects are only one among many interests. You are cordially invited to apply for membership in the Society or to subscribe to its Journal which is published quarterly. Regular meetings are held at 8:00 P.M. on the first and third Tuesdays of each month from October through May at the American Museum of Natural History, the headquarters of the Society. A subject of general interest is discussed at each meeting by an invited speaker. No special training in biology or entomology is necessary for the enjoyment of these talks, most of which are illustrated. Candidates for membership are proposed at a regular meeting and are voted upon at the following meeting. CLASSES OF MEMBERSHIP AND YEARLY DUES Active member: Full membership in the Society, entitled to vote and hold office; with Journal subscription $9.00 Active member without Journal subscription 4.00 Sustaining member: Active member who voluntarily elects to pay $25.00 per year in lieu of regular annual dues. Lije member: Active member who has attained age 45 and who pays the sum of $100.00 in lieu of further annual dues. Student member: Person interested in entomology who is still attending school; with Journal subscription 5.00 (Student members are not entitled to vote or to hold office.) Student member without Journal subscription 2.00 Subscription to Journal without membership 8.00 APPLICATION FOR MEMBERSHIP Date I wish to apply for membership (see classes above). My entomological interests are: If this is a student membership, please indicate school attending and present level. Name Address (Zip Code must be included) Send application to Secretary — iO(o / O ) Devoted to Entomology in General ii>v f i mi\w} QVViv;V/,,i K'A' i '\ ’r \ V v. , t: i \ VV; AC py ' i ) 1 1, yyp\y C:'kyA yA M A! / \ \ \- ' 1 , n " y f ■ : '<■ ' a'- j ■■ ‘li ’/ ■ ,~'S\ ■CP > A i U\ r, / ; Am, , ; fl* V\) L , j. tl V\ 1 i 1 ./ X t Vi ( A ,+ \v\r The 7/ V > Ai-hr :y ,7 v • - ' \ . r'7~ f ( v , New York Entomological Society h\ ■ 7. 'jfcv p •'f ! (\j 1 y (r.( TO vM 'V m v Organized June 29, 1892 — Incorporated February 25, 1893 \ I / Reincorporated February 17, 1943 s 1 i V )r -> 1 I . Ayhk t pc A i w W A 'PP I; The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St., & Central Park W., New York 24, N. Y. „/■ Q [\ A Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. Members of the Society will please remit their annual dues, payable in January, to the Treasurer. , 4 / : , , \ 1 ' P* '< J/.vY' 4 !v7 A '■P \ i -KP ' C mp < P Ai IP u J I.',! Officers for the Year 1967 ' President, Dr. Richard Fredrickson 4 ; PC. Vice-President, Dr. David Miller College of the City of New York 10031 ' ■ ■ i 1 | i/\ 'A' \-hx 'A" ' A r~ A ) • vV/ Vl ' • \ fkhl •• 1 .’I- ' v r:\AP VI College of the City of New York 10031 f \ r , . I , \ j t — ' j ' I • ; ■ 1 *, » ' I *■ ^ i "\ T 1 i IT . ■ , ”1 v Secretary, Mr. Howard Topoff American Museum of Natural History, New York 10024 7 At, A . : \ * " K.-'v, Assistant Secretary, Mr. Albert Poelzl 7# 77, 230 E. 78th Street, New York 10021 Treasurer, Mr. Raymond Brush ,v ; M A A American Museum of Natural History, New York 10024 ■M A , r Assistant Treasurer, Mrs. Patricia Vaurie American Museum of Natural History, New York 10024 I ■ - v > . ■ A\ v V '■ " VbVv \ V; m A\\' _ f ii pr — t , 4i , 1 1 U-lV ’ .,VvV : 1 7 1 711; ? m a. C \ ; p ^ Class of 1967 Dr. Jerome Rozen, Jr. Class) of 1968 •••. Pi - \ ■■ Wf 7 .1 '' A P-S.P 'h 1A m 'iWM ? X'r-P. :Ty ,.yy 4 4y, Tp 1 }'!y% , 4 V1 -PA. Jp-. P ' : < ■ i •//- i -4 , V - AAA |. Vn. • , ' j , 1 , ,A ,7 ' ' ’ Dr. Elsie Klots /, i i' ( ) -4)1-':, cV M Mr. Robert Buckbee Mr. Bernard Heineman ''-N P Mailed October 6, 1967 CP ’r ) ; \. V ‘ T* 1 ' ' V' V 4 C : \A\ 1 1 f 1 lie A $)A: A' '7 ' 1 The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press Inc., 104l Ne|w Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas. I ) (T p* I. p- Oi , v'V s\i 7 v \ \ r it\ xwypt ,v; 'ff i7' f j\ T?\ i i ■■ m 4^7 ■ 7> , a; , , 1 v> P Journal of the New York Entomological Society Volume LXXV October 6, 1967 No. 3 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen College of Pharmaceutical Sciences, Columbia University 115 West 68th Street, N. Y. 10023 Associate Editor James Forbes Fordham University, N. Y. 10458 Publication Committee Dr. Kumar Krishna Dr. Asher Treat Dr. Pedro Wygodzinsky CONTENTS A New Liphistiid Spider from China (Aranae: Liphistiidae) Willis J. Gertsch 114 Activities of Respiratory Enzymes During the Metamorphosis of the Face Fly, Musca autumnalis (De Geer) P. G. Rousell 119 Some Synonyms in American Spiders Wilton Ivie 126 Biology of Dufourea and of its Cleptoparasite, Neopasites (Hymenoptera : Apoidea) Philip F. Torchio, Jerome G. Rozen, Jr., George E. Bohart, and Marjorie S. Favreau 132 Behavior of the German Cockroach, Blattella germanica (L.)? in Response to Surface Textures Robert Berthold, Jr. 148 Two New Species of Crambus (Fabricius) from Western North America (Lep- idoptera: Pyralidae) Alexander B. Klots 154 Perobyscopsylla hamifer (Rothschild) : An Addition to the Entomological Fauna of New York State Allen H. Benton 159 New and Little Known Species of Serica (Coleoptera: Scarahaeidae) X R. W. Dawson 161 Observations of Epicordulia princeps (Hagen) (Odonata: Corduliidae) at a Light Allen M. Young 179 Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera: Tipulidae), XV Charles P. Alexander 183 Book Review 147 A New Liphistiid Spider from China ( Araneae : Liphistiidae ) Willis J. Gertsch The American Museum of Natural History, New York, N.Y. Abstract: A new species of liphistiid spider, Heptathela bristowei, is described on the basis of a female from Szechuan, China. In a discussion the author concludes that the family Heptathelidae cannot be maintained and that the species with a posterior colulus (. Heptathela ) be given only generic ranking. Family Liphistiidae This small family comprising our most generalized spiders was reviewed by Bristowe (1932), who gave comparative data on the then known seven species of Liphistius and two species of Heptathela. In 1939 Petrunkevitch raised the latter genus to full family status on the basis of characters found in the internal anatomy of a female of Heptathela sinensis Bishop and Crosby (1933). The new family Heptathelidae was relegated to synonymy by Gertsch (1949, p. 265) but was recognized by Vachon (1958, p. 431), who contributed important new information on the postembryonic development of Heptathela kimurai Kishida. The family Heptathelidae was based on the following principal features: reduction of the posterior median spinnerets to a functionless vestige, a posterior colulus; reduction of the number of ostia in the heart from five to four pairs; loss of the endocheliceral venom glands. In Liphistius all eight spinnerets are still retained, the heart has five pairs of ostia, and the venom glands, although reduced in size, are still present. Such regressive changes as those credited to Heptathela may have great systematic importance or almost none at all. It should be mentioned that these internal differences are based on knowledge of only half a dozen specimens of at most three or four species. Except for the loss of the posterior median spinnerets, the genus Heptathela shows such close correspondence to Liphistius that it seems undesirable to accord it more than generic distinction. An even more conservative position was taken by Schenkel (1953, p. 1) when he described a species, that should now be listed as Liphistius schensiensis Schenkel, under the following trinomial: Liphistius ( Heptathela ) sinensis (Bishop and Crosby), var. schensiensis , n. var. Since his specimen had eight spinnerets, instead of the seven credited to sinensis, he concluded that this feature was not constant. Further, he saw no need to give even subgeneric recognition to Heptathela (misspelled Heptathele) . Whereas it must be con- ceded that the two genera are remarkably alike, it seems desirable to continue to hold them separate on the basis of the differences in the posterior median 114 September, 1967] Gertsch: New Lipiiistiid Spider Figs. 1-3. Heptathela bristowei, n. sp., female. 1. Carapace and abdomen, dorsal view. 2. Abdomen, ventral view. 3. Epigynum, dorsal view. spinnerets. Thus, Liphistius schensiensis Schenkel is the eighth species of its genus and the species described below is the third for Heptathela. No mention of the internal seminal receptacles of any female liphistiid was made by Bristowe (1932) or any of the principal students who considered the systematics and morphology of the group. This organ (for which I use 116 New York Entomological Society [Vol. LXXV the term epigynum in its broadest connotation) is of the “haplogyne” type. In the Atypoidea (Gertsch, 1949, p. 126, 1128, etc.), there are four primary seminal receptacles. The epigynum of Liphistius malayanus Abraham was illustrated by Schiapelli and Gerschman (1962, pi. 2, figs. 5-6) and shows the four, rather small receptacles, flanking a central pouch, as well as a central cluster of globular organs. The epigynum of Heptathela bristowei is of the same general type and is similar to that of kimurai , the type of the genus. Whereas most of the typical tarantulas (Ctenizoidea) have epigyna with a single seminal receptacle on each side, a few exceptions have been illustrated by Schiapelli and Gerschman (1962, pi. 4, figs. 1-3). Heptathela bristowei, n. sp. Figures 1-3 This interesting species is dedicated to Mr. W. S. Bristowe, colleague and eminent author of “The World of Spiders,” and one who has contributed much to knowledge of the biology and taxonomy of the liphistiid spiders. diagnosis: This species resembles Heptathela sinensis Bishop and Crosby, from Tsinan, Shantung, China, but is readily separated by the following features: The pars cephalica is proportionately narrower in front and its greatest width is only four-fifths the distance to the cervical groove, instead of having these ratios equal. The four median eyes, encircled by the narrowly oval lateral eyes, are closer together. The cervical groove is considerably larger and deeper. The fourth femora are provided below on the retrolateral margin with a double row of short spinules, instead of eight spines. The tergal plates on the abdomen are smaller in size and the first lung plate is of different form, as shown in the figures. female holotype: Total length, including chelicerae, 19.5 mm. Carapace Sternum Labium Maxilla Abdomen Length 7.0 3.5 1.0 3.0 10.0 mm. Width 5.7 2.4 2.0 1.5 8.0 mm. Carapace orange to reddish brown ; pars cephalica dusky and pars thoracica with dusky streaks radiating from median groove ; eye tubercle black. Chelicerae dull reddish brown, pale at base above. Sternum, labium and appendages quite uniform dull orange brown. Abdomen gray; tergites dusky brown. Dorsal view of carapace and abdomen as shown in fig. 1. Structure typical, essentially like that of sinensis. Carapace quite smooth, bare except for tiny setae lying flat on pars cephalica, a middle line of about six stout setae running through and behind median eyes, a series of four setae on clypeal margin, with median pair much longer, and a line of small setae margining carapace. Carapace broadly rounded in front, sharply angled at corners, gently rounded on sides and truncated behind. Pars cephalica strongly elevated, highest just behind eyes; cervical groove deep rounded de- pression smaller than eye turret, situated back five-eighths of length; pars thoracica low, convex, with transverse grooves. Eyes all close together, set on rounded tubercle of typical height. Clypeus inclined forward, narrow, equal to about radius of posterior median eye. Ratio of eyes: ALE : AME : PLE : PME = 62 : 6 : 48 : 35. Front eye row slightly procurved; lateral eyes large, narrowly oval, nearly touching in front ; median eyes minute, lying in front of posterior September, 1967] Gertsch: New Liphistiid Spider 117 median eyes. Posterior eye row moderately recurved; oval median eyes close together, separated by one-fourth their narrow diameter, about as far at narrowest point from larger oval lateral eyes. Median ocular quadrangle broader than long, narrowed in front, with anterior eyes minute. Sternum an elevated sclerite with steep sides, covered with coarse setae, without trace of sigilla. Labium free, separated from sternum by deep, transverse groove, set with black setae. Maxilla truncated at apex, with setae over most of surface and brush of soft hairs along inside margin. Chelicera about 3 mm. long as seen from above, smooth at base, expanded toward apex and set with coarse setae, rounded at apex above claw and without rake; fang of median length, rather stout, lying in indistinct groove margined on prolateral side by row of eight, close-set, black teeth, three of these larger, and on retro- lateral side with thin brush of soft reddish hairs. I II III IV Palpus Femur 4.7 4.1 4.3 6.1 4.3 mm. Patella 2.7 2.6 2.7 3.1 2.4 mm. Tibia 3.0 2.7 2.7 4.2 3.0 mm. Metatarsus 3.0 3.2 3.5 5.8 — Tarsus 1.7 2.0 2.1 2.7 3.7 mm. Total 15.1 14.6 15.3 21.9 13.4 mm. leg formula: 4312. All legs short, clothed sparsely above with hairs and weak spines and below and on sides with more numerous, stouter spines. First and second legs with rows of stout ventral spines on tibiae, metatarsi and tarsi, those on anterior segments nearly lateral in position. Femora with ventral hairs and weak spines; fourth femora with 20 or more stout spinules below in double row near retrolateral edge. Pedipalp with stout sub- lateral spines; tarsus set with even row of seven heavy spines on lateral margins; palpal claw with single tooth at base. Paired claws of legs with two teeth near base ; unpaired claws quite straight, unarmed below. abdomen (figs. 1-2): Globose, covered evenly with tiny setae. Ten tergites visible on dorsum with lateral measurements of these, in millimeters, from front to rear as follows: 3.5; 4.3; 4.2; 4.2; 3.3; 1.8; 1.1; 0.8; 0.7; 0.6; thus, sixth and succeeding tergal plates greatly reduced in size ; each tergal plate with pair of prominent alveoli on caudal edge, bearing long spines. First lung plate gradually produced behind to evenly rounded projection, without special angles or evident grooving. Spinnerets of average size; posterior colulus a small tubercle bearing three tiny setae. epigynum (fig. 3): Consisting of four receptacles; lateral receptacle of each pair larger than inner one. type data: Female holotype from Wanhsien, Yen-Ching-Kao, Szechuan, China, February, 1922 (W. Granger), in the American Museum of Natural History. Literature Cited Bishop, S. C. and Crosby, C. 1932. A New Species of the Spider Family Liphistiidae from China. Peking Nat. Hist. Bull., 6, pp. 5-7, 7 figures. Bristowe, U. S. 1932. The Liphistiid Spiders. With an appendix on their Internal Anatomy by J. Millot. Proc. Zool. Soc. London, pp. 1016-1057, pi. I-VI, text figs. 1-11. Gertsch, W. J. 1949. American Spiders. Van Nostrand, New York. Pp. v-xiii, 1-285, pi. 1-32, PI. I-XXXII. 118 New York Entomological Society [Vol. LXXV Petrunkevitch, A. 1939. Catalogue of American Spiders. Trans. Connecticut Acad. Arts. Sci., 33, pp. 133-338. Schiapelli, R. D., and Gerschman de Pikelin, B. S. 1962. Importancia de las Espermate- cas en la Sistematica de las Aranas del Suborden Mygalomorphae. Physis, 23, No. 64, pp. 69-75, 4 plates. Schenkel, E. 1953. Chinesische Arachnoidea aus dem Museum Hoangho-Peiho in Tienlsin. Bol. Mus. Nac., Rio de Janeiro, No. 119, pp. 1-108, 47 figs. Vachon, M. 1958. Contribution a 1’Etude de Developpement Post-embryonnaire des Araignees. Deuxieme note. Orthognathes. Bull. Soc. Zool. France, 83, pp. 429-461. Received for publication April 17, 1967 Activities of Respiratory Enzymes During the Metamorphosis of the Face Fly, Musca autumnalis De Geer1 P. G. Rousell St. Francis Xavier University, Antigonish, Nova Scotia, Canada Abstract: The activities of alcohol, succinic, malic, glucose, glutamic, alpha-glycerophos- phate, lactic, and isocitric dehydrogenases, the malic enzyme, and cytochrome oxidase were determined during the metamorphosis of the face fly, Musca autumnalis . Total alpha-glycerophosphate, alcohol, malic, and succinic dehydrogenases as well as the malic enzyme exhibited U-shaped activity. Greatest activity was shown by the malic de- hydrogenase. Isocitric dehydrogenase activity was high initially and remained high until the 2-day pupa, and thereafter showed a progressive decline. Glucose dehydrogenase activity was low and remained fairly steady during the entire pupal stage. Alcohol dehydrogenase decreased steadily during the first days of metamorphosis, reached a low value on the third day, and then increased to reach its highest value in the adult stage. Succinic dehydrogenase exhibited a similar pattern, but the level of activity was not as high as most of the other dehydrogenases. Glutamic dehydrogenase showed low activity in the larval stage. It decreased during the first several days of the pupal life and completely disappeared by the fourth day. The activity of lactic dehydrogenase was very low throughout metamorphosis. Malic enzyme exhibited high activity in the larva, prepupa, and again in the adult stage. Cytochrome oxidase activity was also U-shaped during metamorphosis. The 02 consumption of holometabolous insects follows a U-shaped curve dur- ing metamorphosis. This phenomenon was first described by Krogh (1914) for the mealworm, Tenebrio molitor, and subsequently has been confirmed by the following investigators employing a variety of insect species: Clare, 1925; Fink, 1925; Bodine and Orr, 1925; Ludwig, 1931; Dobzhansky and Poulson, 1935; Wolsky, 1938; Sacktor, 1951; Ito, 1954; Cotty, 1956; and Ludwig and Barsa, 1956. Since the causative factors responsible for the U-shaped respiratory curve are not fully understood, various explanations have been advanced. Krogh (1914) and Fink (1925) believed the changes in C)2 consumption to be associated with different degrees of tissue organization. The activity of cytochrome oxidase has been investigated as a rate-limiting factor in respiratory metabolism. Wolsky ( 1938), Williams ( 1950), Ludwig (1953) and Diamantis (1962) found U-shaped activity curves for cytochrome oxidase during the pupal stages of the fruit fly Drosophila melanogaster , the moth' Platysamia cecropia , the Japanese beetle Popillia japonic a, and the flour moth Ephestia kiihniella , respectively. A corre- lation between succinic dehydrogenase activity and respiratory metabolism has been described by Wolsky (1941) for Drosophila melanogaster , Ito (1954) for Bombyx mori , Ludwig and Barsa (1955) for Popillia japonica and for Tenebrio 1 This investigation was financed by a research grant of the National Research Council of Canada. 119 120 New York Entomological Society [Vol. LXXV molitor (1958). Agrell (1949) described total dehydrogenase activity and the activities of malic, citric, and glutamic dehydrogenases as U-shaped during the metamorphosis of the blow fly, Calliphora erythrocephala. Ludwig and Barsa (1958) found malic and succinic dehydrogenases and the malic enzyme activities to be U-shaped during the metamorphosis of Tenebrio molitor. In 1959, they found that with the house fly alcohol and alpha-glycerophosphate dehydro- genases also followed U-shaped curves. Diamantis (1962) described similar activity for alpha-glycerophosphate I and II, malic, isocitric and succinic de- hydrogenases and the malic enzyme. His report of the U-shaped activity of isocitric dehydrogenase is at variance with the findings of Ludwig and Barsa (1959) for the house fly. They reported the isocitric dehydrogenase showed a steady decrease during metamorphosis. Diamantis (1962) also found low glu- tamic dehydrogenase activity at all stages, whereas Ludwig and Barsa (1959) found that it disappeared early in the pupal stage. In the present investigation a study was made of cytochrome oxidase and the various dehydrogenases during the metamorphosis of the face fly Musca autum- nalis. MATERIALS AND METHODS The insects used in this study were obtained from the United States Depart- ment of Agriculture Research Center, Beltsville, Maryland. They were reared in screened cages measuring 30 X 30 X 30 inches. The temperature of the rear- ing room was 25 ± 2°C and the relative humidity varied between 35-60 per cent. The light source consisted of two 160-W General Electric F 40 CW fluo- rescent lamps that gave a light intensity of approximately 150 ft-c measured at the top of the cages. The optimum photoperiod was found to be 16 hours ex- tending from 6 a.m. to 10 p.m. A mixture of skimmed milk and 5 per cent sucrose solution in a 2:1 ratio was placed daily in a petri dish containing a centrally located piece of absorbent cotton which served as a resting place for the flies when they were feeding. Ap- proximately 10 ml of citrated bovine blood was placed in a second dish and 3 ml of 5 per cent maltose solution was also added to this receptacle. Fresh cow dung was placed in a third dish to serve both as a source of food and as an oviposition medium. Each day after being removed from the cages, the dishes of manure were set aside for 48 hours and then examined for the presence of larvae. If larvae were found, the manure was transferred to a porcelain tray (15 X 10 X 3 inches) containing a large central mass of dung surrounded by a fairly thick layer of vermiculite into which the larvae migrated just prior to pupation. These trays were covered with a layer of cheesecloth and placed on shelves in the rearing room. Following pupation, the insects were gently re- moved to a small dish which was put in one of the rearing cages to await emer- gence. September, 1967] Rousell: Respiratory Enzymes of Face Fly 121 The activities of alcohol, succinic, malic, glucose, glutamic, alpha-glycerophos- phate, lactic, and isocitric dehydrogenases and the malic enzyme were deter- mined by the Thunberg method as given by Umbreit, Burris and Stauffer (1957, p. 130). The insects were washed in an alcohol solution, according to the pro- cedure followed by Cotty (1956) to remove surface bacteria before homogeniza- tion. Insects were homogenized by means of a motor-driven glass homogenizer for 1 minute in 0.03 M phosphate buffer, except in the case of isocitric dehydro- genase, where veronal buffer was used since the phosphate ion interferes with the activity of this enzyme. The buffers were adjusted to a pH of 7.4. A3 per cent homogenate (1 ml) was incubated at 30°C for 30 minutes, and when NAD or NADP was used, the homogenate was pre-incubated with 0.5 ml of 0.2 per cent NAD or with 0.5 ml of 0.1 per cent NADP to oxidize the endogenous substrate. The smaller concentration of NADP was used because the addition of larger amounts did not increase enzyme activity. The homogenate was then placed in the side arm cap of the Thunberg tube. In the body of the tube were placed 1 ml of 1/10,000 per cent methylene blue, 1 ml of substrate (0.004 M), and a sufficient amount of buffer to bring the final volume to 6 ml. In measuring the activity of malic dehydrogenase, 0.5 ml of 0.24 M KCN was added to prevent inhibition by the oxalacetate formed (Green 1936). In determining the succinic dehydrogenase activity, 0.5 ml of a mixture of 0.005 M CaCD and 0.005 Alcl3 was added. NADP was used in the studies of the malic enzyme and of isocitric dehydrogenase. In the former determinations, 0.5 ml of 0.033 M MgS04, and in the latter, 0.5 ml of 6 X 10~3 M MnCl2 was added. These supplementary solu- tions were added before the final dilution of the homogenate. The tubes were evacuated for five minutes and were then inverted to add the homogenate con- tained in the side arm to the mixture in the main portion of the tube, thus bring- ing the final concentration of homogenate to 0.5 per cent. Following this the tubes were placed in a constant temperature bath at 30°C, and the time re- quired for 90 per cent reduction of methylene blue to occur was determined by visually matching the color with that of a standard tube. This standard con- tained all of the components of the other tubes except that the methylene blue was diluted to Vio the usual concentration and the homogenate had been previ- ously inactivated by boiling. A control tube containing all the components of the experimental tube except the substrate was used in each determination. Activities of dehydrogenase enzymes were expressed as 1/time in minutes for 90 per cent decoloration of methylene blue. These activities were determined by subtracting the rate of control from that of the experimental tube. The activity of cytochrome oxidase, expressed as A log (CyFe++) / minute, was determined during the same stages of metamorphosis and was measured on tissue homogenates in a final concentration of 1 : 10,000. The insects were homogenized in 0.03 M phosphate buffer which had a pH of 7.4. The spectrophotometric method of Cooperstein and Lazarow (1951) was used to measure the cytochrome oxidase activity. 122 New York Entomological Society [Vol. LXXV Table 1. Dehydrogenase activity expressed as 1/time in minutes for 90% decolorization of methylene blue during the metamorphosis of the face fly, Musca autumnalis. Readings were made at 30° C. (GPD is alpha-glycerophosphate dehydrogenase.) Dehydrogenase Stage Malic Glu- cose Alcohol Lactic Iso- citric Glu- tamic Suc- cinic GPD I GPD II Malic Enzymes Larva 0.375 0.006 0.055 0.024 0.345 0.008 0.019 0.061 0.005 0.120 Prepupa 0.328 0.004 0.050 0.022 0.316 0.006 0.016 0.050 0.006 0.114 Pupa, 1 day 0.280 0.004 0.040 0.018 0.322 0.006 0.008 0.020 0.008 0.105 Pupa, 2 day 0.252 0.006 0.031 0.015 0.282 0.003 0.008 0.011 0.010 0.082 Pupa, 3 day 0.228 0.010 0.022 0.009 0.230 0.002 0.005 0.005 0.010 0.060 Pupa, 4 day 0.230 0.009 0.038 0.006 0.218 0.005 0.004 0.012 0.096 Pupa, 5 day 0.260 0.006 0.046 0.010 0.202 0.009 0.018 0.012 0.104 Pupa, 6 day 0.345 0.005 0.058 0.005 0.180 0.012 0.029 0.025 0.110 Pupa, 7 day Adult, just 0.425 0.005 0.062 0.007 0.156 0.024 0.058 0.032 0.112 emerged 0.785 0.002 0.069 0.007 0.131 0.030 0.074 0.035 0.130 OBSERVATIONS The changes in the activities of the dehydrogenase enzymes during the metamor- phosis of the face fly are shown in Table 1. Each value is an average of ten determinations. Total alpha-glycerophosphate, alcohol, malic, and succinic dehydrogenases as well as the malic enzyme exhibited U-shaped activity. Greatest activity was shown by the malic dehydrogenase with a considerable rise observed in the newly emerged adult. Alpha-glycerophosphate dehydrogenase I (requiring NAD) decreased steadily from the larval stage to the fourth day and then rose gradually during the remainder of the pupal stage. Alpha-glycerophosphate II (not re- quiring NAD) appeared at the first day of the pupal stage and it showed a steady increase with the highest activity being detected in the adult fly. Isocitric dehydrogenase activity was high initially and remained high until 2 -day pupa and thereafter showed a progressive decline. Glucose dehydrogenase activity was very low; it remained fairly steady during the entire pupal stage and decreased slightly in the newly emerged adult. Alcohol dehydrogenase decreased steadily during the first days of the metamorphosis, reaching a low value on the third day, and then increased to reach its highest value in the adult stage. Succinic dehydrogenase exhibited a similar pattern but the level of activity was not as high as most of the other dehydrogenases. The activity of lactic dehydrogenase was low throughout metamorphosis. Malic enzyme exhibited high activities in the larva, prepupa and again in the adult stage. Glutamic dehydrogenase showed low activity in the larval stage. It decreased during the first several days of pupal life and completely disappeared by the fourth day. Cytochrome oxidase activity was also U-shaped during metamorphosis as in- dicated in Table 2. Each value here is also an average of at least ten determina- tions. The larval and prepupal stages were characterized by high activity with a September, 1967] Rousell: Respiratory Enzymes of Face Fly 123 Table 2. Cytochrome oxidase activity during the metamorphosis of Musca autumnalis. Homogenate concentration is 1:10,000. Stage Enzyme Activity A log [CyFe++] / min. Minimum Maximum Average Larva 0.051 0.103 0.084 Prepupa 0.043 0.088 0.061 Pupa, 1 day 0.024 0.042 0.032 Pupa, 2 day 0.014 0.038 0.021 Pupa, 3 day 0.009 0.022 0.014 Pupa, 4 day 0.008 0.017 0.010 Pupa, 5 day 0.027 0.048 0.041 Pupa, 6 day 0.052 0.089 0.076 Pupa, 7 day 0.112 0.151 0.127 Adult, newly emerged 0.124 0.221 0.178 progressive decrease until the fourth day and then a steady increase to a high of 0.178. DISCUSSION The U-shaped activities of malic dehydrogenase and the malic enzyme agree with the results reported for these enzymes during the metamorphosis of the mealworm and of the house fly (Ludwig and Barsa, 1958 and 1959). These findings also agree with those of Agrell (1949) for the blow fly, Calliphora erythrocephala , and of Diamantis (1962) for the Mediterranean flour moth, Ephestia kuhniella. Isocitric dehydrogenase activity in the face fly was slightly lower than that found in the house fly (Ludwig and Barsa, 1959), but similar in that it uniformly decreased during metamorphosis. This differs from the re- sults reported by Agrell (1949) for the blow fly and Diamantis (1962) for the Mediterranean flour moth, both of whom found that isocitric dehydrogenase ex- hibited U-shaped activity. Isocitric dehydrogenase in the presence of NADP and Mn++ catalyzes the oxidation of isocitrate through oxalosuccinate to alpha- ketoglutarate. The high activity of malic dehydrogenase is similar to that re- ported for the house fly by Ludwig and Barsa (1959) and for the flour moth by Diamantis (1962). Malic dehydrogenase and the malic enzyme both catalyze the oxidation of 1-malate. The end products with the malic enzyme are pyruvate and C02? whereas with malic dehydrogenase the end product is oxaloacetate. The high activities of malic dehydrogenase and the malic enzyme coupled with the rather low rate for total lactic dehydrogenase adds additional support to the belief that lactate does not accumulate in insects, but rather pyruvate is reduced to malate which in turn is oxidized to oxaloacetate. The U-shaped activity curves for alcohol and alpha-glycerophosphate I dehydrogenase agree with the results obtained by Ludwig and Barsa ( 1959) with the house fly, but alpha-glycerophos- phate II was found during all stages of metamorphosis in the face fly as con- trasted with the house fly where it does not appear until near the end of the 124 New York Entomological Society [Vol. LXXV pupal stage. The activity curve for succinic dehydrogenase corroborates re- ported results of a number of other insects including Drosophila melanogaster (Wolsky, 1941), Calliphora erythrocephala (Agrell, 1949), Musca domestica (Ludwig and Barsa, 1959), Tenebrio molitor (Ludwig and Barsa, 1958), Ephestia kuhniella (Diamantis, 1962). The low activity of this enzyme indicates that it could be a determining factor in the U-shaped respiratory curve that is character- istic of the metamorphosis of holometabolous insects. The U-shaped pattern of cytochrome oxidase activity here reported for Musca autumnalis agrees with what has been found in the fruit fly, D. melanogaster by Wolsky (1938), in the house fly, M. domestica by Sacktor (1951), in the Japa- nese beetle, P. japonica by Ludwig (1953), in the moth, Platysamia cecropia by Williams (1950), and in the flour moth, Ephestia kuhniella by Diamantis (1912). This would indicate that most of the oxidation during metamorphosis is medi- ated through the cytochrome system. Literature Cited Agrell, I. P. S. 1949. Localization of some hydrogen-activating enzymes in insects during metamorphosis. Nature, 164: 1039-1040. Bodine, J. H., and P. R. Orr. 1925. Respiratory metabolism. Biol. Bull., 48: 1014. Clare, M. R. 1925. A study of oxygen metabolism in Drosphila melanogaster. Biol. Bull., 49: 440-460. Cooperstein, S. J., and A. Lazarow. 1951. A microspectrophotometric method for the determination of cytochrome oxidase. Jour. Biol. Chem., 189: 665-670. Cotty, V. F. 1956. Respiratory metabolism of prepupae of the house fly, Musca domestica L., and of their homogenates. Contrib. Boyce Thompson Inst., 18: 253-262. Diamantis, W. 1962. Activities of respiratory enzymes during the metamorphosis of the Mediterranean flour moth, Ephestia kuhniella Zeller. Jour. N.Y. Ent. Soc., 70: 68-78. Dobzhansky, T., and D. F. Poulson. 1953. Oxygen consumption of Drosophila pupae. II. Drosophila pseudobscura. Z. Vergl. Physiol., 22: 473-478. Fink, D. E. 1925. Metabolism during embryonic and metamorphic development of insects. Jour. Gen. Physiol., 7: 527-543. Green, D. E. 1936. The malic dehydrogenase of animal tissue. Biochem. J., 30: 2095-2110. Ito, T. 1954. The physiology in the metamorphosis of Bombyx mori. I. Respiration. Bull. Sericult. Exp. Sta. (Tokyo), 14: 263-278. Krogh, A. 1914. On the rate of development and CO- production of chrysalides of Tenebrio molitor at different temperatures. Z. allg. Physiol., 16: 178-190. Ludwig, D. 1931. Studies on the metamorphosis of the Japanese beetle ( Popillia japonica Newman). I. Weight and metabolism changes. Jour. Exp. Zool., 60: 309-323. . 1953. Cytochrome oxidase activity during diapause and metamorphosis of the Japanese beetle ( Popillia japonica Newman). Jour. Gen. Physiol., 36: 751-757. — and M. C. Barsa. 1955. The activity of succinic dehydrogenase during diapause and metamorphosis of the Japanese beetle ( Popillia japonica Newman). Jour. N.Y. Ent. Soc., 63: 161-165. . 1956. Oxygen consumption of whole insects and insect homogenates. Biol. Bull., 110: 77-82. . 1958. Activity of dehydrogenase enzymes during the metamorphosis of the meal- worm, Tenebrio molitor Linnaeus. Ann. Ent. Soc. Amer., 49: 103-104. September, 1967] Rousell: Respiratory Enzymes of Face Fly 125 . 1959. Activities of respiratory enzymes during the metamorphosis of the house fly, Musca domestica Linnaeus. Jour. N.Y. Ent. Soc., 67: 151-156. Sacktor, B. 1951. Some aspects of respiratory metabolism during metamorphosis of normal and DDT-resistant house flies, Musca domestica L. Biol. Bull., 100: 229-243. Umbreit, W. W., R. H. Burris, and J. F. Stauffer. 1957. Manometric techniques. A manual describing methods applicable to the study of tissue metabolism. Minneapolis. Williams, C. M. 1950. A hormonal-enzymatic mechanism for control of pupal diapause in the Cecropia silkworm. Abstract of Communication to the XVIII Int. Physiol. Cong. (Copenhagen), 517-518. Wolsky, A. 1938. The effect of carbon monoxide on oxygen consumption of Drosophila melanogaster pupae. Jour. Exp. Biol., 15: 225-234. . 1941. Quantitative changes in the substrate-dehydrogenase system of Drosophila pupae during metamorphosis. Science, 94: 48-49. Received for publication April 3, 1967 Some Synonyms in American Spiders1 Wilton Ivie2 Abstract: New synonyms of one genus and twenty-four species, as well as twenty-one new combinations and a few other notes pertaining to American spiders, most of them in the family Linyphiidae, particularly the sub-family Erigoninae, are recorded. The following notes, concerned with new synonymy and new combinations in the nomenclature of American spiders, are presented herewith so that they may become part of the published record. Many of these notes were accumulated while examining the collections of The American Museum of Natural History, the Museum of Comparative Zoology at Harvard, Cornell University, Ithaca, New York, and parts of the type collections of the University of Utah and The United States National Museum. Cross references are given under the respective genera and families for the names mentioned in the text. Under the literature references, only those not included in Bonnet’s Bibliographia Araneorum are cited. Family CLUBIONIDAE Genus PHRUROTIMPUS Chamberlin and Ivie, 1935. Phrurotimpus alarms (Hentz), 1S47. Phrurotimpus annulatus Chamberlin and Ivie, 1944. New Synonym. Phrurotimpus borealis (Emerton), 1911. Phrurolithus utus Chamberlin and Ivie, 1933. Synonym. Family SALTICIDAE Genus LYSSOMANES Hentz, 1844. Lyssomanes viridis (Walckenaer) , 1837. 7 'etragnatha lutea Walckenaer, 1841. New Synonym. Family THERIDIIDAE Genus DIPOENA Thorell, 1870. PSELOTHORAX Chamberlin, 1948 (Erigonidae) . New Synonym. Dipoena atopa (Chamberlin). New Combination. Pselothorax atopus Chamberlin, 1948. Dipoena daltoni Levi, 1953. New Synonym. Family TETRAGNATHIDAE Genus TETRAGNATHA Latreille, 1804. T etragnatha lutea Walckenaer. See Lyssomanes viridis (Salticidae) 1 This paper was prepared as a phase of a project supported by funds from the National Science Foundation (Grant GB-3880) 2 Research Fellow, Department of Entomology, The American Museum of Natural History, New York. 126 September, 1967] Ivie: American Spider Synonyms 127 Family LINYPHIIDAE Sub-Family Erigoninae Genus ACARTAUCHENIUS Simon, 1884. Acartauchenius columbiensis Crosby. See Maso polita. Genus CERATICELUS Simon, 1884. Ceraticelus anomalus Gertsch and Ivie. See Idionella anomala. Ceraticelus desertus Gertsch and Ivie. See Idionella deserta. Ceraticelus jormosus (Banks). See Idionella jormosa. Ceraticelus guttatus Chamberlin and Ivie. See Idionella anomala. Ceraticelus micropalpus (Emerton). Ceraticelus durus Chamberlin and Ivie, 1939. New Synonym. Ceraticelus nesiotes Crosby. See Idionella nesiotes. Ceraticelus parvulus (Fox). See Ceratinella parvula. Ceraticelus rugosus Crosby. See Idionella rugosa. Ceraticelus titivillitium Crosby and Bishop. See Idionella titivillitium. Ceraticelus tuganus Chamberlin. See Idionella tugana. Genus CERATINELLA Emerton, 1882. Ceratinella brunnea Emerton, 1882. Ceratinella placida Banks, 1892. New Synonym. Ceratinella jormosa Banks. See Idionella jormosa. Ceratinella parvula (Wm. Fox). Erigone ( Ceratinella ) parvula Fox, 1891. Ceratinella sphaerula Emerton, 1911. New Synonym. Ceraticelus parvulus: Crosby and Bishop, 1925. Genus CERATINOPS Banks, 1905. Ceratinops obscura (Chamberlin and Ivie). New Combination. Masonetta obscura Chamberlin and Ivie, 1944. Genus CERATINOPSIS Emerton, 1882. Ceratinopsis disparata (Dondale). New Combination. Grammonota disparata Dondale, 1959. This species is very close to, if not identical with, Ceratinopsis labradorensis Emerton, 1925. Ceratinopsis tybeensis Chamberlin and Ivie. See Masonetta jloridana. Genus CORNICULARIA Menge, 1868. Cornicularia lepida Kulczynski, 1885. Kamchatka. Cornicularia pacijica Emerton, 1923. New Synonym. C ornicularia selma Chamberlin. See Scylaceus selma. Genus EPERIGONE Crosby and Bishop, 1928. Eperigone trilobata (Emerton). Bathyphantes tristis Banks, 1892. Synonymy suggested by Hackman, 1954; con- firmed by examination of type material. Genus EPICERATICELUS Crosby and Bishop, 1931. Epiceraticelus jluvialis Crosby and Bishop. Scylaceus amylus Chamberlin, 1948. New Synonym. Genus ERIDANTES Crosby and Bishop, 1933. Eridantes erigonoides (Emerton). Erigone percisa Keyserling, 1886. New Synonym. Genus ERIGONE Audouin, 1827. Erigone atra (Blackwall), 1833. 128 New York Entomological Society [Vol. LXXV Erigone praepulchra Keyserling, 1886. Synonym. Erigone matei Keyserling. See 0 stearins melanopygius. Erigone minutissima Keyserling. See Scylaceus pallidus. Erigone nigrianus Keyserling. See 0 stearins melanopygius. Erigone percisa Keyserling. See Eridantes erigonoides. Erigone rostratula Keyserling. See Scylaceus pallidus. Genus EULAIRA Chamberlin and Ivie, 1933. Eulaira microtarsus (Emerton). See Hillhousia microtarsus. Genus GONEATARA Bishop and Crosby, 1935. Goneatara nasuta (Barrows). New Combination. Souessa nasuta Barrows, 1943. Genus GRAMMONOT A Emerton, 1882. Grammonota disparata Dondale. See Ceratinopsis disparata. Grammonota sclerata Ivie and Barrows. See Idionella sclerata. Genus HILAIRA Simon, 1884. Hilaira balia Crosby and Bishop, 1929. South America. Microneta maculata Mello-Leitao, 1940. New Synonym. Genus HILLHOUSIA F. P. -Cambridge, 1894. Hillhousia microtarsus (Emerton). New Combination. Tmeticus microtarsus Emerton, 1882. Eulaira microtarsus: Chamberlin and Ivie, 1945. Sciastes microtarsus Bishop and Crosby, 1938. Genus IDIONELLA Banks, 1893. Type: jormosa. Idionella anomala (Gertsch & Ivie). New Combination. Ceraticelus anomalus Gertsch and Ivie, 1936. Male. Ceraticelus guttatus Chamberlin and Ivie, 1939. Female. New Synonym. Idionella deserta (Gertsch and Ivie). New Combination. Ceraticelus desertus Gertsch and Ivie, 1936. Idionella jormosa (Banks). Ceratinella jormosa Banks, 1892. Ceraticelus jormosus : Crosby and Bishop, 1925. Idionella nesiotes (Crosby). New Combination. Ceraticelus nesiotes Crosby, 1924. Idionella rugosa (Crosby). New Combination. Ceraticelus rugosus Crosby, 1905. Idionella sclerata (Ivie and Barrows). New Combination. Grammonota sclerata Ivie and Barrows, 1935. Ceraticelus jormosus : Crosby, 1937 (in part; male, not female). Idionella titivillitium (Crosby and Bishop). New Combination. Ceraticelus titivillitium Crosby and Bishop, 1925. Idionella tugana (Chamberlin). New Combination. Ceraticelus tuganus Chamberlin, 1948. Genus ISLANDIAN A Braendegaard, 1932. Islandiana jalsijica (Keyserling). New Combination. Erigone jalsijica Keyserling, 1886. Tmeticus alatus Emerton, 1919. New Synonym. Islandiana alata: Ivie, 1965. Genus MASO Simon, 1884. Maso polita Banks, 1896. September, 1967] Ivie: American Spider Synonyms 129 Acartauchenius columbiensis Crosby, 1905. New Synonym. Genus MASONCUS Chamberlin, 1948. Masoncus conspectus (Gertsch and Davis). New Combination. Tapinocyba conspecta Gertsch and Davis, 1936. Masoncus nogales Chamberlin, 1948. New Synonym. Genus MASON ETTA Chamberlin and Ivie, 1939. Masonetta floridana (Ivie and Barrows), 1935. Ceratinopsis tybeensis Chamberlin and Ivie, 1944. New Synonym. Masonetta obscura Chamberlin and Ivie. See Ceratinops obscura. Genus OEDOTHORAX Bertkau, 1883. Oedothorax melacra Chamberlin. See Ostearius melanopygius. Genus OSTEARIUS J. E. Hull, 1911. Ostearius melanopygius (O. P. Cambridge). Linyphia melanopygia O. P. Cambridge, 1879. Erigone matei Keyserling, 1886. New Synonym. Erigone nigrianus Keyserling, 1886. New Synonym. Oedothorax melacra Chamberlin, 1916. New Synonym. Scolopembolus melacrus: Bishop and Crosby, 1938. Genus PSELOTHORAX Chamberlin, 1948. See DIPOEN A, (Theridiidae) . Pselothorax atopus Chamberlin. See Dipoena atopa, (Theridiidae). Genus SCIASTES Bishop and Crosby, 1938. Sciastes microtarsus (Emerton). See Hillhousia microtarsus. Sciastes ogeechee Chamberlin and Ivie. See Souessoula parva. Sciastes terrestris (Emerton). See Porrhomma terrestris (Linyphiinae) Genus SCOLOPEMBOLUS Bishop and Crosby, 1938. Scolopembolus melacrus (Chamberlin). See Ostearius melanopygius. Genus SCYLACEUS Bishop and Crosby, 1938. Scylaceus amylus Chamberlin. See Epiceraticelus fluvialis. Scylaceus pallidus (Emerton), 1882. Erigone minutissima Keyserling, 1886. New Synonym. Erigone rostratula Keyserling, 1886. New Synonym. Scylaceus pallas Chamberlin, 1948. New Synonym. Scylaceus selma (Chamberlin). New Combination. Cornicularia selma Chamberlin, 1948. Genus SISICOTTUS Bishop and Crosby, 1938. Sisicottus atypicus Chamberlin and Ivie. See Souessoula parva. Genus SOUESSA Crosby and Bishop, 1936. Souessa nasuta Barrows. See Goneatara nasuta. Genus SOUESSOULA Crosby and Bishop, 1936. Souessoula parva (Banks), 1899. Sciastes ogeechee Chamberlin and Ivie, 1944, female. New Synonym. Sisicottus atypicus Chamberlin and Ivie, 1944, male. New Synonym. Genus T ACHYGYN A Chamberlin and Ivie, 1939. Tachygyna gargopa (Crosby and Bishop). New Combination. Microneta gargopa Crosby and Bishop, 1929. Genus TAPINOCYBA Simon, 1884. Tapinocyba conspecta Gertsch and Davis. See Masoncus conspectus. Genus TMET1CUS Menge, 1886. Tmeticus alatus Emerton. See Islandiana falsifica. 130 New York Entomological Society [Vol. LXXV Tmeticus microtarsus Emerton. See Hillhousia microtarsus. Tmeticus terrestris Emerton. See Porrhomma terrestris (Linyphiinae) . Sub-Family Linyphiinae Genus ALLOMEN GEA Strand, 1912. Allomengea pinnata (Emerton). New Combination. Microneta pinnata Emerton, 1915. Microneta plumosa: Emerton, 1915 (lapsus in caption of figure for M. pinnata.) Linyphia ontariensis Emerton, 1925. New Synonym. Helophora ontariensis : Blauvelt, 1936; Chamberlin and Ivie, 1947. Allomengea scopigera (Grube). Linyphia sitkaensis Keyserling, 1886. New Synonym. Genus BATHYPHANTES Menge, 1866. Bathyphantes pacificus Banks. See Liny phantes pacificus. Bathyphantes tragicus Banks. See Liny phantes tragicus. Bathy phantes tristis Banks. See Eperigone trilobata. Genus LEPTHYPH ANTES Menge, 1866. Lepthy phantes sabulosus (Keyserling), 1886. Lepthy phantes appalachia Chamberlin and Ivie, 1944. New Synonym. Type locality of sabulosus given as Salt Lake City, Utah; probably incorrect. Genus L1NYPH ANTES Chamberlin and Ivie, 1942. Linyphantes aeronautica (Petrunkevitch) . New Combination. Microneta aeronautica Petrunkevitch, 1929. Linyphantes orcinus (Emerton). New Combination. Microneta orcina Emerton, 1917. Linyphantes pacificus (Banks). New Combination. Bathyphantes pacificus Banks, 1905. Linyphantes tragicus (Banks). New Combination. Bathyphantes tragicus Banks, 1898. Baja California. Genus LINYPHIA Sundevall, 1804. Linyphia melanopygia O. P. Cambridge. See Ostearius melanopygius (Erigoninae) . Linyphia ontariensis Emerton. See Allomengea pinnata. Linyphia sitkaensis Keyserling. See Allomengea scopigera. Genus MICRONETA Menge, 1868. Microneta aeronautica Petrunkevitch. See Linyphantes aeronautica. Microneta gargopa Crosby and Bishop. See Tachygyna gargopa. Microneta maculata Mello-Leitao. See H Hair a balia (Erigoninae) Microneta orcina Emerton. See Linyphantes orcinus. Microneta pinnata Emerton. See Allomengea pinnata. Microneta plumosa Emerton. See Allomengea pinnata. Genus PORRHOMMA Simon, 1884. Porrhomma terrestris (Emerton). New Combination. Tmeticus terrestris Emerton, 1882. Sciastes terrestris: Bishop and Crosby, 1938 (Erigoninae). Literature Cited Barrows, W. M. 1943. A New Prairie Spider. Ohio Jour. Science, 43, p. 209. Chamberlin, R. V. 1948. On Some Spiders of the Family Erigonidae. Ann. Ent. Soc. Amer., 41, pp. 483-562, 163 figs. September, 1967] Ivie: American Spider Synonyms 131 Chamberlin, Ralph V., and Wilton Ivie. 1939. Studies on North American Spiders of the Family Micryphantidae. Verh. VII Int. Kongr. Ent., 1, pp. 56-73, 59 figs. . 1942. A Hundred New Species of American Spiders, Bull. Univ. Utah, Biol. Ser., 7, No. 1, pp. 1-117, 231 figs. . 1944. Spiders of the Georgia Region of North America. Ibid., 8, No. 5, pp. 1-267, 217 figs. Dondale, C. D. 1959. Definition of the Genus Grammonota. Canadian Ent., 91, No. 4, pp. 232-242, 26 figs. Hackman, Walter. 1954. The Spiders of Newfoundland. Acta Zool. Fennica, No. 79, pp. 1-99, 121 figs., 5 maps. Ivie, Wilton. 1965. The Spiders of the Genus Islandiana. Amer. Mus. Novitates, No. 2221, pp. 1-25, 53 figs. Levi, Herbert W. 1953. Spiders of the Genus Dipoena from America North of Mexico. Amer. Mus. Novitates, No. 1647, pp. 1-39, 121 figs. Received for publication May 1, 1961 Biology of Dufourea and of its cleptoparasite, Neopasites (Hymenoptera: Apoidea) Philip F. Torchio,1 Jerome G. Rozen, Jr.,2 George E. Bohart,1 and Marjorie S. Favreau2 Abstract: The biologies of four species of Dufourea [ D . mulleri (Cockerell), D. mal- acothricis Timberlake, D. pulchricornis (Cockerell), and D. trochantera Bohart] are de- scribed and compared. The biology of the nomadine bee parasite, N eopasites, family Anthophoridae, is also described. Two species of the parasite are associated with their hosts [Neopasites ( Micropasites ) cressoni Crawford with D. mulleri, and an undescribed species of the subgenus Neopasites with D. trochantera] . The suspected association of an additional species, Neopasites ( Neopasites ) fulviventris (Cresson), on D. dentipes Bohart and an undescribed Dufourea species is included. The subfamilies of Halictidae are com- pared on the basis of biological features in a summary table. The family Halictidae (composed of Halictinae, Nomiinae, and Dufoureinae) is well represented in the biological literature. Most of the information, how- ever, concerns halictines and nomiines. Previous biological studies of the Dufoureinae have been restricted to six species within two Old World genera: Rophites canus Evers (Enslin, 1921; Malyshev, 1925a), Rophites hartmanni Friese (Malyshev, 1925a), R. quin ques pin osus Spinola (Stockhert, 1922), Systropha planidens Giraud and S. curvicornis Scopoli (Malyshev, 1925b), and S. punjabensis Batra and Michener (Batra and Michener, 1966). The holarctic genus, Dufourea , has not been studied biologically, even though it is widely distributed and contains the greatest number of species in the sub- family. The biologies of four Dufourea species (D. mulleri (Cockerell), D. malacothricis Timberlake, D. pulchricornis (Cockerell), and D. trochantera Bohart) are reported. The biology of the New World Neopasites (= Gnathopasites) is also de- scribed. It and its Old World counterpart, Biastes, comprise the nomadine tribe Biastini which are cleptoparasitic primarily on the Dufoureinae. Biastes attacks the nests of Rophites, Systropha , and presumably the eucerine Tetra- lonia (Popov, 1951), and Neopasites attacks those of Dufourea. The literature search for this paper was made with the assistance of the Bibliography of Apoid Biology under the direction of Dr. Charles D. Michener, the University of Kansas, Lawrence. Dufourea mulleri (Cockerell) Description of Habitat: Bohart, Torchio, and Nabil Youssef studied the biology of this species at Tubac, Santa Cruz County, Arizona, between April 1 Entomology Research Division, Agr. Res. Serv., USDA, Logan, Utah, in cooperation with Utah Agricultural Experiment Station. 2 Department of Entomology, the American Museum of Natural History, N. Y., N. Y. 132 September, 1967] Torchio, et al.: Biology of Dufourea 133 Fig. 1. Nesting area of Dufourea mulleri (Cockerell) at 3 miles south-southwest of Rodeo, Hidalgo County, New Mexico. 13 and 17, 1965. Torchio returned to this site on April 27, 1965, and found nesting had been completed. On April 26, 1966, he revisited the nesting site and discovered the nesting population greatly reduced over the previous year. Rozen studied the species 3 miles S.S.W. of Rodeo, New Mexico (Fig. 1) (actually in Cochise County, Arizona), between May 1 and 5, 1965. Rozen and Favreau revisited this site between April 26 and May 5, 1966, at which' time the species was more abundant and nested in various areas along the road between this point and Apache, Arizona. The Tubac site was located adjacent to a gravelly creek bottom which carried water during short periods each year. Phacelia of two species, Les- querella , Malacothrix, Acacia greggii Gray, a tall crucifer, and several grass species were the predominant plants growing along the creek. The surrounding area is typical of the Lower Sonoran. The Rodeo site, a recently disturbed, nearly flat area, half a mile long, was adjacent to a highway running in a S.S.W. direction through the wide San Simon Valley. The nest area was occupied by low, sparsely scattered herbs, including the pollen plant, Phacelia popei T. & G. var. arizonica (Gray) Voss, and a Lepidium species. The vegeta- tion adjoining the nest area was dominated by Prosopis and other xerophilous plants. The soil surface at both nesting sites was unshaded and ranged from horizontal or nearly so near Rodeo to gently sloping (up to 15°) at Tubac. At Tubac, nesting took place in two soil types. One had a 6 mm. layer 134 New York Entomological Society [Vol. LXXV of dry, loose powder covering a hard-packed, sandstone-like layer composed of brownish soil interspersed with' large gravel particles. The hard-packed layer extended below the cell level and contained some moisture below 4.6 cm. The second soil type was light brown, coarsely grained, and loosely packed to 5 cm. below its surface. Large, extremely hard-packed clods found below the surface layer were separated from each other by air spaces or narrow bands of loosely-packed soil. The soil was dry until well below the cell level. The nesting site near Rodeo was sandy and loosely packed from its surface to a depth of 3-4 cm., below which it became hard-packed and pebble-free. Moisture at the cell level varied from slight to moderate, depending upon the depth. Soil temperatures recorded from this site at a depth of 10 cm. on April 24, 1966, were: 9:30 a.m., 69°F; 10 a.m., 69°F; 12:30 p.m., 78°F; 3:15 p.m., 80°F. The time is Rocky Mountain Standard time and the day was clear and warm. Although nests were scattered over extensive areas at both locations, nest concentrations also occurred. The most dense concentrations numbered V2 nest/ sq. ft. at Tubac and 4 nests/sq. ft. near Rodeo. Apparently, the species can be regarded as weakly gregarious. Only a single female occupied a nest. Nest Architecture entrance hole: Some nest entrances occurred in flat, bare ground, but more frequently they were at the lower edges of slight depressions or at the bases of pebbles or rocks. Soil excavated from the nests was deposited on one side of the entrance, forming an asymmetrical tumulus. The typical tumulus at Tubac was heart-shaped and measured 33 mm. long by 27 mm. wide. A weakly defined trail 4 mm. wide, 2 mm. deep, and 18 mm. long extended from the entrance hole to near the apical angle of the tumulus. It was formed as the female swept excavated soil away from the entrance while she backed away repeatedly over the same terrain. At the terminus of the trail, the excavated soil was kicked back and away with rapid, flicking leg movements. The tumulus was continually reshaped and enlarged throughout the period of nesting activity. Entrances were generally kept closed at Tubac but remained open near Rodeo. Possibly the divergent behavior at each location is simply a reflection of adaptability to nesting in different soil types. At Tubac the very loose surface powder tended to fill the entrance holes each time bees entered or left. Returning foragers, however, were able to orient to their respective entrance holes very successfully. They literally dove into the powdered layer, as do some N omadopsis species, and rapidly moved soil about until they found and entered their burrow. The soil near Rodeo was sufficiently granular and hard-packed to allow the entrance holes to remain open. Entrances always lacked turrets. September, 1967] Torchio, et al.: Biology of Dufourea 135 burrows: The main burrow, circular in cross section, was 3.5 mm. in diameter and descended in a meandering fashion. There were no obvious constrictions at or near the entrance hole. The burrow walls were not lined but, at least at the Tubac site, they appeared darker in color and were more tightly packed than the surrounding soil. Their permeability to water was equal to that of the surrounding soil. A vestibule measuring 7 mm. in diameter was found in one nest at Tubac. It was constructed as a pocket in the wall of the main burrow 11 mm. below the soil surface. The main burrow was never plugged and it terminated in a nearly horizontal cell. Lateral burrows were originated along a 15 mm. zone about halfway down the main burrow. The unlined laterals (as many as 9 per nest) radiated horizontally from the main burrow for distances ranging from 5 to 38 mm. Circular in cross section, they had the same diameter as the main burrow except where they narrowed to 3 mm. just before joining the cell. Each lateral was plugged tightly before a new one was excavated. cells: The cells (Figs. 2-6), which were ovoid and broadly rounded distally, were placed from 10 to 40 degrees from the horizontal with the anterior end highest. Their length varied from 6.0 to 8.0 mm. and their width from 4.5 to 5.0 mm. They were carved from the surrounding soil and their inner sur- faces had no apparent “built-in” wall. They were, however, lined with a dull varnish that was nearly transparent upon drying. This lining, less than 0.05 mm. in thickness, filled the space between the sand grains and could not be peeled from the walls of their cells. The lining permitted a moderately rapid absorption of water when a droplet was placed on it. At the Rodeo site a very thin layer of dull, extremely fine, silt-like material coated the depres- sions between the grains of sand. Cells were located between 5 and 10 cm. from the ground surface, with the uppermost cell being excavated first and the lowermost cell last. Cells from previous years were not reconditioned and reused. The unlined cell cap was composed of a moderately packed soil plug which had 3 indistinct spiral rings and a small central micropyle on its concave inner face. Although only one cell per lateral burrow was found at Tubac, two cells (and in one case, three cells) in linear series were commonly found at the end of the lateral burrows near Rodeo. The passage between these cells varied in length from 2.0 to 5.0 mm., and was filled with rather loosely packed soil between the firm rear wall of one cell and the firm cap of the other. Provisioning and Development D. mulleri provisioned its nests with pollen from two Phacelia species at Tubac. One species produced blue pollen and the other, yellow. Since the pollen balls were always either one color or the other, it appears that the bees visited only one host plant species while provisioning a cell. Phacelia popei T. and G. 136 New York Entomological Society [Vol. LXXV var. arizonica (Gray) Voss was the only pollen host near Rodeo. Its dry pollen remained bluish in color while on the bee’s scopae but changed to lavender after it was molded into the provisions. The color of the pollen ball faded to a light tan by the time the first instar hatched from the egg. Approximately three pollen loads were required to complete one pollen ball. The first load, after being transported to the cell, was mixed with nectar and shaped into a small but complete sphere. Each additional load was added to the existing sphere until it became a moist but firm, homogeneous, spherical ball, averaging about 3.5 mm. in diameter and ranging from 2.75 to 3.75 mm. The ball was placed near, but not at, the posterior end of the cell (Fig. 2). The pollen balls of this species resembled those of the panurgine genera N omadopsis and Calliopsis in shape and consistency, but lacked a waterproof covering. The shiny, whitish, strongly arched egg (Fig. 2) was approximately 1.9 mm. long and rested on top of the provision in the sagittal plane of the cell. Both ends were weakly attached to the provisions so that eggs were easily displaced when cells were excavated. In contrast, the eggs of some bees (e.g., certain Panurginae) are attached securely by their posterior ends while the anterior tip rises in the cell or merely touches the provision. In mulleri the broader anterior tip of the egg faced the cell closure. Immediately before the first instar hatched, the egg chorion adhered to the embryo, so that the rather small head and body segmentation were visible on the still strongly arched egg. After hatching, the larva fed and crawled about on the provisions (Fig. 3). The first-stage larva, as well as subsequent ones, was equipped with a pair of dorsolateral tubercles on most body seg- ments, with a somewhat protruding venter on the ninth abdominal segment, and with a posterodorsally directed tenth abdominal segment which could be contracted and expanded somewhat. These modifications assisted the larva as it crawled; by appressing the protruding ninth segment to the pollen ball and the expanded tenth to the cell wall, the larva stationed the posterior part of the body so that it could push its front part forward. While moving forward and bending the anterior portion of its body up and down, the larva fed on the pollen ball and left a wide, shallow groove in its wake. Because the feed- ing larva circled its provisions in random directions, the ball remained nearly spherical almost until it disappeared (Figs. 3-4). After consuming the pollen ball, but before defecating, the larva began spinning a cocoon which, when completed, tightly adhered to the cell walls. When the outer layer of the cocoon was completed, the larva extruded long semi-moist, pale yellow fecal pellets which were applied to the posterior one- half to two-thirds of the cocoon in short strips more or less parallel to the long axis of the cell. During or after the late stages of defecation, the larva applied additional silk over the inner face of the outer cocoon layer and feces September, 1967] Torchio, et al.: Biology of Dufourea 137 Figs. 2-6. Cells of Dufourea mulleri (Cockerell): 2. With pollen ball and egg, side view. 3. With pollen ball and young larva, side view. 4. With nearly mature larva, side view. 5. With postdefecating larva, cocoon, and feces, side view. 6. In linear series, top view. Figs. 7-9. Eggs of Neopasites {Micro pasites) cressoni Crawford: 7. Embedded nearly flush with cell wall, lateral view. 8. Embedded at an angle with cell wall, lateral view. 9. After hatching, showing semicircular split at anterior end, dorsal view. until a complete inner cocoon layer was formed. This very thin inner layer completely isolated the larva from its feces. Most of the fecal pellets, although flattened into ribbons by the pressure of the larva, were still distinguishable. The completed cocoon (Fig. 5) was composed of two layers and assumed the same shape and dimensions as the cell. The parchment-like outer layer was dull, light brown on both of its surfaces but somewhat darker across its anterior face, where it was thicker. The inner layer was composed of a clear matrix interspersed with silk strands. It was very thin and tightly appressed to the inner face of the outer layer except where it incorporated and covered the fecal cake. The exposed surface of this layer was glossy. The cocoon was not 138 New York Entomological Society [Vol. LXXV supplied with a nipple, but individual thread-like silk strands were detected. Soft and easily collapsed anteriorly, it was more rigid where the feces gave it additional support. After the cocoons were spun, cells were difficult to find because they no longer broke open easily during excavation. Although the cocoons imparted extra strength to the cells, the mature larvae may also have secreted a harden- ing substance that permeated the soil adjacent to the cell. In any event, the wall of a cell occupied by a cocoon seemed much tougher than that of a cell containing an egg, an early instar, or an immature of Neopasites, which does not spin a cocoon. Adult Activity D. mulleri and Neopasites cressoni Crawford began flying between 9:00 a.m. and 9:30 a.m. M.S.T. on a warm clear day, and were still active at 2:30 p.m. Males of D. mulleri , presumably in search of mates, were often seen flying swiftly from host plant to host plant. Mating, observed only once, occurred near some host plants and was completed in 5 seconds. The bees did not fly in copula and mating was never observed at the nesting site. Associates Eurystylops (Strepsiptera) was discovered at Tubac as mature females in the abdomens of adult bees and as first instar larvae on the eggs. In one area of the same site, 90 percent of the bee cells contained dead first instars and were infested with a mold complex, including the genus Rhizoctoniump The biology of Neopasites cressoni , which attacked D. mulleri at both nest- ing sites, is described near the end of this paper. One burrow of D. mulleri possessed a unique feature in that it branched at the 2.5 cm. level. The branch, 2.5 mm. in diameter, led to two somewhat smaller cells containing a predefecating and a postdefecating larva belonging to the panurgine genus Perdita. They may well have been Perdita sexmaculata Cockerell, as this species was the only one abundant in the area at that time. Although the Dujourea female was still provisioning its part of the nest whereas the Perdita was not, it is impossible to say which had first started the nest because some offspring of both females had become mature larvae. Dujourea trochanter a Bohart Description of Habitat This species, which is closely related to D. mulleri , was discovered by Torchio nesting gregariously at Newton Dam, Cache County, Utah, on May 27, 1966. The nesting site was located on a 10-foot high, south-facing embankment inclined about 55° from horizontal. The site was made available recently when two Identified by G. M. Baker, Botany Dept., Utah State University. September, 1967] Torchio, et al.: Biology of Dufourea 139 roads converging near the nesting site were cut below the natural terrain of the hillside leading to the reservoir. Nests were mostly confined to an unvegetated 10-foot wide area of the embankment, and most entrances were situated toward the crest of the slope. A few nests were established at the top edge of the embankment where the grade was almost horizontal. Flowering plants growing in the vicinity of the nesting site were: Cirsium lanceolatum (L.), Hill, Oenothera sp., Brassica sp., Sphaeralcea sp., Salix sp., Penstemon sp., and Phacelia leucophila Torr. D. trochantera was utilizing Phacelia leucophila as its pollen and nectar source. The surface layer of the nesting site was composed of a fine, black powdered soil, ranging from 5 to 10 mm. in depth. Below this the black, clay soil be- came extremely hard-packed and contained numerous pebbles and rocks of varying sizes. The soil was dry to below the cell level. Nest Architecture entrance hole: Entrance holes were inclined at 45° angles from the horizontal regardless of the slope characteristics. All entrances, including those on the horizontal surface, faced south to southwest and were kept open. At times, however, winds disturbed the surface layer sufficiently to cause closure of some nests. Returning females associated with these nests landed near the plugged entrances and dug until the burrows were re-exposed. Nesting females kicked excavated soil from the entrances of nests located on steep slopes until indistinct tumuli were deposited below the nests as long strips of soil. If entrances were located on the horizontal surface, each nesting female dragged excavated soil from the nest repeatedly over the same course until a trough or trail was formed. A typical tumulus measured 36 mm. long and 13 mm. wide. The trough was 3 mm. wide and extended about half the length of the tumulus. No obvious constriction or expansion of the burrow occurred at or near the entrance hole. burrows: The unlined, unplugged main burrows were 3.5 mm. in diameter. They descended to depths ranging from 3 to 5 cm. When unobstructed, they spiraled downwards but were often forced to detour around pebbles and rocks. In two of the 25 nests excavated, an unlined vestibule was placed as a carved outpocket on a sharp turn of the main burrow. One of these measured 14 mm. wide by 9 mm. deep, and the second measured 7.5 mm. in diameter. The lateral burrows were also unlined and of the same diameter as the main burrow. They originated along the main burrow at different points and meandered for distances of 4 to 42 mm., where they terminated at cells be- tween 5 and 10 cm. below the surface. The laterals were tightly plugged after the cells were capped. We were unable to determine whether the main burrow terminated at a cell and was subsequently plugged for several centimeters or whether it divided into two or more laterals that were eventually plugged. 140 New York Entomological Society [Vol. LXXV cells: Cells of this species were remarkably similar in shape, form, and manner of construction to those of D. mulleri. The cell lining differed from that of D. mulleri in its somewhat greater impermeability to water. The cell cap, 3 mm. in diameter and composed of a moderately well-packed soil plug, was slightly concave. The unvarnished inner face had two to three indistinct rings surrounding a central micropyle, while the flat outer face had a smooth, unvarnished surface. Most of the lateral burrows terminated at single cells, but others (about 25 percent) led to two cells in linear series. The cells were usually subhorizontal but sometimes dipped to as much as 30° below the horizontal. The passages between those cells in linear series were plugged with soil that varied from loosely to tightly packed. Provisioning and Development The provisions of this species were similar to those of D. mulleri except for their tan color and slightly smaller average size (2.85 to 3.25 mm.). The eggs appeared to be slightly smaller than those of D. mulleri (1.8 mm. long by 0.4 mm. wide) but the samples may have been too small for a reliable comparison. Embryonic development, hatching, and larval shape and mobility all ap- peared to be identical with the same features in D. mulleri. Cocoon formation and structure were quite similar to those of D. mulleri. However, the following differences appeared to be consistent: (1) The outer cocoon layer was somewhat thicker and darker brown toward the anterior end, and there was a very thin, translucent zone about 2 mm. wide anterior to the fecal cake; (2) the fecal pellets composing the fecal cake were more completely fused into a single sheet. Adult Activity During warm, sunny weather, D. trochantera began flying at about 8:30 a.m. M.S.T. By 1:30 p.m. almost all flight ceased. Pollen loads were acquired in from 5 to 18 minutes and the time spent within the nest between loads varied from 2.5 to 36 minutes. This variation in time spent in the field and within the nest appeared to have no correlation with the time of day. Associates In the course of about 15 hours of observation at the nesting site, only two adults of an undescribed species in the subgenus Neo pasties were seen. Sur- prisingly, four of the approximately 40 host cells examined contained quiescent, postdefecating larvae of the parasite. The limited biological information obtained agreed with that of Neopasites cressoni discussed in a separate section below. One cell of D. trochantera contained four dipterous larvae which were con- suming the provision. Unfortunately, this cell was lost in transit from the field to the laboratory. September, 1967] Torchio, et al.: Biology of Dufourea 141 In 1962 a series of Neopasites adults were collected at a D. trochantera site on the Independence Lake Road, Sierra County, California, by M. E. Irwin. We compared specimens from both the California and Utah sites and found them to be distinct but undescribed species. Dufourea malacothricis Timberlake This species, smaller than D. mulleri , was collected from flowers of Mala- cothrix near Rodeo between April 26 and May 5, 1966. It was somewhat less common than D. mulleri , with which it flew, and only two nests were dis- covered by Favreau and Rozen, one at the Rodeo site (described above) and the other in an open area 3 miles north of Apache, Cochise County, Arizona. Nest Architecture entrance hole and burrow: The nest entrance and main burrow near Rodeo remained opened and bore an asymmetrical tumulus. The main burrow was 2.25 mm. in diameter and meandered a short distance before it was lost in the excavation. The second nest occurred on unshaded, nearly horizontal terrain with a 2 cm. -deep surface layer composed of rather loosely packed soil. The soil below was compacted sand free of pebbles. The cells were 10 and 12 cm. deep where the soil was moist. The unlined main burrow, 3.0 mm. in diameter, descended in a meandering fashion to a number of unlined and completely plugged lateral burrows of the same diameter. These laterals were horizontal or somewhat descending and were 4.0 to 4.5 mm. long, although one extended 10 mm. cells: Twelve cells were uncovered from the seven laterals associated with the one nest. Two cells were placed singly and the other 10 were grouped into linear series of two each. The distance between pairs in a series varied from 1.0 to 2.0 mm. All cells were inclined from 10 to 15 degrees from the horizontal with the rear of the cell lower than the front. They were identical in shape to those of D. mulleri and had the same type of lining and con- struction. The lining, however, was even less waterproof than that of D. mulleri in that it almost immediately absorbed a droplet of water. Cells varied from 5.0 to 5.5 mm. in length and from 3.5 to 4.0 mm. in maximum diameter. They were closed with a spiral plug, as in the case of D. mulleri , and the oldest cell was closest to the surface. Provisioning and Development The pollen balls of D. malacothricis differed from those of D. mulleri only in being yellow and in having a smaller diameter (2.75 to 3.0 mm.). The smaller eggs (1.75 mm. long) were identical in shape and placement with those of D. mulleri , and developing larvae practiced the same feeding habits. Unfortunately, no cocoons of this species were obtained. 142 New York Entomological Society [Vol. LXXV Associates The one nest excavated was free of parasites and predators even though N eopasites cressoni occurred in the area. Du jourea pulchricornis ( Cockerell ) Description of Habitat Bohart and Torchio found this species collecting pollen from Lesquerella gordoni (A. Gray) Wats, on the edge of a dry creek bed 14 miles E. of Tucson, Arizona, on April 12, 1965. One active nest was located on a small sandy strip near the center of the gravelly creek bottom. The uneven surface of the strip was sparsely covered with grass and about 10 percent of it was covered by driftwood and other flotsam. Nest Architecture entrance hole: The nest was located near the base of several converging grass plants but it was reasonably well exposed. The burrow entrance was open, faced west, and angled into the soil surface. A small bell-shaped en- largement surrounded the entrance to several millimeters below the surface but it was probably an abnormal structure caused by the collapse of the adjacent, loose, dry sand and its subsequent removal by the nesting bee. A well established asymmetrical tumulus, continually reshaped and en- larged by the nesting female, was present in front of the entrance hole. A shallow trail, 4 mm. wide, extended 3 cm. from the entrance, whereupon it made a 90° turn to the north and continued for an additional 8 mm. The moraine on either side of the trail was quite wide (8 mm.) and contained many pebbles and large sand particles. The trail and associated moraines of the tumulus were formed in the manner described for D. mulleri. burrows: The main difference between the burrow system of D. pulchricornis and that of other species of Du jourea described here was the subdivision of lateral burrows into sublaterals. Unfortunately, the only nest available for study was incomplete and portions of the architecture were lost during excavation. Nevertheless, architecture differed sufficiently to justify description here. The main burrow was unlined, unplugged, and 3 mm. in diameter. It main- tained about a 20° angle from horizontal for 10 mm., whereupon it made a subhorizontal spiral and proceeded vertically. It branched into two lateral burrows about 7 mm. below the surface, but one branch was soon lost. The remaining lateral was difficult to follow because it was partially plugged (possibly in the process of being completely plugged), but it eventually divided into a number of plugged sublaterals radiating short distances from the lateral burrow. Each sublateral terminated at a single cell. The four cells eventually uncovered were 9 cm. below the surface and positioned 3 to 4 mm. apart. cells: The cells were subspherical (5.75 mm. long and 5.0 mm. wide) and varied in position from subhorizontal to a 70-degree inclination from horizontal September, 1967] Torchio, et al.: Biology of Dufourea 143 with the posterior portion lower. As in the other Dujourea studied, the cells were carved from the substrate, but they lacked water-resistant walls as determined by the droplet test. The cell cap was composed of an unlined, tightly-packed soil plug 3.5 mm. wide and 3.0 mm. long. The inner face of the plug was concave and possessed three distinct rings surrounding a 1 mm. wide, central micropyle. The outer face of the cell plug could be distinguished from the plugged sublateral burrow only by its greater compaction. Provisioning and Development The pollen ball closely resembled that of D. mulleri in shape and diameter but differed in being yellow and somewhat drier. In subhorizontal cells, the pollen balls were positioned as were those of D. mulleri but they were at the bottom of the more vertical cells. Adult Activity The female whose nest we studied completed three pollen-carrying trips between 10:30 and 11:07 a.m., and began a fourth trip at 11:10 a.m. The speed with which she collected pollen and deposited it into a cell was remark- able, considering that the day was overcast and the air temperatures never rose above 70 °F. From the above data, it appeared that each pollen ball required at least four pollen loads for its completion. Approximately 100 pollen-collecting females were observed between 8:30 and 9:30 a.m., but an hour’s search throughout the area yielded but one nest. Consequently, D. pulchricornis was not gregarious under the conditions we encountered. Neopasites ( Micropasites ) cressoni Crawford Flight Activity This species of nomadine parasitic bee was encountered both at the Tubac site and at the Rodeo site. The females, more abundant than the males, flew low over the ground in a meandering fashion. They began flying as early in the day as their hosts and continued after the hosts ceased. Their flight, suggestive of that of Oreopasites and Holcopasites, was moderately slow and included frequent stops at apparent nest entrances of the host bee. Several times, two, three, four, or even five females hovered over a nest entrance, though such congregations occurred only where the cuckoo bees were most numerous. They often landed on flat, unshaded surfaces, probably to rest or sun themselves. Males were seen several times at the Rodeo site; their flight was higher and seemed somewhat faster than that of the females. Mating was not observed, but one of us (Torchio) observed it in N. ( Neo- pasites) fulviventris Cresson at Arroyo Seco, Monterey County, California, in 144 New York Entomological Society I Vol. LXXV 1959. The males utilized a small, bare, powdered surface area as a gregarious mating site. It was separated from the nesting area of its suspected host, Dufourea dentipes Bohart, by about 21 meters. Males patrolled the area or landed on it for long periods. Mating was observed in four instances, each occurring on the ground. Copulation lasted from 3 to 10 seconds. Oviposition Over 30 eggs and egg chorions of N. cressoni were encountered in cellsvof D. mulleri at the Rodeo site (by Rozen and Favreau), and all except one were deposited in the cell wall, as is the case with the other nomadine parasitic bees whose biology has been studied. Only once was an egg discovered embedded in the provisions; as there were already six eggs in the wall of this cell, we can only imagine that the female responsible for the seventh egg might have been at a loss to know what to do with it. It is not known how many eggs are normally deposited in a cell; usually one or two were discovered though as many as eight were found in a single cell. The last figure may be abnormally high, for the female Dufourea prob- ably left the cell open, thereby giving numerous Neopasites access to the cell. Frequently deep, rough scratches were observed in a cell which suggested that the host female, upon finding the Neopasites eggs in the cell wall, dug them out; Rozen has observed similar marks in the cell wall of N omadopsis in areas infested with Oreopasites. Eggs were deposited while the cells were being provisioned. The female Neopasites made a groove in the cell wall and inserted the egg, so that it rested with its exposed length flush with or a little higher than the cell wall (Fig. 7). Occasionally the egg was tilted at a slight angle so that one end projected farther than the other (Fig. 8). In all cases the lining of the cell abutted the egg, so that there was never a crack between the cell lining and the egg. This fact indicated that the female Neopasites cemented the crack with fine soil, and, in certain cases, some cement-like material adhered to the exposed part of the egg. The eggs seemed to be placed in almost any part of the cell though they were not ordinarily found near the entrance. The small eggs (Figs. 7-9) had an unusual appearance. About 0.6 mm. long, they were elongate, with the exposed surface being somewhat flattened, whereas the embedded part bowed out; they were thus rather boat-like in shape. The exposed chorion was stiff, thick, opaque white with faint, trans- verse corrugations. The chorion below the cell surface was thin, fragile, trans- parent and without ridges. At hatching, the exposed chorion ruptured (Fig. 9) in a semicircular to nearly circular line at the anterior end of the egg and the first-stage larva crawled out, leaving the chorion and attached door intact. Like the egg, the first-stage larva was very small, being considerably less than half the size of the Dufourea egg. The head was conspicuous, con- September, 1967 I Torchio, et al.: Biology of Dufourea 145 Table 1. The three subfamilies of Halictidae compared on the basis of known biological differences. T3 S-i o G bJj G G3 G CD d O 'in O J— i l+H X3 CD Td Q, cd cd o rG -H 1/1 G G ^ la O cn S-l S-i P-, IS a cd O Jh a & cd -(-i D !— i G _o "cn > O Sh a G O CD 2 s ■5 a G cd cn > o CD cd S ,CD T3 CD N CD _> u "G o c cn , the first and last of these the shortest. Fringe shining, light brown, lighter basally, especially subapically. Terminal line dark brown, very thin, widest subapically, but essentially complete to tornus. Hind wing light brownish, slightly darker terminally and apically, fringe whitish. Palpi, head and thorax brown, slightly lighter dorsally. Tegulae brassy. Frons rather flatly rounded. male genitalia (Fig. 4) : Tegumen narrow, tapering ventrad. Uncus slender, tubular, shorter than gnathos, with very short, fine setation. Gnathos broad basally, long, slender, tubular, distally gradually tapered. Vinculum large and broad, deeply emarginate cephalically. Pseudosaccus large, more than % as long as cephalocaudal width of vinculum, clavate. Costa of valva long, well sclerotized, with a long, slender, tapering, dorsad and mesad curving, free process that exceeds cucullus caudally. Cucullus broad basally, strongly taper- ing and curved dorsad, with abundant, fine, setation on mesal face. Sacculus of valva small and lightly sclerotized, with a short, flat, well sclerotized free process that curves mesad terminally. Aedeagus thick, blunt, shorter than valva, somewhat curved ventrad, a little more sclerotized ventro-distally ; with a small, slender, strongly curved basal cornutus and a very short, heavily sclerotized distal one that arises from a short, discoid base. female genitalia (Fig. 3) : Papillae anales bilobed, the ventral lobes well sclerotized and tapering internally to pronounced, but slender, apophyses posteriores. Eighth segment well sclerotized dorsally and laterally, but not ventrally, lacking apophyses anteriores. Extending caudad from ostium is a well sclerotized trough, complete laterally and ven- trally but open dorsally; from each dorso-caudal corner of this a sharp, slightly curved spine projects caudad. Within this trough is a shorter, rounded, heavily sclerotized, scobinate trough, also open dorsally; from the ventro-caudal edge of this extends a short tongue, concave dorsally and convex ventrally, bearing many very small spines. Entad from this the antrum is more lightly sclerotized, and then broadens to form a bulbous chamber with many complex vermiculations in its wall. Ductus bursae entad of this lightly sclero- tized and narrowing markedly to point of exit of ductus seminalis. A curved, very lightly sclerotized duct between ental end of ductus bursae (at ductus seminalis) and corpus bursae, suddenly enlarging in a short cervix bursae. Corpus bursae large, broadly ovoid, with two rather lightly sclerotized signa, each a group of small scobinations. Eighth abdominal segment undifferentiated. type material: Holotype S and allotype 9, near Dark Canyon, Guadalupe Mts., Eddy Co., New Mexico, July 23, 1959, leg. A. B. Klots. Paratype 3. b ' ■ ■ . t r \ , v IV 1 I: 7 1,’ (-> ) 7w 7a, Voi. lxxv 1 ■ i- i -r v ' • DECEMBER 1967 i&f r /, \ r -«W 1 1 X \ vV A - i'HB ws 777 1 A 77 < ■ '!■; ' ' ■ I J^r w , . ’> //r^ A •, 7 . / ,\ . I iIiH j \’ Vv A i j, \ n ( y Mi-rt ^ v, mk ,'5, •' -J l - , 1 j * \ ms.i'l v > yr 1 £ e N\. >'• , . yi »aj . 7f f ' : ; 7 || ■. A ,f J A A . ' V)7 7] ■ IW/ sm 5 , ;^T7V"' jAL A A! a ;:P * YA ! 7 M i i !>' 7 , 75 ('A/ I l \f 7 , J.. ..y VVj l 7 mm ; ;' M .' |\ '|,.| ,A 1/ 7 . ... MN,. w 7-7. a '-y 7 ( 1 ./ — 1 '/ 1 r- , v y; / ^ i ' » \ ! ' v ; i j ., _ v - v Devoted to Entomology in General iv oV ' l Vi*, t ' -jF Va ■' " 'il- J ' .7 . \» ,1 , M- - ^ ' ■ V i 7 •• /vi 1 ' ; ; .)/;■• : : 1 1| i ' || 4 -A u ' !\ ' I ! l> v? / y r, A. ' 7r - \ i, • Sv'.-lV'. J/'A\ 7 1 1 X- ..H7 1 b ) ,rr 1 1 V BS ,7 ('V, VV.H1? ' ' '■ , !' ' ] I ‘ . i. • ' f \ i H i v, _ ' V « .i 8 I#' • V ‘fri v ' i: r , M/'k , .U'i h AM A'':' i " ■ I / The -N>. New York Entomological Society Organized June 29, 1892 — -Incorporated February 25, 1893 Reincorporated February 17, 1943 vV _ i /, The meetings of the Society are held on the first and third Tuesday of each month (except June, July, August and September) at 8 p.m., in the American Museum of Natural History, 79th St., & Central Park W., New York 24, N. Y. Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00. Members of the Society will please remit their annual dues, payable in January, to the Treasurer. -\ ■( Officers for the Year 1967 President, Dr. Richard Fredrickson Vice-President, Dr. David Miller College of the City of New York 10031 College of the City of New York 10031 Secretary, Mr. Howard Topoff American Museum of Natural History, New York 10024 Assistant Secretary, Mr. Albert Poelzl 230 E. 78th Street, New York 10021 Treasurer, Mr. Raymond Brush American Museum of Natural History, New York 10024 - ..y . 1 / ,( ' , .. y .■ . •. \ " vyyy : . ~ y' ",y I /y / e Assistant Treasurer, Mrs. Patricia Vaurie i American Museum of Natural History, New York 10024 ft K jV-; Trustees ; v’ , r Class of 1967 Dr. Jerome Rozen, Jr. Class of 1968 M: Dr. Elsie Klots ii Mr. Robert Buckbee I' i t v V /C Mr. Bernard Heineman /• j- Mailed December 22, 1967 -K ! The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press Ipc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas. y y \ K Journal of the New York Entomological Society Volume LXXV December 22, 1967 No. 4 EDITORIAL BOARD Editor Emeritus Harry B. Weiss Editor Lucy W. Clausen College of Pharmaceutical Sciences, Columbia University 115 West 68th Street, N. Y. 10023 Associate Editor James Forbes Fordham University, N. Y. 10458 Publication Committee Dr. Kumar Krishna Dr. Asher Treat Dr. Pedro Wygodzinsky CONTENTS Apomyelois bistriatella : A Moth Which Feeds in an Ascomycete Fungus (Lepidoptera : Pyralidae) Jerry A. Powell 190 Melanism in New Jersey Catocala Schrank (Lepidoptera: Noctuidae) Joseph Muller 195 Biological Notes on Dioxys pomonae pomonae and on its Host, Osmia nigro- barbata (Hymenoptera : Megachilidae) Jerome G. Rozen, Jr. and Marjorie S. Favreau 197 A Revision of the Termitophilous Tribe Termitodiscini (Coleoptera: Staph- ylinidae) Part I. The Genus Termitodiscus Wasmann: its Systematics, Phylogeny, and Behavior David H. Kistner 204 The Immature Instars of the Cleptoparasitic Genus Dioxys (Hymenoptera: Megachilidae) Jerome G. Rozen, Jr. 236 Proceedings 249 Index to scientific names 251 Index to authors ... iii Apomyelois bistriatella: A Motli Which Feeds in an Ascomycete Fungus (Lepidoptera: Pyralidae) Jerry A. Powell1 University of California, Berkeley Abstract: A. bistriatella (Phycitinae) , a moth formerly recorded in the eastern United States, has been found to occupy diverse situations in California, feeding in the larval stage on stromata of Hypoxylon occidentale (Xylariaceae) . The moth was not recovered in extensive sampling of Polyporaceae, while records indicate that other species and perhaps other genera of Xylariaceae are used. During the past several years a large number of collections of wood-rot fungi from the western United States and Mexico have been processed for insect material. Early phases of the program were conducted primarily by J. F. Lawrence, now of the Museum of Comparative Zoology, Harvard Uni- versity, who surveyed primarily for Ciidae (Coleoptera). In the last three years an increasing emphasis has been placed on moths, the larvae of which inhabit these fungi. An analysis of host ranges of the Microlepidoptera (Oecophoridae, Oinophilidae, Tineidae) has been prepared (Lawrence and Powell, 1967). A summary of all productive fungus species involved in our collections is given in that paper. Although Polyporaceae (Basidiomyceteae) comprised about 90 per cent of the 480 lots processed, an assortment of other wood-rot fungi was included. Thus, several Thelephoraceae species including some 20 collections and a few lots of Agaricaceae, where these had developed somewhat hardened sporophores, were involved. All of these are members of the Basidieomyceteae, and the only other fungus involved was Hypoxylon occidentale Ellis and Ever- hart,2 ( Ascomyceteae: Xylariaceae ) . The sporophores, or stromata, of this spe- cies are carbonous appearing, hemispherical, about 2 to 4 cm in diameter (Plate I) and are commonly seen on recently killed Quercus agrifolia through- out the coastal foothills of California. Although several species of Tineidae and Oecophoridae were reared from this fungus, it was concluded that it is only an incidental host because the stromata are hard and dry during several months each year. None of the ciidae use H. occidentale. 1 Research conducted in part as a by-product of National Science Foundation grant project GB-4014. L> Hypoxylon occidentale has been treated as a synonym of H. thouarsianum (Lev.), a widespread Neotropical and Nearctic species described from the Galapagos Islands (Miller, 1961). For the present discussion the name occidentale will be used for the California-Oregon segregate. 190 December, 19671 Powell: Fungus-Feeding Moth 191 Upper: Apomyelois bistriatella (Hulst), female (left) and male (right) from Lone Pine, California, reared from Hypoxylon. Actual size: $ 23.5 mm, $ 21.0 mm, wing expanse. Lower: Stromata of Hypoxylon occidentale (Ascomvceteae, Xylariaceae) on bark of Quercus agrifolia from Berkeley, California, showing frass exudations due to feeding of the moth larvae. The 2 cm scale applies to both lower photos. In the fall of 1961 a collection of Hypoxylon occidentale produced two adults of a large phycitid moth. It was assumed that these individuals had only an incidential association with Hypoxylon , perhaps using it as a scavenger or for a pupation site, since Heinrich (1956) lists no American Phycitinae as fungus feeders. However, subsequent collections of this moth, Apomyelois bistriatella (Hulst), indicate that Hypoxylon is a normal host for the larvae. Moreover, Apomyelois was not encountered in any other of the wood-rot fungi which we processed, indicating that the moth is specific to Hypoxylon. Apomyelois bistriatella (Hulst) Dioryctria bistriatella Hulst, 1887, Ent. Americana, 3:136. Apomyelois bistriatella ; Heinrich, 1956, U. S. Natl. Mus., Bull. 207:43. (tax- onomy) . The genus Apomyelois was proposed by Heinrich (1956) to accommodate the single, widespread but poorly known species, bistriatella Hulst, originally described from Washington, D. C. Heinrich had material of the species repre- 192 New York Entomological Society [Vol. LXXV senting several widely scattered stations in the eastern United States and Canada. There was no information on the biology of this moth. Records in the California Insect Survey, University of California, Berkeley, show this species to be widespread ecologically and geographically on the West Coast. adult: The moths are rather large, relative to many Phycitinae, having a wing- spread of 20 to 24 mm. The forewing is dark gray, dusted with whitish, espe- cially on the costal half, and is crossed by two white lines, one at the basal one- third, and a less distinct, somewhat sinuate one beyond the end of the cell (Plate I). Western specimens compare well with Heinrich’s characterization of the species, both in external features and in genitalia form of both sexes. A pair of 60-year-old specimens from Ottawa, Canada, and Massachusetts, sent to me from the U. S. National Museum are paler and have less well defined markings, especially in the terminal area of the forewing. However, these dif- ferences probably are a function of the age of the eastern specimens. The eastern male has a more deeply cleft gnathos (possibly the slide upon which Heinrich’s figure was based) than California examples (four preparations examined). If any of these differences are to be considered sufficient to warrant proposal of a nomenclature designation of the west coast race, this will have to be shown through comparison with typical material in series. The series from Inyo County, California, shows considerable variation in wing color and in size. biology: The life history of this insect is not clearly defined, and it may vary with climatic condition. In eastern areas flight records are available for May, June and July in the north and for March in Florida. Records of field collected adults in the California Insect Survey suggest that the species is multivoltine, the flight perhaps varying with weather conditions and growth of the host. In coastal areas of California the moths have been taken in late April, July, September, and October, and the larvae in May and October producing adults in June and November. At 3500 feet elevation in the Sierra Nevada adults have been collected in June and August. Stromata of Hypoxylon appear in fall after the first rains and grow then and during winter. At this time they are relatively soft, having a consistency similar to damp charcoal, and can be crushed between one’s fingers. Even in late spring, well after winter rains have ceased, visible moisture can be squeezed from sporophores situated in damp areas. During the dry season, however, the stromata harden and desiccate. In summer at most localities where Quercus agri folia serves as a host the hemispherical sporophores are so hard they usually can neither be dislodged nor crushed by hand. Nonetheless, the entostroma is somewhat softer in texture and it appears that at least the larger larvae are able to feed at nearly any time of the year. Neither eggs nor young larvae have been observed. Larger larvae fed in irregular galleries, usually beneath the thin, crust-like, perithecia-bearing sur- December, 19671 Powell: Fungus-Feeding Moth 193 face layer. Often the galleries were somewhat blotch-like, not extending through the whole depth of the entostroma. At times side tunnels radiated outward or more deeply towards the substrate. No evidence of a direct opening to the exterior was noted, and the burrows became filled with frass. The frass some- times extrudes irregularly from the surface of the stromata (Plate I). In the field, the thin surface layer often later collapses or is broken away by external agencies, resulting in a characteristic shallow hollowed out area around the apex of the dome of the stroma. I have noted these evidences of larval feeding at a number of California stations in addition to those from which the moths were reared. No larvae were found to burrow into the bark subtending the Hypoxylon, although they may sometimes wander under normal conditions and seek out crevices, insect burrows, etc. for pupation. In the laboratory pupation usually took place in the burrows, either just under the thin, ectostromal layer and parallel to it, or occasionally in a deeper gallery, perpendicular to the surface. Some individuals formed the loose silken cocoons amongst debris in the rear- ing container, between Hypoxylon pieces, etc. One individual pupated in an abandoned cerambycid gallery some 4 cm from the emergence hole of the beetle. This exit was also successfully used by the moth upon emergence. geographical distribution: Available records show a disjunct range, in eastern North America from Ontario and Wisconsin to the District of Columbia and Iowa, in Florida (Heinrich, 1956) and in California. The diverse ecological situations occupied by the species in California are not representative of austral or boreal distributional patterns typical of many insects. Probably Apomyelois bistriatella occurs over much of temperate North America at in- termediate elevations. Specimens of Hypoxylon collected from Populus at Lone Pine were not submitted for identification, having been assumed to be H. occidentale. For H. thouarsianum , including occidentale , however, Miller (1961) states that Celtis , Piersea and Quercus are known hosts. Thus it may be that the Inyo County fungus was a different species. The range of H. thouarsianum in the eastern United States does not extend north of North Carolina (Miller, 1961), indicating that at least one additional host is involved. Hypoxylon species with relatively bulky stromata (as opposed to species with little or no development of entostromal tissue) may be generally used. In addition, I have seen herbarium specimens of Daldinea , a related genus of Xylariaceae, with evidences of lepidopterous feeding, suggesting the possibility that Apomyelois uses ascomycetes other than Hypoxylon. California material examined: Contra Costa Co.: Pleasant Hill, 1 2 IX- 15-58 (W. E. Ferguson); Orinda, 1 $ X-ll-61, at 15 watt blacklight (P. A. Opler) ; Walnut Creek, 1 $ VIII-5-65, 1 2 VIII-23-66 (J. Powell). Inyo Co.: 5 mi. W. Lone Pine 25 S S , 33 2 2 VI-13-65, r. f. Hypoxylon on poplar, emgd. 194 New York Entomological Society [Vol. LXXV VII-6 to VIII-1-65 (J. T. Doyen Collr; JAP 65 G5). Marin Co.: 1 mi. SE Inverness, 15,1$ X-8-61, r. f. Hypoxylon occidentale on Quercus agrifolia, emgd. XI-7 and XI-20-61 (C. W. O’Brien collr.; JFL 979); Inverness, 1 5 IX-8-62, at light (C. A. Toschi). Santa Barbara Co.: Prisoner’s Harbor, Santa Cruz Island, 2 $ $ V-l-66, r. f. Hypoxylon occidentale on Quercus agrifolia, emgd. VI-7 and VI-13-66 (J. Powell, A. Slater, J. Wolf collrs.; JAP 66E4); Central Valley, Santa Cruz Is., 1 5 IV-28-66, at light (J. Powell). Sonoma Co.: Hacienda, 1 $ VII-9-61 (C. Slobodchikoff). Tuolumne Co.: Twain Harte, 1 $ VI-19-59, 1 $ VIII-18-60 (M. Lundgren). Specimens are deposited in the collections of the California Insect Survey and U. S. National Museum. Acknowledgments: Thanks are extended J. F. Lawrence, Museum of Comparative Zoology, who provided some of the early data for this study, and to J. T. Doyen and C. W. O’Brien, University of California, Berkeley for field collections. Further field and laboratory observations were made by P. A. Rude, A. J. Slater, and J. Wolf, assistants with the National Science Foundation project (GB-4014) which supported part of the study. Identifications of the Hypoxylon were provided by I. I. Tavares, University of California, Berkeley, Herbarium. Specimens of the moth were examined by W. D. Duck- worth, U. S. National Museum, Washington, D. C., and acknowledgment is also made for use of comparative material which was sent from that institution. Literature Cited Heinrich, C. 1956. American moths of the subfamily Phycitinae. U. S. Natl. Mus., Bull. 207 : 581 pp. Lawrence, J. F. and J. A. Powell. 1967. Host relationships in North American fungus feeding moths (Oecophoridae, Oinophilidae, Tineidae). Bull. Mus. Comp. Zook, Harvard, in press. Miller, J. H. 1961. A monograph of the world species of Hypoxylon. LTniv. Georgia Press, Athens; xii + U8 pp. Received for Publication June 5, 1967 An Information Desk for Scientists and Technologists The Library of Congress with the support of the National Science Founda- tion has set up a National Referral Center for Science & Technology (Library of Congress, Washington, D.C. 20540). The Referral Center is designed to pro- vide a single place for advice on where and how to obtain information of any facet of the physical, biological, social, or engineering sciences. It serves as the intermediary to direct persons or organizations seeking information on specific topics to those who can furnish the information, and there is no charge. The Center does not provide technical details nor bibliographic assistance. Melanism in New Jersey Catocala Schrank (Lepidoptera, Noctuidae) Joseph Muller Lebanon, New Jersey Abstract: Brief discussions, counts, and descriptions are given of reared melanic forms of Catocala micronympha Guen. and C. minuta W. H. Edwards. In the Spring of 1966, seventy-five Catocala micronympha were reared from eggs laid by 4 females obtained in July, 1965 at black lights in Lebanon, New Jersey. The female parents were all more or less brownish black or melanic. All but 6 of these reared individuals were more or less melanic. They may be characterized best by comparison with the 9 figures of C. micronympha given by Barnes and McDunnough (1918, Mem. Amer. Mus. Nat. Hist., n. ser., Vol. Ill, part 1, PL 9, figs. 22-30). Of the specimens figured there, one is a brownish black form, gisela Meyer, and the other 8 are brown and grey with a complete absence of black. Among the reared specimens 6 are of the gisela form, and several are like gisela but have the white sordid rather than clear. Twelve specimens resemble the form hero Henry Edwards, but have the wing bases greyish rather than brownish and the apices of the fore wings black rather than brown. Forty-two specimens are all black with only a faint whitish sub- terminal line; and the remainder of the specimens are more or less evenly grey- black. Dr. A. E. Brower of Augusta, Maine has commented (in litt.) that gisela is a genetic form of C. micronympha known long before any appreciable melanism appeared in the genus, and that now we have melanic specimens of gisela. Thirty-one specimens of C. minuta W. H. Edwards were reared from pupae, also collected in Lebanon. Compared with the forms figured by Barnes and McDunnough (loc. cit. figs. 1-6) 20 specimens resemble fig. 5, which is mostly dark brown, but are darker and show no brown; and 11 specimens resemble f. parvula W. H. Edwards (fig. 2) but have the brown replaced by blackish grey, and the inner margin black. In 1960 I described the melanic f. broweri of C. connubialis pulverulent a Brower from Lebanon (Jour. Lepid. Soc., 14: 177). Until 2 years ago this was the commonest form at Lebanon. Since then, however, both broweri and the nominate form, pulverulenta , have almost disappeared from this area, being replaced by C. micronympha which was first seen here about five years ago. This region of New Jersey Hunterdon County, is mostly farm land with hilly areas of deciduous woods. Industries are 30-60 miles distant. Many melanic forms of various species of Lepidoptera, especially of Catocala , have been taken here, which indicates that air pollution extends this far. However, melanic forms of Catocala are also numerous in northern New Jersey where 195 196 New York Entomological Society [Vol. LXXV there are mountains and continuous, dense woods mixed with huge hemlocks and pines. In the latter case it is thought that the Lepidoptera have become adapted to their surroundings, and that industrial air pollution does not extend this far. On the other hand, no melanic forms of Catocala have been collected in the dense pine woods of the pine barrens of southern New Jersey, although many nights have been spent there sugar-baiting and light-collecting. During the last 6 seasons many different ways have been tried to mate the normal and melanic forms. Most of the females mated do lay eggs, but these have always collapsed and failed to hatch. Received for Publication December 16, 1966 Insect Attractants Two acrylic auto paints have been reported to be effective attractants for sap beetles in Science, 156: 946-947 (May 19, 1967). Biological Notes on Dioxys pomonae pomonae and on its Host, Osmia nigrobarbata ( Hymenoptera : Megachilidae ) Jerome G. Rozen, Jr.1 and Marjorie S. Favreau1 Abstract: Biological observations on the parasitic bee Dioxys pomonae pomonae Cockerell are presented covering the following points: searching habits of female, oviposition, elimina- tion of immatures of the host, feeding habits, and cocoon. Additional observations, including nest structure, are given for the host bee Osmia nigrobarbata Cockerell. With the exception of a paper by Micheli (1936), apparently nothing was known heretofore concerning the biology of the cleptoparasitic bee genus Dioxys beyond the host associations of some of the species (Hurd, 1958; Jaycox, 1966). For this reason, we present the following observations concerning Dioxys pomonae pomonae Cockerell, a North American representative of this dis- tinctive Holarctic genus. Brief notes are also given on the biology of the host bee, Osmia ( Acanthosmioides ) nigrobarbata Cockerell. An accompanying paper (Rozen, 1967) describes the immature stages of D. pomonae pomonae. We would like to thank the following people for identifications of adults associated with this study: Dr. Paul D. Hurd, Jr., University of California, Berkeley; Dr. Elbert R. Jaycox, University of Illinois, Urbana; and Dr. Charles D. Michener, the University of Kansas, Lawrence. The literature search was aided by the Bibliography of Apoid Biology under Dr. Michener’s supervision. This study was carried out at the Southwestern Research Station of The Ameri- can Museum of Natural History, Portal, Arizona. description of nesting area: All observations were made at 3 miles north of Apache, Cochise County, Arizona, between April 28 and May 5, 1966. The Osmia burrows were widely scattered over nearly horizontal ground sparsely covered by low vegetation consisting of Malacothrix , Gaillardia, Phacelia , a number of grasses, and other low-growing plants (Fig. 1). Several possible hosts of Dioxys were active including Osmia ( Acanthosmioides ) nigrobarbata Cockerell (determined C. D. Michener) and Anthidium emarginatum (Say) (determined E. R. Jaycox). Both the Osmia and Anthidium collected pollen from Astragalus. Three species of Dioxys flew in the area: D. productus subruber (Cockerell), D. pomonae pomonae Cockerell, and D. pacificus paci- jicus Cockerel] (all identified by P. D. Hurd). Females of D. pomonae pomonae were seen both entering and waiting by the burrows of Osmia nigrobarbata , and a female was reared from an Osmia cell. The hosts of the other species are not known. 1 Dept. Ent., Amer. Mus. Nat. Hist. 197 198 New York Entomological Society [Vol. LXXV Fig. 1. Nesting area of Osmia nigrobarbata Cockerell. observations on the biology of Osmia nigrobarbata: Nests of this species were widely scattered and entrances were usually found at the bases of low plants or at the edge of shallow depressions. The burrows entered the ground at a slight angle from the horizontal and each tumulus was piled on one side of the entrance. Burrows were open and their direction was unpredictable, for some turned sharply to the side or downward. They were short, measuring only a few inches long, and the cells were situated within two or three inches of the surface. Some cells were encountered barely below the loose, dry surface layer of soil. The nearly horizontal cells are constructed from a mastic of plant tissue. The source of this material is unknown, but because it was uniform for all cells encountered, it must be gathered from a particular plant. At first bright green, its color fades, so that cells several months old are nearly brown. The cell wall, approximately 0. 5-1.0 mm thick, is quite hard; the inside cell dimensions are approximately 8.0 mm long and 5.0 mm in maximum diameter. The cell closure consists of the same plant material as that of the wall and is nearly flat on the inside and concave on the outside. The arrangement of the cells is extremely variable. Some single cells were found which were probably the beginning of a nest series; the other cells were December, 1967] Rozen and Favreau: Parasite Bee (Dioxys) Biology 199 Fig. 2. Nests of Osmia nigrobarbata Cockerell. Swellings represent individual cells. Fig. 3. Opened cell of Osmia nigrobarbata Cockerell showing food loaf and egg, from side. arranged in a basically linear series that branched in an infinite number of ways (Fig. 2). Cells in the series were all interconnected so that four or five cells could often be removed from the ground without their separating. Each cell was a complete unit in that the rear end (or side) of one cell was not the 200 New York Entomological Society [Vol, LXXV Figs. 4, 5. Cell of Osmia nigrobarbata Cockerell, with front end removed. 4. Freshly deposited egg of Dioxys pomonae pomonae Cockerell adhering to anterior end of the Osmia egg. 5. Same cell, viewed from above, just before Dioxys egg hatched. Notice chorion adhering closely to the Dioxys embryo. December, 1967] Rozen and Favreau: Parasite Bee (Dioxys) Biology 201 cap of the previous cell. Hence in a series, individual cells could be broken off without any of the cells being damaged. The strings of cells lay approximately horizontally in the ground. Only one female was responsible for a cell series, and each cell was constructed, provisioned, and closed before the next one was started. Provisions of nectar and pollen were formed into a large, elongate, moist loaf (Fig. 3) occupying most of the cell. All eggs were uniformly placed on top of the provisions, forward of the center, in the sagittal plane of the cell. The eggs were laid either on the surface of provisions or with the rear of the egg slightly embedded. The anterior end rested on or, perhaps more frequently, projected into the lumen of the cell and pointed toward the cell closure. The mature larva of Osmia spins a well-developed cocoon which consists of a loosely woven, tan outer layer and a tough (leathery), polished (on the inner surface) inner layer that is almost black. The cocoon lacks a nipple at the anterior end. biological notes on Dioxys : The females of D. pomonae pomonae and pacijicus pacificus fly slowly close to the surface of the ground and stop briefly at spots that presumably have certain characteristics of the nest entrances of the hosts. The flight appears “deliberate” and unhurried. Occasionally a female suddenly flies swiftly a short distance and then again starts her slow searching. Although the path meanders, it tends to lead in one direction, so that the female travels a considerable distance. As the Osmia nests were widely scattered over a number of acres, this behavior pattern of D. pomonae pomonae appears to be functional. In contrast, the meanderings of such parasitic bees as Oreo- pasites, Holcopasites, and Neopasites carry the bee back and forth over a limited area; this restricted search pattern appears to be an adaptation to the gregarious nesting habits of host species. Now and then, the Dioxys females land on the ground and clean their wings and antennae as do females of the nomadine genera. Once, after finding a burrow of Osmia , a female of D. pomonae pomonae examined the entrance, then retreated a few inches, and sat on a twig where it waited, as if for the departure of the host female. Several other times a female was noticed entering an Osmia burrow but came out within a half a minute. Over 470 cells of Osmia were opened during our search for the immatures of Dioxys , with the result that we found seven larvae and two eggs of the para- site. One egg (Fig. 4) adhered loosely to the anterior end of the host egg. A small slit in the cell wall above the posterior end of the Dioxys egg apparently marked the spot through which the egg was inserted into the sealed cell. The other egg was partly embedded lengthwise in the under surface of the pollen- nectar mass so that somewhat more than half of it was visible. The chorion is shiny and translucent white. Resembling the host egg in almost all respects, the egg of Dioxys is somewhat smaller: length, 1.5-1. 8 mm, width, 0.6 mm. 202 New York Entomological Society [Vol. LXXV Figs. 6, 7. Cells of Osmia nigrobarbata Cockerell. 6. Same cell as in Figs. 4 and S. First instar Dioxys with its large head next to Osmia egg, which has been recently killed. 7. Intermediate stage larva of Dioxys pomonae pomonae Cockerell. December, 1967] Rozen and Favreau: Parasite Bee (Dioxys) Biology 203 Each of the Dioxys larvae was found in a cell with a dead egg, first instar, or second instar of Osmia. The host is killed with the sharp mandibles which are present during the first three larval stages (Rozen, 1967). One first instar larva was discovered on the underside of the pollen-nectar loaf, whereas the other larvae, presumably second instars, rested on the side or top part of the food. The egg found adhering to the Osmia egg hatched in the laboratory, and the first instar immediately killed the host egg (Figs. 4-6). However, at least the first and second instars were active and, if touched with forceps, opened their jaws widely and actively moved the anterior part of their bodies from side to side. These actions, plus the large, sharp-pointed mandibles of the first three instars, suggest that the host may be eliminated by the second or third instar as well as the first. Never more than one Dioxys was found in a cell; the female Dioxys probably deposits only a single egg in a nest. In contrast, females of many of the Nomadinae lay more than one egg per cell. As with most other bee larvae, the duration of the feeding period is short, lasting for two to three weeks. The larva, while feeding, moves about on the provisions (Fig. 7). Four larval instars were observed (but see Rozen, 1967). The fourth instar begins to defecate before it finishes feeding; the feces are extruded as elongate semisolid pellets. The thin outer layer of the cocoon is composed of very loose strands of silk to which some of the fecal pellets adhere. Defecation is completed before the next layer is deposited. The second layer, black in color, is comparable to the inner, leathery layer of the Osmia cocoon but is thicker and imparts a greater rigidity to the finished case. The innermost face of the one Dioxys cocoon examined consisted of yet another layer, at least toward the anterior end of the cocoon. Loose and light brown, it formed a cellophane-like coating even though some individual silk strands were detected. Except toward the rear where the inner layer adhered more or less closely to the rigid layer, the inner face did not possess the polished, nearly black surface of the Osmia cocoon. The cocoon of D. pomonae pomonae possessed a distinct nipple at the anterior end, so that the shape of the cocoon was identical to that of Dioxys cincta (Jurine) (Micheli, 1936, Fig. 6). Literature Cited Hurd, P. D., Jr. 1958. American bees of the genus Dioxys Lepeletier and Serville (Hymenoptera: Megachilidae) . Univ. California Publ. Ent., 14: 275-302. Jaycox, E. R. 1966. Observations on Dioxys productus productus (Cresson) as a para- site of Anthidium utahense Swenk (Hymenoptera: Megachilidae). Pan-Pacific Ent., 42: 18-20. Micheli, L. 1936. Note biologiche e morfologiche sugli imenotteri (VI Serie). Atti Soc. Italiana Sci. Nat. e Mus. Civ. Stor. Nat., 75: 5-16. Rozen, J. G., Jr. 1967. The immature instars of the cleptoparasitic genus Dioxys (Hymenoptera: Megachilidae). Jour. New York Ent. Soc., LXXV(4): 236-248. Received for Publication June 13, 1967 A Revision of the Termitophilous Tribe Termitodiscini ( Coleoptera : Staphylinidae ) Part I. The Genus Termitodiscus Wasmann ; its Systematics, Phylogeny, and Behavior1 David H. Kistner Department of Biological Sciences Chico State College Chico, California 95926 Abstract: The genus Termitodiscus Wasmann is redescribed, illustrated, and a key differ- entiating this genus from the other two genera of the tribe is provided. All of the previously described species of the genus are redescribed and new characters illustrated. Six new species are herein described, T. eoatoni from South Africa, T. emersoni from the Congo Republic, T. krishnai from Burma, T. latericius from South Africa, T. sheasbyi from Southwest Africa and T. vansomereni from Kenya. Distribution maps are presented which show the distribution of all species. Diagrams are presented showing the relationships among the species using both the phylogenetic and the phenetic approach. A summary of the host relationships is presented showing 100% host specificity to species of Odontotermes of the species now known. Observations on the behavior and distribution of selected species within the nests are presented which support the interpretation of the species as integrated termite guests whose principal adaptation to life within the nest is that of avoidance. The relation- ship of the tribe Termitodiscini with the Mvrmedoniini is documented and discussed. INTRODUCTION AND TAXONOMIC HISTORY The termitophilous tribe Termitodiscini was reorganized as a tribe of the subfamily Aleocharinae by Seevers (1957) to contain the genera Termitodiscus Wasmann, T ermitogerrus Bernhauer, and Discoxenus Wasmann. Prior to Seevers’ revision, the group had been recognized as a separate subfamily of the Staph- ylinidae. I here concur with Seevers’ judgment that there is no character or group of characters which could separate them absolutely as a subfamily distinct from the Aleocharinae. I shall show that the group probably arose from some free- living or loosely integrated termitophile of the aleocharine tribe Myrmedoniini. Seevers did not attempt to revise the species due to the paucity of material avail- able. Since that time, a lot of new material has been collected due to the field efforts of Dr. William Coaton and his colleagues of the Plant Protection Research Institute, Pretoria; Dr. Alfred E. Emerson, University of Chicago; Dr. Kumar Krishna, American Museum of Natural History, New York; Dr. A. de Barros Machado and his colleagues, Museu do Dundo, and myself. Most of the new ma- terial belongs to the genus Termitodiscus, so that this revision is confined to that 1 This study was financed in part by the National Science Foundation (Grant GB-3396). Some of the data reported herein were collected during the tenure of a post doctoral fellow- ship of the John Simon Guggenheim Foundation. 204 December, 1967] Kistner: Termitophile Revision 205 genus and revision of the other two genera will be deferred until a reasonable amount of new material becomes available. The careful study of new material has revealed characteristics which make it necessary that the key to the genera provided by Seevers be revised and this is done here. While collecting Termi- todiscus in the field, various observations were made on their behavior, par- ticularly in relation to their termite hosts, which bear on the integration of the termitophiles into the termite colonies. These observations and their interpre- tation are presented in this paper. The remainder is organized into the follow- ing sections: (1) Methods and materials; (2) Key to the genera of the tribe; (3) Redescription of the genus; (4) Key to species; (5) Descriptions of the species; (6) Relationships of the species; (7) Behavorial observations; (8) Host specificity; (9) Relationship of the tribe to the aleocharine tribe Myrme- doniini; (10) Acknowledgments; (11) Literature cited. METHODS AND MATERIALS Most of the routine methods used in my laboratory have been described several times, most recently by Koblick and Kistner, 1965, and Kistner, 1966. The only major change has been the substitution of a Nikon F camera with 55 mm, 50 mm, 35 mm, and 28 mm lenses plus bellows and extension tubes for the Exacta equipment used in the past. For ultra close-up photos of minute insects, this has proven superior because the corners are not chopped off the pictures and the lenses are easier to reverse to eliminate spherical aberration. The special techniques involved in the computer analysis of the relationships between the species are discussed later. The programs themselves are not included as most laboratories have developed their own and our programs are changed just about every time we use them. Current print-outs in Fortran II will be sent to anyone requesting them. The field techniques used vary according to the way in which the Odonto- termes hosts make their nests. Some species such as Odontotermes taprobanes Walker and Odontotermes culturarum Sjoestedt make well defined nests of which the bulk is located above the level of the surrounding ground. The queens are usually located at or near the ground level with the fungus gardens arranged in semispherical layers above the royal cell. The fungus gardens immediately above the royal cell usually yield the most specimens of Termito discus, but the other fungus combs may yield T ermitodiscus or other species of associated insects. We try to keep the layers separate as we dig in, but individual idio- syncracies of the nests prevent absolute accuracy. The fungus gardens are removed and taken back to the laboratory or other dwelling where the fungus is carefully pulverized over a yellow plastic tray. The yellow contrasts well with the termites and the termitophiles and permits the investigator to see the termitophiles and to aspirate them up or to pick them up with a camel’s hair brush. It takes about 4 to 5 times as long to sort through the fungus gardens of 206 New York Entomological Society [Vol. LXXV . IB ■ Figs. 1-2. Overall appearance of dorsal surface of beetle: 1. T ermito discus braunsi Was- mann; 2. T. escherichi Wasmann, Cotype. Scale arbitrary, see descriptions for measurements. a productive nest than to dig it up, so it pays to at least assay the fungus in the field before taking it back. Unless collecting is extremely poor in general, I usually abandon a nest if I don’t see at least a few specimens of termitophiles during the field assay. Other species of termites such as Odontotermes montanus Harris or Odonto- termes transvaalensis Sjoestedt build nests which are completely or almost completely submerged under the ground, often with little evidence on the surface of their position. Working with such nests can be extremely productive but is often extremely frustrating because a sizable investment of time and labor has to be made before one can tell if there are any termitophiles there or not, or even if the nest is there or not. The procedure we used and which is also used by Dr. Coaton and his colleagues is to dig a trench about 4 feet wide, 6 feet long and 4 feet deep to the side of where you think the nest is. Then dig in toward the nest from the side until you (hopefully) hit it. If you dig in from the top, you eventually fall into the nest which complicates the sorting process and partially destroys the ecological data. After you reach the fungus gardens, the fungus is gathered and sorted as above. I might add that I have dug until I could not throw the dirt out of the hole over my head and still not reached the nest, so I usually keep an open mind about abandoning a hole if nothing shows up quickly. A gung-ho attitude of, “I’m going to find that nest if December, 1967] Kistner: Termitophile Revision 207 iK'n Figs. 3-9. Antennae and mouthparts: Termitodiscus escherichi Wasmann: 3. 10-segmented antenna; 8. Maxilla; 9. Labrum and mentum. T. angolae Seevers: 6. Mandible. T. machadoi Seevers: 4. 9-segmented antenna; S. Labrum; 7. Mandible. Scale arbitrary, photos were taken at 100 X magnification. it kills me,” (my original attitude) just will not make economic sense in the long run. It is thus more productive to abandon a potentially dry hole while the investment in time and labor is still minimal and put that time and labor into another potential nest. The judgment necessary to make that decision came hard for us and is still based on so many subjective factors that finding the nests and then the termitophiles is still in the realm of art rather than science. KEY TO THE GENERA OF THE TRIBE 1. Mesocoxae widely separated; antennae 9, 10, or 11 segmented, short, very slightly visible from above; antennae segments other than 1 and 2 compressed and in- crassate 2 208 New York Entomological Society [Vol. LXXV Figs. 10-13. Termitodiscus escherichi Wasmann: 10. Prosternum and mesothoracic peri- tremes; 11. Abdominal segment VIII; 12. Abdominal segment IX and spermatheca. T. transvaalensis Silvestri: 13. Abdominal segment IX and spermatheca. Scale arbitrary, photos taken at 100 X magnification. Mesocoxae narrowly separated; antennae 11 -segmented, elongated, easily visible from above; antennal segments 3-11 with the sides meeting each other and covering the petiolar connections Discoxenus Wasmann 2. Antennae 9 or 10-segmented ; antero-lateral margin of pronotum slightly flared; mesosternum slightly declivous in middle Termitodiscus Wasmann Antenna 11-segmented ; antero-lateral margin of pronotum not flared; mesosternum almost vertical at the middle and thus scarcely visible from below Termitogerrus Bernhauer December, 19671 Kistner: Termitophile Revision 209 Figs. 14-16. Legs of Termito discus transvaalensis Silvestri; 14. Proleg; IS. Mesoleg; 16. Metaleg. Scale arbitrary but equal for all legs; photos taken at 100 X magnification. note: Termito germs seems to be confined to Central and West Africa as careful searches of Macrotermes nests in South Africa and the Orient have not revealed this genus so far. Discoxenus has only shown up in Odontotermes nests from the Orient in spite of careful searches of Odontotermes nests in Africa. The revision of these two genera will be delayed until there are far more new specimens available for study. 210 New York Entomological Society [Vol. LXXV REDESCRIPTION OF THE GENUS Genus Termitodiscus Wasmann Termitodiscus Wasmann 1899, Deutsch. Entomol. Zeitschr., 1899: 147; 1912, Zeitschr. Wissensch. Zool., 101: 92; 1916, Zool. Jahrb. System., 39: 179; Cameron, 1932, Fauna of Brit. India, Staph., 3: 317; Silvestri, 1947, Arch. Zool. Ital., 31: 125; Seevers, 1957, Fieldiana Zool., 40: 259; 1965, Publ. cult. Companh. Diam. Angola, 69: 129. Type species: Termito- discus heimi Wasmann ( Blackwelder, 1952: 377). Overall body shape limuloid, broad and flat as in figs. 1 and 2. Head broad and short, sub- triangular in form with the foramen magnum totally ventral in position. Eyes present, well developed and forward and laterally directed. Antennae inserted between the eyes with grooves developed on the genae for the reception of the large basal antennal segments. Submentum and gula extremely short. Antennae 9 or 10-segmented, shaped as in fig. 3 and 4. Labrum short, shaped as in fig. 5. Mandibular shape somewhat variable by species but the form is relatively constant, two extremes shown in figs. 6 and 7, note the one central and one apical tooth with the short stubby prostheca (barely visible in the photographs below the central tooth). Maxillae shaped as in fig. 8, palpi 4-segmented. Labium and mentum extremely small, shaped as in fig. 9, palpi 3-segmented. Pronotum semi-circular in shape (figs. 1 and 2) such that there is no distinction between anterior and lateral margins which are henceforth referred to as the anterolateral margins. Prosternum small, carinate in the middle, shaped as in fig. 10. Mesothoracic peritremes reduced in size but present and shaped as in fig. 10. Mesosternum and metasternum both short, metasternum somewhat shorter than the mesosternum. Mesothoracic coxal cavities relatively widely separated by a smooth mesothoracic and metathoracic process. Leg axis short compared to the width of the body. Proleg shaped as in fig. 14, with a large coxa but without flanges on the femur to accept the tibia in repose. Mesoleg shaped as in fig. 15, without femoral flanges. Metaleg shaped as in fig. 16, without well developed femoral flanges. Tarsal formula 4-5-5. Abdomen flattened, overall shape tapering gradually from segment III to segment IX. Apparent differences as in figs. 1 and 2 due to relative telescoping of segments. Segment II represented by a very reduced tergite only. Segments III-VII entire with 2 pairs of para- tergites each. Segment VIII represented by the tergite and sternite only which may or may not be pointed as a secondary sexual character, shaped as in fig. 11. Abdominal segment IX trilobed with 2 lateral portions and split median portion, shaped in the female as in figs. 12 and 13. The male has longer asymmetrical projections from the anterior border of the venter. Median lobe of the male genitalia variable by species. Lateral lobe of the male genitalia somewhat variable by species but always of the same general form as in figs. 17 and 18. KEY TO SPECIES OF TERMITODISCUS 1. Pronotum with an even covering of setae 2 Pronotum without setae or with at most a single row along the posterior border 7 2. Antennae with 9 segments 3 Antennae with 10 segments 4 3. Male genitalia shaped as in fig. 27, with a median spine sheasbyi n. sp. Male genitalia shaped as in fig. 23, without a median spine machadoi Seevers 4. Size very small, pronotum length 0.33-0.38 mm 5 Size larger, pronotum length 0.47-0.55 mm 6 5. Pronotal setae rather sparse, male genitalia shaped as in fig. 24 .... krishnai n. sp. Pronotal setae dense, male genitalia unknown minutus Cameron December, 1967] Kistner: Termitophile Revision 211 6. Male genitalia shaped as in fig. 25, with a lateral ventral spine on each side heimi Wasmann Male genitalia shaped as in fig. 22, without latral ventral spines on each side escherichi Wasmann Male genitalis unknown but most probably unlike either heimi or escherichi , from colonies of Odontotermes ( Hypoternies ) obscuriceps Wasmann in Ceylon (see description) butteli Wasmann 7. Elytra and abdomen with setae having bifurcated tips 8 Elytra and abdomen with setae having straight tips 11 8. Size small; pronotum length, 0.36-0.41 mm 9 Size larger; pronotum length, 0.47-0.55 mm 10 9. Sternites with 2 macrochaetae at each lateral edge; male genitalia shaped as in fig. 19 angolae Seevers Sternites without macrochaetae except for sternite VII which has 1 on each side; male genitalia shaped as in fig. 28 splendidus Wasmann 10. Spermatheca shaped as in fig. 33 emersoni n. sp. Spermatheca shaped as in fig. 38 vansomereni n. sp. 11. Pronotum with a single row of very fine setae along posterior border transvaalensis Silvestri Pronotum without any setae whatsoever 12 12. Abdominal tergites III-VII with no macrochaetae, male genitalia shaped as in fig. 21 eoatoni n. sp. Abdominal tergites III-VII with some macrochaetae 13 13. Macrochaetotaxy of abdominal tergites III-VII, 4, 4, 4, 4, 4; male genitalia shaped as in fig. 26 latericius n. sp. Macrochaetotaxy of abdominal tergites III-VII, 6, 6, 6, 6, 6; male genitalia shaped as in fig. 20 braunsi Wasmann DESCRIPTION OF THE SPECIES Termito discus angolae Seevers Figs. 6, 19, 44 Termitodiscus angolae Seevers, 1965, Publ. cult. Comph. Diam. Angola, 69: 134, figs. 6 and 7. Museu do Dundo (Angola: Dundo, R. Capemba, ex fungus gardens of Odontotermes nolaensis Sjoestedt, April, 1962, Coll. Machado and Sanjinje). Most closely related to T. emersoni n. sp. from which it is distinguished by its smaller size and the shape of the male genitalia. Related to T. splendidus Wasmann through its similar size, but separable therefrom by the abdominal chaetotaxy. Color light yellowish brown throughout with the antero-lateral edges of the pronotum and elytra a little lighter than the rest of the body. Dorsal surface of the head and pronotum smooth and shiny without setae of any kind but with fine punctures evenly but sparsely scattered about. Dorsal surface of the elytra and abdomen with an even covering of yellow, recumbent, short, stiff setae with bifurcated tips. No tergal macrochaetotaxy. Sternites III-VII with a double row of black macrochaetae on each lateral edge. Sternite VIII with the one row of black macrochaetae on each lateral edge and with the mesial row toward the middle. Apex of tergite VIII pointed in the female. Median lobe of the male genitalia shaped as in fig. 19. Antennae 9-segmented. measurements: Pronotum length, 0.33 mm; elytra length, 0.18 mm; pronotum width, 0.51. Number measured, 1. 212 New York Entomological Society [Vol. LXXV Figs. 17-21. Male genitalia: Lateral lobes: 17. Termitodiscus escherichi Wasmann; 18. T. vansomereni n. sp. Median lobes: 19. T. angolae Seevers; 20. T. braunsi Wasmann; 21. T. coatoni n. sp. Scale is equal to 0.25 mm. material examined: 3 specimens of the type series (C.N.H.M., D.K.). Dis- tribution shown in fig. 44. Termitodiscus braunsi Wasmann Figs. 1, 20, 31, 43 Termitodiscus braunsi Wasmann, 1912, Zeitschr. Wiss Zool., 101: 94 — Naturhistorisch Museum, Maastricht (Republic of South Africa: Orange Free State, Bothaville, with Odontotermes transvaalensis Sjoestedt) ; Seevers, 1957, Fieldiana Zool., 40: 262 (key, list). Most closely related to T. latericius n. sp. from which it is distinguished by the 9-seg- December, 1967 I Kistner: Termitophile Revision 213 merited antennae, abdominal macrochaetotaxy, and the shape of the male genitalia and spermatheca. Color light reddish brown throughout with the antero-lateral edges of the pronotum and elytra still lighter, approaching yellowish brown. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head and pronotum without setae of any kind. Dorsal surface of the elytra and abdomen with an even covering of yellow, erect setae with straight nonbifurcated tips. Macrochaetotaxy of abdominal tergites II-VII as follows: II, 0; III, 6; IV, 6; V, 6; VI, 6; VII, 6. Tergite VIII with two rows of 4 macrochaetae each. Sternites III-VII with a row of two setae on each side. Spermatheca shaped as in fig. 31. Median lobe of the male genitalia shaped as in fig. 20. Antennae 9-segmented. measurements: Pronotum length, 0.47-0.52 mm; elytra length, 0.23-0.24 mm; pronotum width, 0.80-0.85 mm. Number measured, 10. material examined: Republic of South Africa: Orange Free State, 1, Holo- type, T. braunsi Wasmann, det. E. Wasmann, Bothaville, Coll. Brauns, bearing label, “ Termes rubricola Wasmann,” (N.H.M.). Transvaal: 8, 36 mi. ex Pre- toria-Warmbad, 18 February 1963, Coll. J. Sheasby, T-12 (N.C.I., D.K.); 1 (coll.), 34 mi. ex Pretoria-Pienaars River, 8 March 1963, Coll. J. Sheasby, T-37, (N.C.I.); 3, Rooikop, Rus de Winter, 30 June 1963, Coll. J. Sheasby, T-102 (N.C.I., D.K.); 7, 32 miles ex Pretoria-Pienaars River, 7 August 1963, Coll. J. L. Sheasby, T-132 (N.C.I., D.K.); 4, 30 mi. ex Pretoria-Pienaars River, 8 January 1964, Coll. J. L. Sheasby, T-238 (N.C.I., D.K.); 1, Rooikop, Rus de Winter, 19 March 1964, Coll. J. L. Sheasby, T-325 (N.C.I.) 2, 7 miles ex Pienaars River — Rus de Winter, 20 May 1964, Coll. J. L. Sheasby, T-345 (N.C.I. , D.K.) ; 7, Rooikop, Rus de Winter, 10 March 1965, Coll. J. L. Sheasby, T-379 (N.C.I., D.K.); 1, 30.5 mi. ex Pretoria-Warmbad, 17 March 1966, ex fungus gardens, Coll. W. Coaton, J. L. Sheasby, and D. Kistner, No. 1438 (D.K.). notes: All of the hosts of the Transvaal specimens listed above were identified as Odonto- termes transvaalensis Sjoestedt by Dr. W. G. H. Coaton. The accession numbers of the termites, should future workers wish to check the hosts are as follows (in the same order as the data above) : S-6, S-16, S-22, S-30, S-56, TM. 13360, TM. 14169, & unaccessioned, all in the National Isoptera Collection of South Africa. The last numbered 1439, nest T-160, in the collection of D. Kistner. The distribution of the species is shown in fig. 43. Termitodiscus butteli Wasmann Fig. 44 Termitodiscus butteli Wasmann, 1916, Zool. Jahrb. System., 39: 181, pi. 4, fig. 10, pi. 5, fig. 10a, Naturhistorisch Museum, Maastricht (Ceylon: Peradeniya, ex fungus gardens of Odontotermes ( Hypotermes ) obscuriceps Wasmann, Coll, by von Buttel-Reepen, December 1911) ; Seevers 1957, Fieldiana Zool., 40: 262 (key and list). Closely related to T. escherichi Wasmann and T. heimi Wasmann from which it is dis- tinguishable only by its smaller size (1.4 mm vs. 1 .6—1 .9 mm). See notes below. Color yellowish brown throughout, yellower toward the antero-lateral edge of the pronotum than elsewhere. Dorsal surface of the head, pronotum, and elytra smooth and shiny with 214 New York Entomological Society [Vol. LXXV fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum, elytra, and abdomen with an even covering of fine yellow, recumbent, short, stiff setae with bifurcated tips. Macrochaetotaxy of abdominal tergites II-VII: 0, 0, 0, 0, 0, 0. Macrochaetotaxy of sternites and abdominal segment VIII unknown. Male genitalia and female spermatheca unknown. Antennae 10- segmented. measurements: Pronotum length, 0.45-0.46 mm; elytra length, 0.22-0.23 mm; pronotum width, 0.85-0.92 mm. Number measured, 2. material examined: Type and cotype (N.H.M.); 1 cotype (B.M.N.H.). The distribution is shown in fig. 44. notes: Because dissection material was not available, sufficient characters are not known to distinguish this species from either T. heimi or T. escherichi. The overall size difference was taken from the original description, but actual measurements made are all on the low side of the range for T. escherichi. I found and dissected one nest of O. obscuriceps in Kandy, Ceylon, but unfortunately did not get any specimens. No new material of this species has been collected since the original capture. The clustering program on the basis of the characters available show that it is very closely related to heimi and escherichi (1.000 correlation) and I do not believe that new material will greatly alter the association although it would undoubtedly lower the coefficient of relationship. Because heimi and escherichi are now well known, it should be easy to place this species, once material from O. obscuriceps colonies from reasonably close to Peradeniya is available. Termitodiscus coatoni n. sp. Figs. 21, 32, 43 Most closely related to T. transvaalensis Silvestri from which it is distinguished by the absence of a row of fine setae on the posterior edge of the pronotum, its 9-segmented antennae, its abdominal macrochaetotaxy, and the shape of the male genitalia and spermatheca. Color reddish brown throughout with the antero-lateral edge of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head and pronotum without any setae of any kind. Dorsal surface of the elytra and abdomen with an even covering of yellow setae with nonbifurcated straight tips. Macro- chaetotaxy of abdominal tergites II-VIII as follows: 0, 0, 0, 0, 0, 0, 2. Macrochaetotaxy of abdominal sternites III— VIII as follows: III, 2; IV, 2; V, 2; VI, 4; VII, 4; VIII, 6, all lateral except for the mesial 2 on VIII. Female tergite VIII evenly rounded on posterior edge. Spermatheca shaped as in fig. 32. Median lobe of male genitalia shaped as in fig. 21. Antennae 9-segmented. measurements: Pronotum length, 0.48-0.51 mm; elytra length, 0.21-0.25 mm; pronotum width, 0.80-0.85 mm. Number measured, 10. holotype: 1 male, No. 12515, South Africa, Transvaal, Rooikop, Rus de Winter, 19 March 1963, Coll. J. L. Sheasby No. T-47. In the National Collection of Insects, South Africa. paratypes: South Africa: Transvaal: 20, same data as holotype (N.C.I., D.K.) ; 4, 14 mi. ex Pretoria-Pienaars River Dam, 9 August 1960, Coll. W. G. H. Coaton, TM7433 (N.C.I., D.K.); 6, Pretoria West, 14 August 1963, Coll. Rorke No. T-145 (N.C.I., D.K.). December, 19671 Kistner: Termitophile Revision 215 notes: The hosts of all the captures were determined as Odontotermes badius (Haviland) by Dr. W. G. H. Coaton. The accession numbers of the termites are S-18, TM7433, and S-32 respectively and the specimens are in the National Collection of Isoptera, South Africa. The distribution of the species is shown in fig. 43. Termitodiscus emersoni n. sp. Figs. 33, 44 Most closely related to T. angolae Seevers from which it is distinguished by its larger size. Also related to T. vansomereni n. sp. from which it is distinguished by the shape of the female spermatheca. Color reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head and pronotum without any setae of any kind. Dorsal surface of the elytra and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. No macrochaetae on abdominal tergites II-VIII. Macrochaetotaxy of abdomenal sternites III-VIII, 4, 4, 4, 4, 4, 4, all lateral in position. Spermatheca shaped as in fig. 33. Male unknown. Antennae 9-segmented. measurements: Pronotum length, 0.47 mm; elytra length, 0.24-0.25 mm; pronotum width, 0.80-0.85 mm. Number measured, 2. holotype: 1 female, No. 12228, Congo Republic, Kivu, Keyberg, 25 April 1948, Coll. Alfred E. Emerson. In the collection of the author. paratype: 1 female, same data as the holotype (D.K.). notes: The host colony was identified as Odontotermes patruus Sjoestedt by Dr. A. E. Emerson. Specimens of the host colony are in the Emerson collection of the American Museum of Natural History, New York. The distribution of the species is shown in fig. 44. Termitodiscus escherichi Wasmann Figs. 2, 3, 8, 9, 10, 11, 12, 17, 22, 44 Termitodiscus escherichi Wasmann, 1911, Termitenleben auf Ceylon: 231 Naturhistorisch Museum, Maastricht (Ceylon, Perandeniya, with Odontotermes redemanni Wasmann) ; 1912, Zeitschr. wissensch Zook, 101: 94 (no additional data added) ; 1916, Zook Jahrb. Syst., 39: 181, pi. 4, fig. 9, pi. 5, fig. 9a (key) ; Cameron, 1932, Fauna Brit, India, Staphyk, 3: 318 (key); Seevers, 1957, Fieldiana Zook, 40: 260 (key). Termitodiscus escherichi var. picea Wasmann, 1916, Zook Jahrb. Syst., 39: 181 Natur- historisch Museum, Maastricht (Ceylon, Peradeniya, with Odontotermes ceylonicus Wasmann, 8 January 1912, Coll. H. von. Buttel-Reepen) ; Seevers, 1957, Fieldiana Zook, 40: 260 (synonymized variety). Most closely related to T. heimi Wasmann from which it is distinguished by the lack of ventral spines on the median lobe of the male genitalia and presence of 2 more macrochaetae on the sternites of each of abdominal segments VI, VII, and VIII, as well as the shape of the median lobe of the male genitalia. Color light reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without any setae of any kind. Dorsal surface of the pronotum, elytra, and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with 216 New York Entomological Society [Vol. LXXV Figs. 22-26. Median lobes of male genitalia: 22. Termitodiscus escherichi Wasmann; 23. T. machadoi Seevers; 24. T. krishnai n. sp.; 25. T. heimi Wasmann; 26. T. latericius n. sp. Scale is equal to 0.25 mm and applies to all figures except fig. 23. Fig. 23 after Seevers (1965). bifurcated tips. Macrochaetotaxy of abdominal tergites II— VIII : 0, 0, 0, 0, 0, 0, 4. Macro- chaetotaxy of abdominal sternites III— VIII : 2, 2, 4, 6, 6, 4. Median lobe of male genitalia without ventral spines, shaped as in fig. 22. Spermatheca shaped as in fig. 12. Antennae 10- segmented. measurements: Pronotum length, 0.45-0.50 mm; elytra length 0.22-0.25 mm; pronotum width, 0.90-1.00 mm. Number measured, 15. December, 1967] Kistner: Termitophile Revision 217 material examined: Ceylon i Holotype and Cotype, T. escherichi Wasmann, det. E. Wasmann, Peradeniya, with Odontotermes redemanni Wasmann (N.H.M.); Holotype, T. escherichi var. picea Wasmann, det. E. Wasmann, Peradeniya, with Odontotermes ceylonicus Wasmann (N.H.M.); 161, Sigiriya, ex fungus gardens of nest T 22, 25 August 1960, Coll. D. H. and A. C. Kistner (D.K.) ; 2, Sigiriya, ex fungus gardens of nest T24, 25 August 1960, Coll. D. H. and A. C. Kistner (D.K.) ; 2, Sigiriya, ex fungus gardens of nest T23, 24 August 1960, Coll. D. H. and A. C. Kistner (D.K.); 4, Sigiriya, ex fungus gardens of nest T21, 24 August 1960, Coll. D. H. and A. C. Kistner (D.K.). The dis- tribution of the species is shown in fig. 44. notes: The termite hosts of our Sigiriya captures were identified as Odontotermes taprobanes Walker by Dr. A. E. Emerson who stated that O. redemanni Wasmann is a synonym of that species. The specimens of the hosts are deposited in the Emerson collection of the American Museum of Natural History, New York. The royal cells of the above colonies were all located, opened, and were devoid of termitophiles. Termitodiscus heimi Wasmann Figs. 25, 34, 44 Termitodiscus heimi Wasmann, 1899, Deutsches Entomol. Zeitschr. 1899: 147, pi. 1, fig. la-f; Naturhistorisch Museum, Maastricht (India: Ahmednagar District, Wallon, and Sangamner with Odontotermes obesus Rambur and Odontotermes wallonensis Wasmann) ; 1912, Zeitschr. wissensch. Zook, 101: 93, pi. 5, fig. 4; 1916, Zool. Jahrb. Syst., 39: 181, pi. 4, fig. 8a-b, pi. 5, fig. 8c; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 318 (key); Silvestri, 1947, Arch. Zool. Ital., 31: 127, fig. 1 (1-7); Seevers, 1957, Fieldiana Zook, 40: 260 (key). Termitodiscus heimi var. vicinior Silvestri, 1947, Arch. Zook Ital., 31: 127, fig. 2, (India: Barkuda Island, with Odontotermes sp.) ; Seevers, 1957, Fieldiana Zook, 40: 260 (synonymized variety) . Most closely related to T. escherichi Wasmann from which it is distinguished by the presence of ventral spines on the median lobe of the male genitalia and the presence of 2 less macrochaetae on the sternites of each of abdominal segments VI, VII, and VIII, as well as the shape of the median lobe of the male genitalia. Color light reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum, elytra, and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. Macrochaetotaxy of abdominal tergites II— VIII, 0, 0, 0, 0, 0, 0, 2. Macrochaetotaxy of abdominal sternites III-VIII: 2, 2, 4, 4, 4, 2, all on the lateral edges. Median lobe of the male genitalia with 2 ventral spines, 1 on each side, shaped as in fig. 25. Spermatheca shaped as in fig. 34. Antennae 10-segmented. measurements: Pronotum length, 0.50-0.55 mm; elytra length, 0.25-0.26 mm; pronotum width, 0.95-1.07 mm. Number measured, 10. material examined: India! Holotype and 1 cotype, Ahmednagar District, Wallon, with Odontotermes obesus Rambur (N.H.M.); 11, Bombay Province, Wallon, Coll. J. B. Heim, with Odontotermes obesus (D.K.); 4, Bombay 218 New York Entomological Society [Vol. LXXV Province, Khandala, ex fungus gardens to nest T20, 21 August 1960, Coll. D. H. and A. C. Kistner (D.K.); 11, Khandala, with Odontotermes obesus , 1913, Coll. J. Assmuth (D.K.). The distribution is shown in fig. 44. notes: Host colony T20 was determined as Odontotermes obesus Rambur by Dr. A. E. Emerson and the termite specimens are deposited in the Emerson Collection of the American Museum of Natural History, New York. No specimens were found in the royal cell of this nest either. Termitodiscus krishnai n. sp. Figs. 24, 35, 44 Most closely related to T . minutus Cameron from which it is presently distinguishable only by the more sparse setae on the pronotum, elytra, and abdomen of T. krishnai. When dis- section material of T. minutus is available other characters will undoubtedly emerge. Color yellowish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum, elytra, and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. Macro- chaetotaxy of abdominal tergites II-VIII, 0, 0, 0, 0, 0, 0, 2. Macrochaetotaxy of abdominal sternites III— VIII, 0, 0, 0, 0, 0, 2. Median lobe of the male genitalia shaped as in fig. 24. Spermatheca shaped as in fig. 35. Antennae 10-segmented. measurements: Pronotum length, 0.33-0.38 mm; elytra length, 0.17-0.18 mm; pronotum width, 0.63-0.64 mm. Number measured, 2. holotype: 1 male, No. 12518, Burma, 21 mi. ex Mandalay, 23 October 1961, Coll. K. Krishna. In the collection of the author. paratype: 1 female, same data as the holotype (D.K.). notes: The host of the above specimens was identified as Odontotermes hainanensis (Light) by Dr. Kumar Krishna. The specimens of the host are deposited in the American Museum of Natural History, New York. The distribution of the species is shown in fig. 44. Termitodiscus latericius n. sp. Figs. 26, 36, 44 Most closely related to T . braunsi Wasmann from which it is distinguished by its 10- segmented antennae, the tergal macrochaetotaxy, the shape of the spermatheca, and the median lobe of the male genitalia. Color reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head and pronotum without setae of any kind. Dorsal surface of the elytra and abdomen with an even covering of long yellow setae with non-bifurcated tips which are not recumbent but not erect either. Macrochaetotaxy of abdominal tergites II-VIII: 0, 4, 4, 4, 4, 4, 2. Sternites III-VII with 2 macrochaetae on each lateral edge. Sternite VIII with 1 macrochaeta on the lateral edge and 1 about half way toward the middle on each side. Tergite VIII with the posterior edge pointed. Median lobe of the male gentalia shaped as in fig. 26. Spermatheca shaped as in fig. 36. Antennae 10-segmented. December, 19671 Kistner: Termitophile Revision 219 measurements: Pronotum length, 0.47-0.55 mm; elytra length, 0.22-0.25 mm; pronotum width, 0.70-0.85 mm. Number measured, 10. holotype: 1 male, No. 12506, Republic of South Africa, Transvaal, 33 mi ex Pretoria-Pienaars River, 22 February 1965, Coll. J. L. Sheasby, No. T378. In the National Collection of Insects, South Africa. paratypes: Republic of South Africa, Transvaal: 10, Pretoria, Waverly, 20 February 1963, Coll. J. L. Sheasby, No. T17 (N.C.I., D.K.); 4, Pretoria, Derdepoort, 4 March 1963, Coll. J. L. Sheasby, No. T31 (N.C.I., D.K.); 5, Derdepoort, 9 July 1963, Coll. J. L. Sheasby, No. T110 (N.C.I., D.K.); 1, Derdepoort, 20 January 1964, Coll. J. L. Sheasby, No. T258 (N.C.I.); 2, 9 mi ex Pretoria-Pienaars River, 2 March 1964, Coll. J. L. Sheasby, No. T306 (N.C.I., D.K.); 1, Derdepoort, 6 March 1964, Coll. J. L. Sheasby, No. T313 (N.C.I.). notes: The host colonies of all the above specimens were determined as Odontotermes latericius (Haviland) by Dr. W. G. H. Coaton. The host specimens are in the South African National Collection of Isoptera under the following accession numbers: S-7, S-14, S-23, S-59, S-65, S-66, unaccessioned (T378). The distribution of the species is shown in fig. 44. Termitodiscus machadoi Seevers Figs. 4, 5, 7, 23, 44 Termitodiscus machadoi Seevers, 1965, Publ. Cult. Comph. Diam. Angola 69: 136, figs. 8, 9, Museu do Dundo, Angola (Angola, Dundo, R. Capemba, 23 March 1962, from nest of Odontotermes interveniens Sjoestedt, Coll. A. De Barros Machado). Most closely related to T. sheasbyi n. sp. from which it is distinguished by its slightly smaller size and the absence of ventral spines from the median lobe of the male genitalia as well as by the shape of the median lobe of the genitalia. Color reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum, elytra and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. No macrochaetae on either sternites or tergites. Median lobe of the male genitalia shaped as in fig. 23. Spermatheca unknown. An- tennae 9-segmented. measurements: Pronotum length, 0.41-0.43 mm; elytra length, 0.30-0.22 mm; pro- notum width, 0.76-0.80 mm. Number measured, 3. material examined: 6 paratypes (F.M.N.H., D.K.). The distribution of the species is shown in fig. 44. Termitodiscus minutus Cameron Fig. 44 Termitodiscus minutus Cameron, 1926, Trans. Entomol. Soc. London, 74: 171 — British Museum (N.H.), London (India: Dehra Dun, in nest of termites, Coll. M. Cameron); 1932, Fauna Brit. India, Staphyl., 3: 319 (key); Seevers, 1957, Fieldiana Zook, 40: 262 (key, list) . 220 New York Entomological Society [Vol. LXXV Figs. 27-30. Median lobes of male genitalia: 27. Termitodiscus sheasbyi n. sp.; 28. T. splendidus Wasmann ; 29. T. transvaalensis Silvestri; 30. T. vansomereni n. sp. Scale is equal to 0.25 mm. Most closely related to T. krishnai n. sp. from which it is presently distinguishable only by the presence of more setae with bifurcated tips on the pronotum, elytra, and abdomen. In this regard, it is also closely related to T. escherichi and T. heimi from which it is distinguished by its much smaller size. When dissectable material is ultimately available, more definitive characters are almost certain to be found as the range of the host of T. krishnai does not extend to Dehra Dun. Color yellowish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum, elytra, and abdomen with an even covering of fine, yellow, re- cumbent, stiff, short setae with bifurcated tips. No macrochaetae on any of the tergites. Macrochaetotaxy of the sternites unknown. Male genitalia and spermatheca unknown. Antennae 10-segmented. measurements: Pronotum length, 0.33 mm; elytra length, 0.18 mm; pronotum width, 0.66-0.70 mm. Number measured, 2. material examined: Holotype plus 1, India, Uttar Pradesh, Dehra Dun, 19 March 1924, Coll. M. Cameron, from the nest of a termite (B.M.N.H.). notes: A search of the termite collection of the British Museum (N.H.) by Mr. W. A. Sands did not yield any Odontotermes bearing data corresponding to the type label. If there is any sample of the termites associated with these specimens, they might be at the Forest Research Institute at Dehra Dun, but other than that possibility, only further collections are likely to yield the host data. The distribution of the species is shown in fig. 44. December, 19671 Kistner: Termitophile Revision 221 \ 1 Figs. 31-38. Spermathecae: 31. Termitodiscus braunsi Wasmann; 32. T. coatoni n. sp.; 33. T. emersoni n. sp.; 34. T. heirni Wasmann; 35. T. krishnai n. sp. ; 36. T. latericius n. sp.; 37. T. transvaalensis Silvestri; 38. T. vansomereni n. sp. Scale is equal to 0.25. Termitodiscus sheasbyi n. sp. Figs. 27, 44 Most closely related to T. machadoi Seevers from which it is distinguished by its slightly larger size and the presence of a ventral spine from the median lobe of the male genitalia as well as by the shape of the median lobe of the male genitalia. Color reddish brown throughout with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum, elytra, and 222 New York Entomological Society IVol. LXXV abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. No tergites with macrochaetae. Macrochaetotaxy of abdominal sternites III— VIII : 0, 0, 0, 0, 0, 8. Median lobe of the male genitalia with a spine on the median ventral posterior border, shaped as in fig. 27. Female unknown. Antennae 9-segmented. measurements: Pronotum length, 0.45 mm; elytra length, 0.18 mm; pronotum width, 0.75-0.90 mm. Number measured, 3. holotype: 1 male, No. 12503, South West Africa, 30 miles ex Tsumeb- Tsinsabis (15°, 45-59' S., 17°, 45-59' E.), 26 September 1966, Coll. J. L. Sheasby, No. T502, ex fungus gardens. In the National Collection of Insects, South Africa. paratypes: 2 males, same data as holotype (N.C.I., D.K.). notes: The host of the above species was determined as Odontotermes (c.f.) latericius (Haviland) by Dr. VV. G. H. Coaton. The sample bears the accession number TM. 20457 and is in the National Isoptera Collection, South Africa. The distribution of the species is shown in fig. 44. Termitodiscus splendidus Wasmann Figs. 28, 43 Termitodiscus splendidus Wasmann, 1899, Deutsch. Entolmol. Zeitschr. 1899: 401. Natur- historisch Museum, Maastricht (Republic of South Africa: Natal, Shivyre, with Odontotermes vulgaris Haviland, Coll. Haviland); 1912, Zeitschr. wissensch. Zook, 101: 94, pi. 5, fig. 5; Seevers, 1957, Fieldiana Zool., 40: 26 2 (key, list). The (c.f.) designation given in the determination was used to indicate morphological similarity to latericius from South Africa. The nest however was constructed differently. Not very closely related to any other species but bears similarity to T. vansomereni, T. emersoni, and T. angolae by having setae with bifurcated tips on the elytra, but dis- tinguishable by its smaller size, the abdominal macrochaetotaxy and the shape of the male genitalia. Related to the sheasbyi-machadoi group through its size, macrochaetotaxy of the abdomen, and the antennal segmentation but separable therefrom by the lack of setae on the pronotum as well as genitalic characters. Color light reddish brown throughout with the antero-lateral edges of the pronotum just about the same color as the rest of the body. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head and pronotum without setae of any kind. Dorsal surface of the elytra and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. No tergites with macrochaetae. Macrochaetotaxy of abdominal sternites III— VIII : 0, 0, 0, 0, 0, 2. Median lobe of the male genitalia small, shaped as in fig. 28. Spermatheca unknown, as the female dissected lacked a spermatheca for some inexplicable reason. Antennae 9-segmented. measurements: Pronotum length, 0.36-0.41 mm; elytra length, 0.17-0.19 mm; pronotum width 0.67-0.70 mm. Number measured, 2. material examined: Holotype and 2 cotypes on a single pin, top specimen herewith designated hololectotype, Natal (Shivyre), November 1898, Coll. G. D. Haviland, with Odontotermes vulgaris Haviland (N.H.M.) ; 2, same locality, host and collector, 16 February 1898 (D.K.). notes: The distribution of the species is shown in fig. 43. December, 1967 I Kistner: Termitophile Revision 223 Termito discus transvaalensis Silvestri Figs. 13-16, 29, 37, 43 Termito discus transvaalensis Silvestri, 1947, Arch. Zool. Ital., 31: 129, fig. 3, (Transvaal, ex nest of Odontotermes angustatus Rambur, Coll. C. Fuller) ; Seevers, 1957, Fieldiana Zool., 40: 262 (key, list) . Not very closely related to any other species. Closely related to T. vansomereni through its size and abdominal macrochaetotaxy, but separable therefrom by its straight-tipped setae and its 10-segmented antennae. Closely related to T. laterieius n. sp. but separable there- from by the macrochaetotaxy of the abdominal tergites. Separable from all species by the presence of a row of fine setae with straight tips at the posterior edge of the pronotum as well as the shape of the male genitalia. Color reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body, approaching yellow. Dorsal surface of the head, pronotum, and elytra smooth and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head without setae of any kind. Dorsal surface of the pronotum generally without setae, but bearing one row of fine, short yellow setae at the posterior border. Dorsal surface of the elytra and abdomen with an even covering of fine yellow setae with straight, non- bifurcated tips. Macrochaetotaxy of abdominal tergites II-VIII: 0, 0, 0, 0, 0, 0, 2. Macro- chaetotaxy of abdominal sternites III-VIII: 6, 6, 6, 4, 4, 4, 4. Median lobe of male genitalia shaped as in fig. 29. Spermatheca shaped as in fig. 13. One aberrent spermatheca was shaped as in fig. 37, whereas other members of the same population matched fig. 13. Antennae 10-segmented. measurements: Pronotum length, 0.47-0.51 mm; elytra length, 0.24-0.26 mm; pronotum width, 0.80-0.87 mm. Number measured, 10. material examined: South Africa: Transvaal: 3, 3 mi. ex Morgenson- Standerton, 10 September 1963, Coll. J. L. Sheasby, No. T160 (N.C.I., D.K.) ; 5, 3 mi. ex Morgenson-Standerton, 10 September 1963, Coll. J. L. Sheasby, No. T161 (N.C.I., D.K.); 1, 10 mi. ex Morgenson-Standerton, 11 September 1963, Coll. J. L. Sheasby, No. T167 (N.C.I.); 2, 13 mi. ex Morgenson-Ermelo, 12 September 1963, Coll. J. L. Sheasby, No. T168 (N.C.I., D.K.). Cape Province: 11, 6 mi. ex Sterkstroom-Tarka, 8 October 1963, Coll. J. L. Sheasby, No. T206 (N.C.I., D.K.); 13, 10 mi. ex Cala-Indwe, 7 October 1963, Coll. J. L. Sheasby, No. T202 (N.C.I., D.K.) . notes: The hosts of all of the above specimens were determined as Odontotermes angustatus (Rambur) by Dr. W. G. H. Coaton. The hosts bear the accession numbers S-37, S-40, S-41, TM 13045, TM 13059, and are in the National Collection of Isoptera, South Afirca. The distribution of the species is shown in fig. 43. Termitodiscus vansomereni n. sp. Figs. 18, 30, 38, 44 Most closely related to T. emersoni n. sp. and T. angolae Seevers from which it is dis- tinguished by its larger size, the abdominal macrochaetotaxy and the shape of the male genitalia. Closely similar to O. transvaalensis Silvestri from which it is distinguished by the presence of setae with bifurcated tips. Color light reddish brown throughout, with the antero-lateral edges of the pronotum lighter than the rest of the body. Dorsal surface of the head, pronotum, and elytra smooth 224 New York Entomological Society [Vol. LXXV Figs. 39-40. Setae on elytra: 39. Straight tipped setae, Termitodiscus transvaalensis Silvestri; 40. Bifurcated tipped setae, T. escherichi Wasmann. Scale is arbitrary, photos were taken at 440 X . and shiny with fine punctures evenly but sparsely scattered about. Dorsal surface of the head and pronotum without setae of any kind. Dorsal surface of the elytra and abdomen with an even covering of fine, yellow, recumbent, stiff, short setae with bifurcated tips. Tergites with no macrochaetae. Macrochaetotaxy of abdominal sternites III— VIII : 6, 6, 6, 6, 6, 6,. Median lobe of the male genitalia shaped as in fig. 30. Spermatheca shaped as in fig. 38. Antennae 9-segmented. measurements: Pronotum length, 0.50-0.53 mm; elytra length, 0.26-0.30 mm; pronotum width, 0.85-0.95 mm. Number measured, 10. holotype: 1 male, No. 12224, Kenya, Karen, 18 June 1966, ex fungus gardens of nest T185, Coll. G. R. Cunningham-Van Someren, No. 1559. In the Collec- tion of D. H. Kistner. paratypes: 54, same data as the holotype (D.K.). notes: The host of the above specimens was determined as Odontotermes montanus Harris by Mr. W. A. Sands. The termite sample is in the collection of the British Museum (Natural History), London. This nest was being raided by Dorylus ( Doryhis ) helvolus L. at the time of excavation. The distribution of the species is shown in fig. 44. RELATIONSHIPS OF THE SPECIES AND HOST SPECIFICITY In the early days of describing species, various authors made a big point about the relative size of the last joint of the antennae in relation to the length of the rest of the segments as well as the absolute length of the entire specimen. Careful slide preparations have revealed that this is an almost useless character as the size of the terminal segment is always proportionate to the rest of the antenna. Figures 3 and 4 show this well, even though the number of antennal segments vary. The length of the entire specimen is another useless character December, 1967] Kistner: Termitophile Revision 225 as the abdomen is able to be telescoped a great deal. Hence both of these char- acters were dropped. In searching for new characteristics, microscopic examination revealed the following: Not all the species had 10-segmented antennae as was previously supposed. I first discovered this on T. machadoi. I then remembered that Silvestri had shown a 10-segmented antenna on both T. transvaalensis and T. heimi. This worried me as I have never known Silvestri to be wrong on a ques- tion of fact. Sure enough, both of the species studied by Silvestri had 10-seg- mented antennae. I then surveyed the antennae of all the species and could find no correlation of the antennal segmentation with any other character. Hence this character is here interpreted as a species specific character, and a new genus is not erected on this basis. Selection has obviously been working to compress the antennae of this beast, so a segment has been lost now and then on what appears to be a hit or miss basis. I believe that the segment has been lost between segment 3 and 5 and on one species, T. vansomereni, one can see what appears to be a fine line of fusion on the third segment. Close study of the setae revealed that there are two types. One type is a perfectly ordinary kind with straight pointed tips as shown in fig. 39. The other type has bifurcated tips as shown in fig. 40. The difference between the setae was noted by Seevers ( 1957, p. 260) as being feebly notched. He inter- preted this as being only in the Indian species which was not true. Among the species described at that time, T. splendidus also had such setae. Using traditional phylogenetic methods, it is possible to construct a phylogeny of the species groups as shown in fig. 4. Group A consisting of T. braunsi, T. latericius, T. coatoni, and T. transvaalensis would be interpreted as the most primitive species because they have setae with straight tips which is the usual situation in the Staphylinidae and particularly true in the primitive groups. Where deviations have occurred as in Phyllodinarda (see Kistner 1965), the deviations are of a different nature than for Termito discus and can therefore be assumed to be of independent origin. Of the four species, T. transvaalensis is probably the most primitive as it still has a short row of setae on the pronotum whereas the others lack pronotal setae entirely. Again, the complete absence of setae on the pronotum of a Staphylinid is an unusual condition and is there- fore interpreted as being a derivative condition. This view is reinforced by the fact that groups C, D, and E have pronotal setae, albeit modified, and modified setae had to be derived from some pre-existing setae, hence I am supposing that the common ancestor had to have setae, most likely unmodified setae on its pronotum. None of the presently known species quite fills the bill, but T. transvaalensis comes closest. T. latericius is more closely related to T. transvaalensis in that it has 10-segmented antennae whereas the other species ( T . braunsi and T. coatoni) have 9-segmented antennae. Groups B, C, D, and E are all related in having setae with bifurcated tips. 226 New York Entomological Society [Vol. LXXV Group A Group B Group C Group D Group E no pronotal setae small genitalia small size 9 segmented antennae setae with bifurcated tips 10 segmented antennae large genitalia large or intermediate size setae with straight tips Fig. 41. Proposed phytogeny of species groups of T ermitodiscus using traditional methods. Group A includes T. braunsi, T. latericius, T. eoatoni, and T. transvaalensis. Group B includes T. angolae, T . emersoni, T. vansomereni, and T. splendidus. Group C includes T. machadoi and T. sheasbyi. Group D includes T. krishnai and T. minutus. Group E includes T. butteli, T. escherichi, and T. heimi. Slide preparations of most of the species revealed that the bifurcated tips are all of the same type. T. minutus and T. butteli were not so examined but the dry preparations revealed no differences. Group B has no pronotal setae, but the elytra and abdomen have the bifurcated setae. All the members of this group (T. angolae , T. emersoni, T. splendidus , and T. vansomereni) have 9-segmented antennae which would link them to part of group A, and also to group C. Group C has setae with bifurcated tips on the pronotum as well as the elytra and abdomen. A careful examination of the diagram (fig. 41) will reveal that we are assuming that the common ancestor of B, C, D, and E had setae on all three regions, that this became bifurcated, and then was lost on the pronotums of group B. Thus group C would be more primitive than group B. Groups D and E share with group C the property of having setae with bifurcated tips but differ in having 10-segmented antennae. Hence I interpret that groups D and E were split off earlier in the evolution of the groups before segment reduction. Groups D and E are very closely related to one another but differ in size and in the size of the genitalia (where known). December, 1967] Kistner: Termitophile Revision 227 It is obvious from the foregoing that I did not use characters such as the macrochaetotaxy of the abdomen or the various characters of the male genitalia (other than gross size) in the construction of the phylogenetic tree. These characters, while useful for discriminating species, are presently of no use in determining phylogenies, as there is no way of determining or guessing the primitive and derivative states of such characters. Should an ancestral type be found in nature, it might be possible to judge this in the future, but this is not so at present. Computer methods were then used to see if a more precise statement of the relationships of the species could be constructed. To do this, it was necessary to develop a list of unit characters following the general outline of Sokal and Sneath (1963). After eliminating characteristics which were redundant or invariant, the following list of 31 characters was used and coded 0 for absence, 1 for presence, and 3 for no comparison. The no comparisons arose when a male character was listed and the species was known only from a female or the ma- terial studied could not be dissected to yield the desired comparison. LIST OF CHARACTERS USED FOR NUMERICAL ANALYSIS 1. Pronotum with setae with bifurcated tips 2. Elytra with setae with bifurcated tips 3. Abdomen with setae with bifurcated tips 4. Tergite VIII of male pointed 5. Male genitalia small 6. Pronotum with posterior edge with 1 row of straight tipped setae. 7. Ten antennal segments 8. Male genitalia with median spines 9. Tergite VIII of female pointed 10. Pronotum length, 0.47-0.55 mm 1 1 . Pronotum length, 0.43-0.45 mm 12. Pronotum length, 0.33-0.41 mm 13. Elytra length, 0.25-0.30 mm 14. Pronotum width twice pronotum length 15. No macrochaetae on tergites II-VII 16. Tergal macrochaetotaxy (II-VII) 0, 6, 6, 6, 6, 6 17. Tergal macrochaetotaxy (II-VIII) 0, 4, 4, 4, 4, 4 18. No macrochaetae, sternite III 19. 2 macrochaetae, sternite III 20. No macrochaetae, sternite IV 21. 2 macrochaetae, sternite IV 22. No macrochaetae, sternite V 23. 2 macrochaetae, sternite V 228 New York Entomological Society [Vol. LXXV 24. 4 macrochaetae, sternite V 25. No macrochaetae, sternite VI 26. 2 macrochaetae, sternite VI 27. 4 macrochaetae, sternite VI 28. No macrochaetae, sternite VII 29. 2 macrochaetae, sternite VII 30. 4 macrochaetae, sternite VII 3 1 . Pronotum with setae of any type The distribution of these characteristics in the 15 species is given in Table 1. These data were then loaded into an IBM 1620 computer with a program to produce the simple matching coefficients described by Sokal and Michener (1958). The output of this program was then used to cluster the data using the weighted pair-group method described by Sokal and Sneath (1963). The results of these analyses are presented in fig. 42. Only the matrix values where the groups join are indicated. The perfect correlation between T. butteli and T. heimi (and for that matter between T. butteli and T. escherichi also) is due to the large number of no comparisons in the original data. It is also probable that the correlation between T. krishnai and T. minutus will not be as high when dissection material of T. minutus is available. It will be noted that there are some major discrepancies between the phylo- genetic diagram and fig. 42. The most serious discrepancies are the clustering of T. vansomereni with T. transvaalensis , and the clustering of T. splendidus with the cluster T. sheasbyi-T. machadoi. Less serious but still important is the clustering of T. latericius to T. braunsi rather than to T. transvaalensis . All of these are due to a weighting of size and chaetotaxy factors as equal to kinds of setae and antennal segmentation. Table 1. Distribution of unit characters in Termitodiscus species. Characters are arranged sequentially from left to right. Species No. Species name Characters 01 T. angolae Seevers 02 T. braunsi Wasmann 03 T. butteli Wasmann 04 T. coatoni n. sp. 05 T. emersoni n. sp. 06 T. escherichi Wasmann 07 T. heimi Wasmann 08 T. krishnai n. sp. 09 T. latericius n. sp. 10 T. machadoi Seevers 11 T. minutus Cameron 12 T. sheasbyi n. sp. 13 T. splendidus Wasmann 14 T. transvaalensis Silvestri 15 T. vansomereni n. sp. 0111100010010010001010100100100 0001100011001001001010100100100 1113301331001110033333333333331 0001100001001010001010100010010 0113300301001010001010100100100 1111001011001110001010010000001 1111001111001110001010010010011 1111101010010110010101000100011 0000101011001000101010100100100 1 1 10100000100110010101001001001 1113301300010110033333333333331 1 1 10100101000110010101001001001 0110100000010010010101001001000 0000 11100100 1010000000000000000 01 1 1100001001010000000000000000 December, 1967] Kistner: Termitophile Revision 229 vansomereni 15 transvaalensis 14 coatoni 4 latericius 9 braunsi 2 emersoni 5 angolae I splendidus 13 sheasbyi 12 machadoi 10 krishnai 8 minutus II butteli 3 heimi 7 escherichi 6 Coefficient of Association Fig. 42. Diagram of the phenetic relationships between the species of Termito discus. The realtionships diagrammed in fig. 42 are probably less accurate as a phylogenetic scheme than the relationships shown in fig. 41. However, the information in fig. 42 is very useful as purely taxonomic information. It took only 45 minutes to put the problem through the computer and that 45 minutes saved hours of time in constructing the keys to species. As the species are known now, there is complete host specificity. The host information is summarized in Table 2. If we make the assumption that the rates of evolution of the termites and termitophiles are about the same and that there were no accidental host changes in evolutionary history, both hand- some assumptions, then we should expect that the termites would be related to each other in the same manner as the termitophiles. Thus we would expect Odontotermes heimi and Odontotermes taprobanes to be more closely related to each other than to Odontotermes hainanensis . We would expect the species O. latericius from S. W. Africa that is the host of T. sheasbyi to be more closely related to O. interveniens than to O. latericius from South Africa. It will be 230 New York Entomological Society [Vol. LXXV Table 2. Host relationships of Termito discus. Termitophile Host T. angolae Seevers Odontotermes nolaensis Sjoestedt T. braunsi Wasmann Odontotermes transvaalensis Sjoestedt T. butteli Wasmann Odontotermes ( Hypotermes ) obscuriceps Wasmann T. eoatoni n. sp. Odontotermes badius Haviland T . emersoni n. sp. Odontotermes patruus Sjoestedt T. escherichi Wasmann Odontotermes taprobanes Walker T. heimi Wasmann Odontotermes obesus Rambur T. krishnai n. sp. Odontotermes hainanensis Light T. laterieius n. sp. Odontotermes laterieius Haviland T. machadoi Seevers Odontotermes interveniens Sjoestedt T. minutus Cameron not known T. sheasbyi n. sp. Odontotermes laterieius Haviland T. splendidus Wasmann Odontotermes vulgaris Haviland T. transvaalensis Silvestri Odontotermes angustatus Rambur T. vansomereni n. sp. Odontotermes montanus Harris interesting to see whether either of the arrangements given here corresponds with the relationships between the species of Odontotermes , when this genus is revised. BEHAVIORAL OBSERVATIONS Two species were studied closely in the field, T. heimi and T. escherichi , especially to see what transpired when the termitophile came into contact with the termite host. To do this, living specimens of the termitophiles and their hosts were placed in petri dishes with moist filter paper on the bottoms and some pieces of fungus gardens. The termitophiles and termites were observed after a couple of hours had elapsed to give them time to accommodate to the container. The chief behavorial adaptation of the termitophile appeared to be avoidance. The termitophile is small in relation to the size of the termite workers or soldiers, it has good eyesight whereas the termite does not, and it is fast on its feet whereas the termite is slow and clumsy. In every termitophile-termite encounter, the termitophile was able to maneuver out of range of the mandibles before the termite was even aware of its existence. We maneuvered some termites into position with a camel’s hair brush and then tried to prevent the termitophile from escaping with another camel’s hair brush, but in every instance, the beetle was able to crawl under or around the termite without getting caught or even attract- ing attention. We were thus unable to acquire any insight into the possible adaptive function of the limuloid body shape. These same observations were confirmed in a limited way on T. braunsi in South Africa. However, future studies should be directed to see if there is any difference in the behavior of those forms with setae with bifurcated tips and those with straight tips. December, 1967] Kistner: Termitophile Revision 231 Fig. 43. Distribution of certain South African species of Termito discus. 232 New York Entomological Society [Vol. LXXV butteli ★ emersoni latericius minutus Fig. 44. Distribution of certain species of Termito discus. December, 1967] Kistner: Termitophile Revision 233 Observations in the field of T. heimi and T. escherichi revealed that both species ate the fungus in the fungus gardens. Subsequent gut smears confirmed this. The above observations should be combined with another field observation before any conclusions are drawn. Invariably, the most Termitodiscus were found in the fungus gardens immediately adjacent to the royal cell. These are the fungus gardens which contain the most eggs and young termites and hence there is more termite activity. Termite activity decreases in the more peripheral fungus gardens and one seldom finds termitophiles in these. Termitophiles were never taken in the royal cells of Odontotermes. Because of their association with the termites in areas of high termite activity, their perfect host specificity, and the fact that no accidental capture of Ter- mitodiscus outside the termite nest has ever been made, I am interpreting the genus as integrated termitophiles whose principal adaptation to the termite hosts is avoidance of direct contact. Wasmann (1895 and elsewhere) erected the category of “trutztypus” or defensive forms and placed the genus Termitodiscus in that category in 1912 and 1916 based on the morphology alone. There is no evidence that the ter- mitophiles lead a harried existence in the nest. The avoidance of the termites under observation never led to a confrontation even when we tried to manipulate one. What seems to prevail is a kind of wary but completely dependent co- existence on the part of the termitophile and an unawareness on the part of the termites. RELATIONSHIP OF THE TRIBE TO OTHER ALEOCHARINAE The closest free-living aleocharine tribe to the Termitodiscini is the tribe Myrmedoniini. The following characters link the two tribes: (1) Nature of the teeth on the mandibles; (2) Structure of the legs; (3) Tarsal formula; (4) Structure of the prosternum; (5) The tri-lobed nature of the ninth abdominal segment. The only termitophilous tribe that is close to the Termitodiscini is the sub- tribe Termitondina of the Myrmedoniini which may share common origins. More material of the Termitondina will be necessary before these relationships can be checked. The relationship of the Termitodiscini to the Myrmedoniini does not destroy the tribal status of the Termitodiscini; it merely gives some idea of what the ancestral type must have been like. Acknowledgments A study like this is only possible with the cooperation and active interest of many people. For help in my field work, I cheerfully thank Dr. W. G. H. Coaton, Plant Protection Research Institute, Pretoria, South Africa; Dr. R. Lawrence and his son, Natal Museum, Pieter- 234 New York Entomological Society [Vol. LXXV maritzburg, South Africa; Mr. G. R. Cunningham-Van Someren, Karen, Kenya; Dr. T. Fletcher, Institute for the Study of Malaria and Arthropod-borne Diseases, Amani, Tanzania; and Dr. Joseph De Sa, Bombay, India. Thanks are also given to my wife, Alzada Carlisle Kistner, for hours and hours of fungus sorting in the field. For providing specimens collected by themselves and colleagues, I wish to thank par- ticularly, Dr. W. G. H. Coaton. He and his colleagues, particularly Mr. J. L. Sheasby, have amassed a termitophile collection over the past six years for the Republic of South Africa and its dependency, Southwest Africa which is without equal for any other area of the world. For other specimens, I thank Dr. Alfred E. Emerson, University of Chicago, Dr. Kumar Krishna, American Museum of Natural History, and Mr. G. R. Cunningham-Van Someren. I am also grateful to individuals for providing facilities and space during type study trips and also for cordial hospitality while in their institutions. These are Professor J. K. A. Van Boven, Universite de Louvain, Belgium, also Curator of the Wasmann Collection at the Naturhistorisch Museum, Maastricht (N.H.M.) ; Mr. P. Basilewsky, Chef de la Section d’Entomologie, Musee Royal de l’Afrique Centrale Tervuren (M.R.A.C.) ; Mr. J. Balfour- Browne, British Museum (Natural History), London (B.M.N.H.), and Dr. Rupert L. Wenzel, Field Museum of Natural History, Chicago (F.M.N.H.). The initials given above indicate the institution wherein specimens cited are deposited. Specimens deposited in the collection of the author are indicated (D.K.) while those in the National Collection of Insects, Pretoria, South Africa are indicated (N.C.I.). Termite host identifications are all credited in the text but I am extremely grateful to Dr. W. G. H. Coaton, Dr. A. E. Emerson, Dr. Kumar Krishna, and Mr. W. A. Sands, Termite Research Unit at the British Museum (Natural History) for taking the time to make the determinations. Thanks are given to Mr. William Lane of the Computer Center and Division of Engineer- ing, Chico State College, for helpful suggestions and patient instruction on the IBM 1620 computer. Thanks are also given to Mr. Herbert Jacobson and Mr. Robert Banfill for help on the programming necessary for the numerical analyses and for help in debugging (no pun intended) the programs after modification. I am very grateful to Mr. R. Gary Malin and Mr. David Harwood of Chico State College for assistance in the mounting, labelling, dissecting, and other operations necessary in this kind of work. Literature Cited Blackwelder, Richard E. 1952. The generic names of the beetle family Staphylinidae. U.S. Nat. Mus. Bull. 200: IV + 484 p. Cameron, Malcolm. 1926. New species of Staphylinidae from India. Part II. Trans. Entomol. Soc. London, 1925 (1926): 341-372. — . 1932. The fauna of British India, including Ceylon and Burma. Staphylinidae, 3: 443 p. London. Kistner, David H. 1965. A revision of the species of the genus Phyllodinarda Wasmann with notes on their behavior ( Coleoptera: Staphylinidae) . Pan-Pacific Entomol., 41(2) : 121-132. — . 1966. A revision of the African species of the Aleocharine tribe Dorylomimini ( Coleoptera: Staphylinidae) . II. The genera Dorylominus, Dorylonannus, Dorylogaster, Dorylobactrus, and Mimanomma, with notes on their behavior. Ann. Entomol. Soc. Amer., 59(2) : 320-340. — . 1967. The biology of termitophiles. In Krishna, Kumar and F. W. Lechleitner, The Biology of Termites, Chapter 32. Academic Press, N.Y. (In press). December, 1967] Kistner: Termitopiiile Revision 235 Koblick, T. A., and D. H. Kistner. 1965. A revision of the species of the genus Myrmechusa from tropical Africa with notes on their behavior and their relationship to the Pygo- stenini (Coleoptera:Staphylinidae) . Ann. Entomol. Soc. Amer., 58(1): 28-44. Seevers, Charles H. 1957. A monograph on the termitophilous Staphvlinidae (Coleoptera) . Fieldiana: Zool., 40: 1-334. . 1965. New termitophilous Aleocharinae from Angola (Coleoptera:Staphylinidae) . Publ. Cult. Comph. Diam. Angola, Lisbon, 69: 129-138. Silvestri, Filippo. 1947. Contributo alia conoscenza dei Termitodiscinae e Cephaloplectinae (Staphylinidae, Coleoptera) termitofili. Arch. Zool. Ital., 31: 123-149. Sokal, R. R., and C. D. Michener. 1958. A statistical method for evaluating systematic relationships. Univ. Kans. Sci. Bull., 38: 1409-1438. Sokal, R. R., and P. H. A. Sneath. 1963. Principles of Numerical Taxonomy. Freeman & Co., San Francisco, XVIII + 360 pp. Wasmann, Erich. 1895. Die Myrmecophilen und Termitophilen. Compt. Rend. Ill Congr. Internat. Zool. Leyden., 1896: 410-440. — . 1899. Neue Termitophilen und Myrmekophilen aus Indien. Deutsch. Entomol. Zeitschr., 1899: 145-169. 2 plates. . 1911. Termitophile Coleopteren aus Ceylon. In Escherich, Termitenleben auf Ceylon: 231-232. . 1912. Neue Beitrage zur Kenntnis der Termitophilen und Myrmekophilen. Zeitschr. Wissenschaft. Zool., 101: 70-115. Plates V-VII. — — — . 1916. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien, V. Ter- mitophile und myrmecophile Coleopteren, gesammelt von Herrn Prof. Dr. V. Buttel-Reepen, 1911-1912. Zool. Jahrb. System., 39: 169-210. Plates 4 and 5. Received eor Publication July 13, 1967 The Immature Instars of the Cleptoparasitic Genus Dioxys (Hymenoptera: Megachilidae) Jerome G. Rozen, Jr.1 Abstract: The last-stage larva of Dioxys pomonae pomonae and D. productus productus? are described taxonomically and compared with the previously published account of the larva of D. cincta (Jurine). The three other larval instars and the pupa of D. pomonae pomonae are also described and the adaptive significance of some of the anatomical features of the larvae are discussed. A preliminary key is presented to distinguish among the genera of parasitic megachilid bees on the basis of the last larval instar. The purposes of this paper are ( 1 ) to describe taxonomically the immature instars of the parasitic bee genus Dioxys and (2) to compare the external anatomy of the four larval instars of Dioxys pomonae pomonae. At the end, a key is given that may help in the identification of mature larvae of parasitic Megachilidae. Cleptoparasitism (social parasitism) has evolved in at least three separate cases in the family Megachilidae. Coelioxys, usually a parasite of Megachile but also associated with Centris , Anthophora , and probably others, obviously arose from a Megachile- like ancestor. Stelis, sensu lato (including Euaspis and Parevaspis) and Dioxys (and related genera) presumably evolved from separate lineages in the Anthidiini. Most Stelis, sensu lato, apparently attack mega- chilids, although some (and perhaps all) species of the subgenus Odonto- stelis attack Euglossa (Friese, 1925; Bennett, 1966). The biology and larvae of Stelis are sufficiently diverse to raise the question whether this genus is monophyletic (Rozen, 1966). Insofar as known, all members of the Dioxys complex parasitize the Megachilinae (Hurd, 1958; Jaycox, 1966), but our lack of knowledge of their immature stages and biology does not permit us to specu- late on the origin of parasitism in this group. I hope that data recorded here, as well as biological information presented in the accompanying report (Rozen and Favreau, 1967) will eventually be used for this purpose. The number of larval instars in bees has been open to question because of the difficulty in rearing these animals. However, Hackwell and Stephen (1966) claim on the basis of carefully accumulated data that the halictid Nomia melanderi Cockerell has five instars. These men observed that the egg chorion encased the entire first instar except for most or all of the head capsule and that the first and second instars were similar in size. The first and second instars moved their mandibles back and forth and occasionally consumed liquid and pollen grains. Rozen (1964) stated that the embryo of the anthophorid 1 Chairman and Curator, Dept. Ent., Amer. Mus. Nat. Hist. 236 December, 1967] Rozen: Immature Instars of Dioxys 237 Figs. 1-8. Mature larva of Dioxys pomonae pomonae Cockerell. 1. Predefecating larva, lateral view (setae not shown). 2. Spiracle. 3-5. Right mandible, dorsal, inner, and ventral views, respectively. 6. Head, front view. 7. Labium, with mandibles removed, showing hypopharyngeal lobes, front view. 8. Head, lateral view. Scale refers to Fig. 1. Svastra obliqua obliqua (Say) ingested liquid just before the chorion was cast off; shortly after eclosion a transparent embryonic cuticle was shed. The “first instar” of N omia melanderi and the late “embryo” of Svastra obliqua obliqua are probably the same stage. If this is true, then the cryptic early stage may be a widespread phenomenon among bees, as N omia and Svastra belong to separate families and as this stage has also apparently been observed 238 New York Entomological Society [Vol. LXXV in the Panurginae (Rozen, 1967). Whether it represents the first instar, per- haps especially adapted to the task of ingesting fluid prior to casting off the chorion, or whether it is a late embryo can be determined only by further studies. In the case of Dioxys pomonae pomonae four distinct larval instars were observed. The early cryptic stage was not noticed though it may have been present. For the purpose of this paper, the “first instar” is the first actively moving stage, to which the chorion no longer adheres. The specimens of Dioxys productus productus? described below were kindly made available by Dr. Elbert R. Jaycox, University of Illinois, Urbana. Dr. Paul D. Hurd, Jr., University of California, Berkeley, identified the adult Dioxys. The literature search was facilitated by the Bibliography of Apoid Biology directed by Dr. Charles D. Michener, the University of Kansas, Lawrence. Mrs. Marjorie Favreau ably assisted me in the field investigations and laboratory work which culminated in this study. My wife, Barbara, helped prepare the scientific illustrations, and Mrs. Rose Ismay carefully typed the manuscript. MATURE LARVAE Only a single account of an immature of this genus occurs in the literature; Micheli (1936) provided a useful description of the mature, fourth-stage larva of the European Dioxys cincta (Jurine), the type of the genus. Grandi (1934) reported on an unknown bee larva associated with Chalicodoma muraria (Fa- bricius), and although Michener (1953a) tentatively assigned it to Dioxys , the hairy mandible identifies it as a Coelioxys. I am describing here the mature larvae of two other species of Dioxys , D. pomonae pomonae and D. productus productus? . The three known species have a number of features that may prove diagnostic for the genus. Unlike mature larvae of other megachilids, which are heavily pilose, those of Dioxys possess only widely scattered setae on the postcephalic region. The setae, sparse on the thorax, are even sparser on the abdominal seg- ments. The bidentate mandibles (Figs. 3-5, 11-13) of the three species lack the apical concavity and cusp of other members of the family (except for some Stelis, Rozen, 1966) and differ from those of other Anthidiini (except some Stelis , ibid., and Trachusa, Michener, 1953a) in that there are no small teeth on the margin between the apical teeth; of the three forms, only D. pomonae pomonae (Figs. 3-5) has such teeth on the upper and lower mandibular edges. The antennae of D. cincta apparently are not abnormally large for a megachilid but those of D. pomonae pomonae (Fig. 8) are distinctly greater in size than those of members of other genera. The antennae of D. productus productus? (Fig. 15), however, are the largest of any bee larva that I have seen. Antennal size therefore is helpful, both for species separation and, in some cases, for identification of the genus. December, 1967] Rozen: Immature Instars of Dioxys 239 Figs. 9-15. Mature larva of Dioxys productus productus (Cresson) ? 9. Larva, lateral view (setae not shown). 10. Spiracle. 11-13. Right mandible, dorsal, inner, and ventral views. 14, 15. Head, frontal and lateral views. Scale refers to Fig. 1. In other respects, the fourth instar of Dioxys seems to possess the features of other members of the family. Whether the distinctive characters mentioned above warrant placing the genus in a separate tribe as contemplated by Michener (1944) after studying the adults, is open to question. In general the larvae of megachilids appear so similar that it is difficult to imagine that larval features will be of much assistance in arranging the higher classification of the family. Dioxys pomonae pomonae Cockerell Figures 1-8 head: (Figs. 6-8) Integument with numerous scattered long setae and without spicules; antennae, labrum, pleurostomal ridges, hypostomal ridges, mandibles, cardines, stipites, palpi, 240 New York Entomological Society [Vol. LXXV and base of prementum conspicuously pigmented. Tentorium complete and well developed; posterior pits conspicuous and normal in position; posterior thickening of head capsule and hvpostomal ridge well developed ; pleurostomal ridge and lateral arms of epistomal ridge moderately developed but not so sharply defined as hypostomal ridge; epistomal ridge fading just mesiad of anterior tentorial pits; longitudinal thickening of head capsule, cleavage lines, and parietal bands not evident; head constricted behind as in D. productus productus? but dorsolateral angles of capsule less produced. Antennal papilla elongate, apparently more so than that of D. cincta (Micheli, 1936) but papilla distinctly smaller than that of D. productus productus? ; papilla slightly shorter than three times basal diameter; each papilla arising from inconspicuous prominence; these prominences distinctly less pronounced than those of D. productus productus? . Labrum without tubercles and with apical margin emarginate medially. Mandible (Figs. 3-5) without conspicuous setae, more elongate than that of D. cincta (Micheli, 1936), and apically bidentate with ventral tooth longer; margin between apical teeth smooth (i.e., nonserrate) ; dorsal apical edge with small but distinct teeth; ventral apical edge with inconspicuous serrations; apical concavity and cusp not present. Maxilla with basal part somewhat enlarged and with apex produced adorally; galea absent; palpus elongate but shorter and narrower than antennal papilla; cardo and stipes sclerotic. Labrum projecting, divided into prementum and postmentum and bearing salivary opening at apex; salivary opening a transverse slit with projecting lips; labial palpi perhaps slightly more slender than maxillary palpi; hypopharynx (Fig. 7) with prominent lobe on each side of maxilla. body: Form (Fig. 1) moderately robust; most body segments divided dorsally into low cephalic annulet and elevated caudal annulet on postdefecating larva; annulations on predefecating form indistinct; caudal annulets on postdefecating form low medially so that larva appears to have paired transverse dorsolateral tubercles; middorsal tubercles absent; lateral tubercles (below spiracles) well developed (at least on postdefecating form). In- tegument soft ; scattered setae (not shown in illustration) found on caudal annulets, lateral tubercles, and venter; these setae approximately as dense as those of D. productus productus? , but much sparser than those of host Osmia nigrobarbata and other megachilids. Spiracular atrium (Fig. 2) large, with ridges; atrium projecting somewhat above body wall and with rim ; peritreme present but narrow so that opening appears large ; primary tracheal opening without distinct collar; subatrium normally long. Tenth abdominal segment moderate in length and with anus situated dorsally. material studied: One postdefecating larva, 3 miles north of Apache, Cochise County, Arizona, April 30 through May 4, 1966; larva preserved October 14, 1966; from nest of Osmia nigrobarbata Cockerell (J. G. Rozen and M. Favreau) ; two predefecating mature larvae, same data except preserved at time of collection. Dioxys productus productus (Cresson) ? Figures 9-15 These larvae were discussed by Jaycox (1966). head: (Figs. 14, 15) As described for D. pomonae pomonae except for following: Dorso- lateral angles of head produced, apparently as in D. cincta (Micheli, 1936), and more so than in D. pomonae pomonae. Antennal papilla enormously elongate, being a little over three times longer than basal diameter; each papilla arising from restricted but pronounced prominence. Mandible (Figs. 11-13) like that of D. pomonae pomonae except dorsal and ventral apical edges without teeth or serrations. December, 1967] Rozen: Immature Instars of Dioxys 241 Figs. 16-22. First instar of Dioxys pomonae pomonae Cockerell. 16. Larva, lateral view. 17. Spiracle. 18-20. Head, frontal, lateral, and ventral views, respectively. 21, 22. Right mandible, dorsal and inner views. Scale refers to Fig. 16. 242 New York Entomological Society [Vol. LXXV body: (Fig. 9) As described for D. pomonae pomonae except spiracular subatrium short (Fig. 10). material studied: One mature larva, Smithfield, Cache County, Utah, June 30, 1962, from Anthidium nest in small plastic tube; fixed July 2, 1962 (E. R. Jaycox) ; one mature larva, same locality, June 31, 1961 ; from nest of Anthidium ; fixed July 19, 1962 (E. R. Jaycox). OTHER INSTARS None of the other immature instars of this genus has been described before; all of the following belong to Dioxys pomonae pomonae. First-Stage Larva of Dioxys pomonae pomonae Cockerell Figures 16-22 head: (Figs. 18-20) Head hypognathous, not prognathous as in Coelioxys. Integument slightly pigmented and, unlike that of host, with scattered long setae. Tentorium complete, including thin dorsal arms; posterior tentorial pits normal in position; posterior thickening of head capsule and hvpostomal ridge slender but evident ; pleurostomal ridge weak except at mandibular articulations; epistomal ridge scarcely evident, both mesiad and laterad of anterior tentorial pits; these pits well developed; longitudinal thickening of head capsule faint; cleavage lines and parietal bands not evident; head somewhat constricted behind; genal area, unlike that of Coelioxys, not produced anteroventrally into long tubercle-like projection. Antennal papilla greatly elongate, length about four times basal diameter; each papilla arising from conspicuous prominence. Labrum without tubercles and with apical margin emarginate medially and with sensilla. Mandible (Figs. 21, 22) elongate, without conspicuous setae, and with apex simple, tapering, curved, and pigmented. Maxilla with basal part greatly enlarged and sclerotized (Fig. 19); apical part directed adorally; palpus elongate; galea absent. Labium, unlike that of other bee larvae, not extending ventrally so far as maxillae; labium recessed, not divided into prementum and postmentum, and ap- parently somewhat sclerotized though not so strongly sclerotized as that of first instar of Coelioxys ; salivary opening a small transverse slit; palpi shorter than maxillary palpi, about as long as basal diameter. body: Form (Fig. 16) moderately slender and straight; some body segments possibly with intrasegmental lines; middorsal tubercles absent; distinct lateral tubercles (i.e., “ventral lateral tubercles” of Odontostelis , Rozen, 1966) conspicuous on most segments. Integument with scattered setae (in contrast with integument of first instar of host which lacks setae) ; setae on anterior part of body longer than those on posterior part; on most abdominal segments setae situated on posterior part of segment dorsally, at apices of lateral tubercles, and widely scattered on venter; integument finely spiculate in numerous areas, including most of the tenth abdominal segment. Spiracles moderately large except for second pair which are distinctly smaller than others; atrium (Fig. 17) not projecting above body wall, with a peritreme, and slightly wider than deep; atrial wall apparently with indistinct ridges; primary tracheal opening apparently without collar. Tenth abdominal segment without large lobes or other modifications; anus slightly dorsal in position. material studied: One first-stage larva, 3 miles north of Apache, Cochise County, Arizona, April 30, 1966; from nest of Osmia nigrobarbata (J. G. Rozen and M. Favreau). December, 1967] Rozen: Immature Instars of Dioxys 243 Figs. 23-29. Dioxys pomonae pomonae Cockerell. 23. Right mandible of second instar, dorsal view. 24. Second instar, lateral view. 25. Head of second instar, lateral view. 26. Head of third instar, lateral view. 27, 28. Right mandible of third instar, dorsal and inner views. 29. Third instar, lateral view. Scales refer to Figs. 24 and 29, respectively. 244 New York Entomological Society [Vol. LXXV Figs. 30, 31. Pupa of Dioxys pomonae pomonae Cockerell, lateral and dorsal views. Second-Stage Larva of Dioxys pomonae pomonae Cockerell Figures 23-25 head: (Fig. 25) As described for first instar except for following: Posterior thickening of head capsule, hypostomal ridges, pleurostomal ridges, epistomal ridge, and longitudinal thickening of head capsule slightly more evident. Mandible (Fig. 23) not quite so slender and slightly shorter in relation to size of head. body: (Fig. 24) As described for first instar. material studied: One second-stage larva, 3 miles north of Apache, Cochise County, Arizona, April 30, 1966; from nest of Osmia nigrobarbata (J. G. Rozen and M. Favreau). Third-Stage Larva of Dioxys pomonae pomonae Cockerell Figures 26-29 head: (Fig. 26) As described for first instar except for following: Internal ridges of head capsule more distinct than those of second instar. Mandible (Figs. 27, 28) stouter than that of either first or second instar and shorter in relation to size of head. Both dorsal and ventral subapical inner edges faintly and indistinctly dentate. body: (Fig. 29) As described for first instar. material studied: One third-stage larva, 3 miles north of Apache, Cochise County, Arizona, May 4, 1966; from nest of Osmia nigrobarbata (J. G. Rozen and M. Favreau). December, 1967] Rozen: Immature Instars of Dioxys 245 Pupa of Dioxys pomonae pomonae Cockerell Figures 30-31 Length, 6.75 mm; body curved so that tip of tongue almost touching tip of metasoma. head: Scape and frons without tubercles; vertex without tubercles except for low mounds of ocelli; scattered small, unpigmented setae occurring mesiad of upper inner orbits but not above level of anterior ocellus; setae less abundant than those on head of Stelis bilineolata (Rozen, 1966). mesosoma: Lateral angles of pronotum somewhat produced; posterior lobes not produced; mesepisternum, mesoscutum, mesoscutellum, and axillae without tubercles and not produced; metanotum produced as distinct median rounded tubercle; slightly pigmented setae present on mesoscutum and mesoscutellum but not on axillae ; these setae fewer than those of Stelis bilineolata and somewhat longer than those of head; tegula not produced; wing without tubercles; fore tibia with apical tubercle; mid and hind tibia each with somewhat smaller apical tubercle ; other leg segments without distinct tubercles. metasoma: Terga I-VI with apical bands of short, unpigmented setae rising from minute tubercles; sterna without tubercles or setae; terminal spine absent. material studied : One live female pupa, 3 miles north of Apache, Cochise County, Arizona, larva collected May, 1966, pupated approximately September 1, 1966; from cell of Osmia nigrobarbata Cockerell (J. G. Rozen and M. Favreau). DISCUSSION The larvae of most nonparasitic bees superficially seem to change merely in size as they develop. Indeed, the four instars exist in the same environment, and their behavior, concerned primarily with feeding, is quite uniform. It would be surprising, therefore, if marked differences occurred from one instar to the next. A number of workers have noticed, however, changes with respect to the various tubercles on the postcephalic region in some groups. The tubercles seem to be associated with the feeding habits of the larva; the changes are presumably adaptations to the modifications in the shape, consistency, and size of the pollen mass as it is being consumed. Conspicuous changes also appear in the larvae of cocoon-spinning bees; such features as long palpi, projecting labiomaxillary region, and protruding salivary lips, that appear in the later instars are adaptations to cocoon spinning. More pronounced differences among instars have been noted with certain parasitic bees, such as the Nomadinae. The mode of parasitism in this group indicates that the first instar kills the egg or larva of the host and subsequent instars consume the pollen-nectar mixture. The first instar is equipped with a pigmented, more or less prognathous head capsule and greatly elongate, sickle- shaped mandibles with which it destroys the host’s offspring. The tip of the abdomen, at least in some cases, is modified into a pygopod-like structure enabling the larva to move about in search of its prey. Not only is the host’s egg or larva eliminated but also sibling larvae, for a female nomadine often lays more than one egg in a cell. The second and subsequent instars are much more 246 New York Entomological Society [Vol. LXXV “normal,” lacking most of the specialized modifications of the first stage. This pattern of parasitism seems to be the most common in the Apoidea and has arisen de novo a number of times. Another mode of parasitism occurs in the subgenus Odontostelis (Bennett, 1966) and apparently in Sphecodes ; the adult cuckoo bee removes the host egg or young larva before depositing her own egg, and the first instar hatches as a “normal” type. In Dioxys still another pattern of parasitism seems to be represented: the host’s offspring may be killed by the first, second or third instar of the cuckoo bee. Not only the first instar but also the second and third possess large sickle-shaped mandibles, and at least the first and second instars display an aggressive behavior when touched with forceps (Rozen and Favreau, 1967). None of the instars has an obvious pygopod-like structure for pushing itself around the cell or on the pollen mass. These facts suggest that the larva, be- cause of a slow mobility, may pass through several stages before it encounters and eliminates the host larva. Also, the egg of Dioxys is apparently inserted through the cell wall, probably after the cell is closed. Hence parasitism of a cell may take place over a considerable period of time. The first three instars of Dioxys are equipped to kill eggs of other Dioxys when and if they are in- troduced into an already parasitized cell. The ability of the intermediate instar to eliminate host and siblings may also be found in Coelioxys (Michener, 1953b) and in those Stelis which have apically simple mandibles in the last larval stage (Rozen, 1966). The changes that occur from one larval instar to the next in Dioxys pomonae pomonae involve the change in body size and form (Figs. 1, 16, 24, 29); the width of the head capsule of the four instars is as follows: first, 0.65 mm (one datum); second, 0.875 mm (one datum); third, 0.95 mm (one datum); fourth, 1.10-1.13 mm (three data). The antennae become relatively smaller with each instar though they are still large even in the fourth instar. The mandibles become shorter in relation to head size and the denticles on the upper and lower subapical edges first appear in the third stage. However, the dorsal apical tooth is a feature solely of the last instar as are the projecting enlarged labium, the division of the labium into a prementum and a postmentum, the protruding salivary lips, and the annulations of the spiracular subatria. The basal part of the maxilla is greatly enlarged in the first instar, a condition that holds for the second and third stages and persists to some extent in the last larval instar. The internal ridges of the head, including the stipites and the cardines, appear to become successively more pronounced with each instar. In other respects the larval instars of D. pomonae pomonae are remarkably similar. Even the setae which become shorter in relation to the body size, from instar to instar, maintain the same general distribution on the body through all instars. Indeed, the overall constancy of the external anatomical features is December, 1967] Rozen: Immature Instars of Dioxys 247 a more surprising result of the study than are the changes that take place in the development of the larva. Key to Some Genera of Cleptoparasitic Megachilidae, Based on the Mature Larvae Although this key is based only on the few species that I have examined, it may be of some value in separating the genera of megachilid cuckoo bees. Mature megachilid larvae, as a group, can be recognized because of the setae found on the postcephalic region; only the anthophorid genus Allodape and its relatives also bear conspicuous setae. 1. Mandible with more than four conspicuous setae on outer surface (Michener, 1953a, Figs. 160-161) ; gena, at least usually, produced into downward-pointing tubercle immediately behind posterior mandibular articulation (Michener, 1953b, Fig. 26) Coelioxys (two species) Mandible with at most one or two inconspicuous setae (Figs. 3, 5, 11, 13) ; gena with- out tubercle (Figs. 8, 15) 2 2. Body setae widely scattered and few; dorsal body setae restricted to caudal annulets on middle segments; vertex depressed medially; basal part of maxilla somewhat enlarged (Figs. 8, 15) Dioxys (two species) Body setae abundant; dorsal body setae numerous on both the caudal and cephalic annulets of middle segments; vertex not abnormally depressed medially; basal part of maxilla normal in size (Rozen, 1966, Figs. 5, 10) Stelis (three species) Literature Cited Bennett, F. D. 1966. Notes on the biology of Stelis ( Odontostelis ) bilineolata (Spinola), a parasite of Euglossa cordata (Linnaeus) (Hymenoptera: Apoidea: Megachilidae). Jour. New York Ent. Soc., 74: 72-79. Friese, H. 1925. Neue neotropische Bienenarten, zugleich II. Nachtrag zur Bienenfauna von Costa Rica (Hym.). Stettin, Ent. Ztg., 86: 1-41. Grandi, G. 1934. Contributi alia conoscenza degli imenotteri melliferi e predatori. XIII. Boll. 1st. Ent. Univ. Bologna, 7: 1-144. Hackwell, G. A. and W. P. Stephen. 1966. Eclosion and duration of larval develop- ment in the alkali bee, Nomia melanderi Cockerell (Hymenoptera: Apoidea). Pan- Pacific Ent., 42: 196-200. Hurd, P. D., Jr. 1958. American bees of the genus Dioxys Lepeletier and Serville (Hymenoptera: Megachilidae). Univ. California Publ. Ent., 14: 275-302. Jaycox, E. R. 1966. Observations on Dioxys productus productus (Cresson) as a parasite of Anthidium utahense Swenk (Hymenoptera: Megachilidae). Pan-Pacific Ent., 42: 18-20. Micheli, L. 1936. Note biologiche e morfologiche sugli imenotteri (VI Serie). Atti Soc. Italiana Sci. Nat. e Mus. Civ. Stor. Nat., 75: 5-16. Michener, C. D. 1944. Comparative external morphology, phylogeny, and a classification of the bees (Hymenoptera). Bull. Amer. Mus. Nat. Hist., 82: 151-326. -. 1953a. Comparative morphological and systematic studies of bee larvae with a key to the families of hymenopterous larvae. Univ. Kansas Sci. Bulk, 35: 987-1102. -. 1953b. The biology of a leafcutter bee ( Megachile brevis ) and its associates. Ibid., 35: 1659-1748. Rozen, J. G., Jr. 1964. The biology of Svastra obliqua obliqua (Say), with a taxonomic description of its larvae (Apoidea, Anthophoridae) . Amer. Mus. Novitates, no. 2170, pp. 1-13. 248 New York Entomological Society [Vol. LXXV . 1966. Taxonomic descriptions of the immature stages of the parasitic bee, Stelis ( Odontostelis ) bilineolata (Spinola) (Hymenoptera: Apoidea: Megachilidae) . Jour. New York Ent. Soc., 74: 92-94. . 1967. Review of the biology of panurgine bees with observations on North American forms (Hymenoptera, Andrenidae). Amer. Mus. Novitates, no. 2297, pp. 1-44. Rozen, J. G., Jr. and Marjorie S. Favreau. 1967. Biological notes on Dioxys pomonae pomonae and on its host, Osmia nigrobarbata (Hymenoptera, Megachilidae). Jour. New York Ent. Soc., LXXV(4): 197-203. Received for Publication June 13, 1967 Exchange Opportunities in Eastern Europe The National Academy of Sciences invites applications from American sci- entists who wish to visit Poland, Romania, and Yugoslavia for varying periods during the 1967-68 academic year. Through arrangements with the academies of these countries, the NAS will be able to select Americans for one-month survey visits or for research visits of from 3 to 12 months. Applicants for all programs must be U.S. citizens and have a doctoral degree or its equivalent in physical, biological, or behavioral sciences, mathematics, or engineering sciences. Applicants should specify which country they wish to visit since combined visits to two or more cannot be conveniently arranged. Participants will receive transportation to and from the foreign country. Those making research visits of 3 months or longer will receive grants to offset the loss of salary. Those making visits of 5 months or longer may also receive support for travel of dependents. Allowances from the receiving academy vary accord- ing to individual programs. Full information and applications may be obtained from the National Academy of Sciences, Office of the Foreign Secretary (USSR/ EE), Washington, D.C. 20418. Proceedings of the New York Entomological Society (Meetings held in Room 129 of the American Museum of Natural History unless otherwise indicated.) Meeting of April 4, 1967 Dr. Fredrickson presided; 23 members and 7 guests were present. Dr. Pinter of Harvard University, an expert on spiders, was introduced as a guest. program. Entomological Wanderings in Africa. Dr. Jerome Rozen, Chairman of the Department of Entomology at the Museum, described his recent trip to Africa where he searched for nests and immature stages of bees. The trip included short visits to Egypt and Nairobi, and a more extensive excursion in South Africa. He described the terrain in these areas, and commented on the flora, the fauna, and some points of interest along the way. His talk was illustrated with many colored slides. Howard R. Topoff, Sec. Meeting of April 18, 1967 President Fredrickson presided; 15 members and 4 guests were present. Dr. Alexander Klots reported for the Auditing Committee, stating that the records of the Society for 1966 were examined, and the accounts were found to be accurate and complete. The report was accepted, and the Committee was thanked. Dr. Michael Emsley of the Philadelphia Academy of Natural Science was proposed for active membership in the Society. Mrs. Betty Slater, wife of the speaker of the evening, was introduced. program. Zoogeography, Classification, and Evolution of the Chinch Bugs. Dr. James A. Slater, Chairman of the Department of Zoology and Entomology, University of Connecticut, gave a resume of the classification of chinch bugs and indicated their general importance. He spoke about the zoogeography and the evolution of these insects. He pointed out that there is a close relationship between the chinch bug fauna in South America and that in Africa; this opened a discussion of continental drift. Dr. Slater’s talk was illustrated with slides. Howard R. Topoff, Sec. Meeting of May 2, 1967 President Fredrickson called the meeting to order; 21 members and 7 guests were present. Dr. Michael Emsley of the Philadelphia Academy of Natural Sciences was elected to active membership, and Dr. Allen Benton of the State University College at Fredonia, New York was proposed for membership. program. Coding of Chemical Informtaion by Insects. Dr. Edward S. Hodgson of the Department of Zoology, Columbia University, illustrated his talk with slides. (An abstract follows.) Howard R. Topoff, Sec. CODING OF CHEMICAL INFORMATION BY INSECTS Recent developments in studies of the chemical senses of insects were described. The electron microscope has shown that sensory structures typically have pores which allow 249 250 New York Entomological Society [Vol. LXXV chemical stimuli direct access to receptor neurons. Electrical events in receptor excitation are best studied in sensilla on mouthparts of flies. Four taste receptor types have been identified: receptors for cations, anions, sugars, and water. Only the anion receptor mediates rejection responses under all conditions. Sensitivity of the chemoreceptors is affected by internal hormonal state as well as by external stimuli. E. S. Hodgson Meeting of May 16, 1967 The meeting was called to order by Vice-president David Miller in the absence of the Presi- dent; 16 members and 7 guests were present. Dr. Allen Benton of the State University College at Fredonia, New York was elected to membership. Dr. James Forbes, the Society’s delegate to the 11th Annual Conference of Biological Editors which was held at the Barbizon-Plaza Hotel, May 7-9, presented his report. Some of the problems and the topics considered by the Biological Editors this year were what constitutes primary publication, costs of printing journals, the use of key words in the titles of articles for properly designating their contents, and unreferred publications. He feels that the participation in these meetings over the past years has improved the quality of our Journal. He thanked the members for the oppor- tunity to represent them. program. Of Mice, Malaria, and Mosquitoes. Dr. Jerome Vanderberg of the Department of Preventative Medicine of the New York University Medical School illustrated his talk with slides. (An abstract follows.) Howard R. Topoff, Sec. OF MICE, MALARIA, AND MOSQUITOES Studies in the Department of Preventative Medicine during the past several years have been aimed at developing a model system of mammalian malaria which could be easily maintained and studied in the laboratory. The rodent malarial parasite, Plasmodium berghei, is a suitable organism in this regard, and the parasite can be transmitted through the mosquito, Anopheles stephensi , under controlled conditions. An important factor determining the success of this transmission is the temperature at which infected mosquitoes are kept. An inbred strain of mice from the Jackson Memorial Laboratory (Strain A/J) is a highly susceptible host for this parasite. By utilizing this system it has been possible to perform studies on basic physiology and morphogenesis of the malarial parasite, and in the applied area attempts have been made to develop a vaccine for malaria. J. Vanderberg December, 1967] Index to Volume LXXV 251 INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS VOLUME LXXV Generic names begin with capital letters. New genera, subspecies, and varieties are printed in italics. This index does not include the genera and subgenera of the Tortricidae and Phaloniidae, pp. 2-11; the aphids and their food plants, pp. 72-92; synonyms in American spiders, pp. 126-131. Acacia greggii, 133 Acamptopappus, 170 Acrolophus morus, 18 Adenostoma fasciculatum, 165 Aedes aegypti, 22 Aenictus, 107 Anopheles stephensi, 250 Anthidium emarginatum, 197 manicatum, 68 Anthophora, 236 Apomyelois bistriatella, 190 Aserica, 168 Astragalus, 197 Autoserica, 168 Biastes, 132 Blatella germanica, 148 Bombus, 69 Bombyx mori, 45, 119 Brachyspasta, 97 Brasilostreptus gracilis, 59 Brassica, 139 oleracea, 12 Calliopsis, 136 Calliphora erythrocephala, 119 Calospasta, 97 Capua lentiginosana, 34 Caryopteris clandonensis, 68 Catocala connubialis pulverulenta, 195 c. p. broweri, 195 micronympha, 195 m. gisela, 195 m. hero, 195 minuta, 195 Celtis, 193 Centris, 236 Chalicodoma muraria, 238 Chrysanthemum, 68 Cirsium lanceolatum, 139 Cochylis fernaldana, 34 Coelioxys, 236 Colias eurytheme, 12 philodice, 12 Conorhinopsylla stanfordi, 159 Cordylospasta, 97 Crambus bigelovi, 154 bolterellus, 158 cyrilellus, 158 harrisi, 154 leachellus, 158 praefectellus, 155 oslarellus, 155 Ctenopseustis flavicirrata, 34 Cysteodemus, 97 Daldinea, 193 Dioxys cincta, 203, 236 pacificus pacificus, 197 pomonae pomonae, 197, 236 productus productus?, 236 subruber, 197 Discoxenus, 204 Dorylus helvolus, 224 Drosophila, 20 melanogaster, 119 Dufourea dentipes, 132 malacothricis, 132 maura, 146 mulleri, 132 pulchricornis, 132 spinifera, 146 trochantera, 132 Eciton, 107 burchelli, 101 hamatum, 102 Epagoge schausiana, 34 spadicea, 34 vinolenta, 34 252 New York Entomological Society [Vol. LXXV Ephestia kiihniella, 119 Lythrum salicaria, 68 Epicordulia princeps, 179 Lytta, 97 regina, 179 Euaspis, 236 Macrotermes, 209 Euglossa, 236 Malacothrix, 133, 197 Eupompha, 97 Maladera castanea, 167 Eurvstylops, 138 Megachile, 236 Megetra, 97 Gaillardia, 13, 197 Melanoplus differentialis, 45 Galleria mellonella, 105 Meloe, 97 Glaucomys sabrinus, 159 Mentha, 68 volans, 159 Microtus pennsylvanicus, 159 Gnophomyia (Gnophomyia) diacaena, 183 Monopsyllus vison, 31 eupetes, 184 Musca autumnalis, 119 Gonepityche pacaraimae, 56 Myrmecia pyriformis, 35 Gonomyia (Lipophleps) pentacantha, 183 tarsata, 35 nissoriana, 184 vindex, 35 Gvmnastes (Gymnastes) anticaniger, 24 cyaneus, 26 Nanostreptus, 56 nUgiricus , 24 Negalius, 97 latifusciis, 24 Neivamyrmex, 106 ornatipennis, 28 Neopasites, 201 tridens, 24 (Micropasites) cressoni, 132 Gynaecomeloe, 97 (Neopasites) fulviventris, 13 Nepytia janetae , 74 Heliconious erato, 109 regulata, 76 melpomene, 109 Nomadopsis, 134 Heptathela bristowei , 114 Nomia melanderi, 236 kimurai, 114 sinensis, 114 Oenothera, 139 Heterocampa pulverea, 62 Odontostelis, 236 umbrata, 63 Odontotermes, 204 Holcopasites, 143, 201 angustatus, 223 Hyalophora gloveri, 105 badius, 215 Hvmenolepsis diminuta, 19 ceylonicus, 215 nana, 21 culturarum, 205 Hypoxylon occidentale, 190 hainanensis, 218 thouarsianum, 190 interveniens, 219 latericius, 222 Juniperus pachyphloea, 158 montanus, 206 nolaensis, 211 Lapara, 44 obesus, 217 Larix, 43 obscuriceps, 211 Lepidium, 132 patruus, 215 Lesquerella, 133 redemanni, 215 gordoni, 142 taprobanes, 205 Liphistius, 114 transvaalensis, 206 malayanus, 115 vulgaris, 222 schensiensis, 114 wallonensis, 217 sinensis schensiensis, 114 Oreopasites, 143, 201 December, 19671 Index to Volume LXXV 253 Osmia lignaria, 108 nigrobarbata, 197, 240 Panthea furcilla, 43 Parevaspis, 236 Pelargonium, 69 Penstemon, 139 Perdita sexmaculata, 138 Peromyscopsylla h. hamifer, 159 Phacelia, 197 popei arizonica, 133 leucophila, 139 Philosamia cynthia, 105 Phodaga, 97 Pieris rapae, 12 Piersea, 193 Pinus banksiana, 44 resinosa, 44 rigida, 44 scopulorum, 158 strobus, 43 Plasmodium berghei, 250 gallinaceum, 22 Platysamia cecropia, 119 Pleurospasta, 97 Popillia japonica, 45, 119 Populus, 193 Potentilla, 68 Prosopis, 133 Protoparce sexta, 105 Pseudomeloe miniaceomaculata, 93 Pteroptyx, 104 Pyrota, 97 Quercus agrifolia, 190 coccinea, 62 Rhizoctonium, 138 Rophites canus, 132 hartmanni, 132 quinquespinosus, 132 Salix, 139 Salvia farinacea, 68 splendens, 68 Sciaphila indivisana, 34 Semiothisa, 44 Serica adversa , 161 alabama , 161 alleni, 161 anthracina, 161 atracapilla, 161 atricapilla, 161 aspera, 171 atratula monita, 171 aviceps, 161 barri, 161 blatchleyi, 163 bruneri, 161 caliginosa, 167 Carolina, 171 castanea, 161 contorta, 171 diablo, 161 elusa, 163 ensenada, 161 errans, 166 fimbriata, 161 floirdana, 161 frosti, 161 heteracantha , 161 howdeni, 161 imitans, 1 7 1 joaquinella, 161 laguna, 171 mackenziei, 161 mendota, 161 michelbacheri, 161 oliveri, 161 peregrina, 161 perigonia eremicola, 161 pilifera, 161 porcula, 161 prava, 171 pruinipennis, 161 pullata, 161 rossi, 161 searli, 161 sericea, 161 sericeoides, 161 sculptilis, 161 solita, 167 stygia, 161 texana, 1 7 1 tristis, 163 trociformis blatchleyi, 161 vespertina accola, 171 watsoni, 171 Solidago, 69 Sphaeralcea, 139 254 New York Entomological Society [Vol. LXXV Sphecodes, 246 Stelis, 236 bilineolata, 245 Svastra obliqua obliqua, 236 Synaptomys cooperi, 159 Systropha curvicornis, 132 planidens, 132 punjabensis, 132 Tortrix baboquavariana, 34 biocellata, 34 desmatana, 34 triplagata, 34 Toxorhina (Ceratocheilus) bistyla , 183 capnitis, 186 fulvicolor, 183 juscolimbata, 183 simplicistyla, 183 Tamiasciurus hudsonicus, 31, 159 Tegrodera, 97 Tenebrio molitor, 20, 45, 119 Termitodiscus angolae, 207 braunsi, 206 butteli, 211 coatoni, 204 emersoni, 204 escherichi, 206 heimi, 210 krishnai , 204 later icus , 204 machadoi, 207 minutus, 210 sheasbyi, 204 splendidus, 211 transvaalensis, 209 vansomereni, 204 vicinior, 217 Termitogerrus, 204 Tetralonia, 132 Tetraonyx, 97 luteibasis, 185 mesorhyncha, 185 monostyla, 185 tuberifera, 185 Trachusa, 238 Trentepohlia (Mongoma) amphinipha, 24 flava, 25 horiana, 25 patens, 24 subtenera, 25 (Trentepohlia) bellipennis, 26 camillerii, 26 injernalis, 24 ornatipennis, 26 Tribolium confusum, 19 Trypargilum, 108 Urechis caupo, 45 Urostreptus, 56 Vicia cracca, 13 Xylocopa, 71 INVITATION TO MEMBERSHIP The New York Entomological Society was founded 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