Americana A Journal of Entomology. Volume XXX (Niew Series) PUBLICATION COMMITTEE JOSEPH C. BEQUAERT, EDITOR GEORGE S. TULLOCH EDWIN WAY TEALE PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY ENTOMOLOGICA AMERICANA VOL. XXX (N. S.), 1950 CONTENTS PAGE The Scutate Ticks, or Ixodidae, of Indonesia. George Anastos. August 14, 1950 1 DIV. INS.' U.S. NATL. HUS 7 Nos. 1 to 4 VOL. XXX (New Series) A Journal of Entomology. PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY PUBLICATION COMMITTEE JOSEPH C. BEQUAERT, Editor GEORGE S. TULLOCH E. W. TEALE Published for the Society by the Business Press Inc. N. Queen St. and McGovern Ave., Lancaster, Pa. Subscription, $5.00 per year Date of issue, August 14, 1950 Vol. XXX Americana Nos. 1-A THE SCUTATE TICKS, OR IXODIDAE, OF INDONESIA1 By George Anastos Department of Zoology, Miami University, Oxford, Ohio Contents Page Introduction 2 Morphology 4 General Range of Species 6 Distribution According to Islands 7 Distribution According to Hosts 9 Key to Genera 14 Genus Ixodes Latreille, 1795 16 Genus Ilaemaphysalis C. L. Koch, 1844 23 Genus Dermacentor C. L. Koch, 1844 51 Genus Rhipicephalus C. L. Koch, 1844 59 Genus Boophilus Curtice, 1891 71 Genus Amblyomma C. L. Koch, 1844 77 Genus Aponomma Neumann, 1899 120 Species of Doubtful Occurrence in the East Indies 133 Index 135 1 The material in this paper was included in a thesis, submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, at Harvard University. 1 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Introduction This work is a revisional study of the ticks of the family Ixodidae occurring in Indonesia (former Netherlands East Indies). It is based upon the collections of museums in the United States and upon the material sent to Dr. J. C. Bequaert in 1940 by Professor F. C. Kraneveld, at that time at the Veeartsenij kundig Instituut, Buitenzorg, Java, now at the Rijksuniversiteit, Utrecht, Netherland. Material was also received from Dr. E. de Boer of the Veterinary Institute of the Department of Agriculture and Fisheries at Buiten- zorg, Java. The East Indian tick collection of the Museum of Com- parative Zoology, Cambridge, Massachusetts, is the largest studied, and it contains twenty species. The Rocky Mountain Laboratory, Hamilton, Montana, has an equally fine collection containing eighteen species. The collection of the American Museum of Nat- ural History, New York, is small and contains only three East Indian species. The collection of the Chicago Natural History Museum has only one species represented. The material received from Dr. Kraneveld, hereafter referred to as the Kraneveld col- lection, consists of approximately twenty thousand specimens with twelve species represented. I also saw two species sent to the Rocky Mountain Laboratory by the late Dr. P. Schulze of Rostock, Germany. The collection of Dr. de Boer contained only one species of interest. The synonymy is extralimital in all cases, except for the species world-wide in distribution, such as RJiipicepkalus sanguineus and to a lesser extent Boopkilus microplus. The synonymy of previous workers is critically examined, and a discussion of the new synonymy is included in the text under each species. In the spelling of place names, I follow the National Geographic Society, Map of Southeast Asia and Pacific Islands, 1944. Names not located on this map are taken from Stielers Hand Atlas, 1930- 1931, and the Century Atlas, 1901. In this work I take a broad view of a “species”, and I sink many of the purely geographical subspecies described by Schulze. For example, the characters used by Schulze to separate four subspecies of Amklyomma testudinarium are color, pattern of ornamentation and size ; all are variable and have little, if any, taxonomic value. The primary reason for his subspecies appears to be geographical. Since these ticks were introduced into many areas throughout the Oriental Region in recent times, and since they are still being spread by livestock, there remains no basis for recognizing these sub- 2 ENTOMOLOGICA AMERICANA species. I examined approximately one hundred specimens of A. testudinarium from numerous localities in the Oriental Region, and since I can find no reliable characters by which they can be sepa- rated, I regard them all as one species. In the case of Rhipicephalns haemaphysaloides, where reliable morphological characters can be correlated with geographical distribution, there is a basis for recog- nizing subspecies. I also take a broad view regarding genera, and I believe that they should be subdivided only when absolutely neces- sary. The history of acarology shows clearly that great care should be exercised in the erection of new genera or higher groups. The arrangement in the list of reptilian hosts follows Loveridge (1945, Reptiles of the Pacific World), and the names are according to the Catalogue of Reptiles in the Museum of Comparative Zoology. In the list of avian hosts, which is very small, the arrangement is that of Chasen (1935, Bull. Raffles Mus., 11, pp. 1-389). The ar- rangement and the names in the list of mammalian hosts are for the most part taken from Chasen (1940, Op. cit., 15, pp. 1-209), with the rodent names based upon Ellerman (1940, The Families and Genera of Living Rodents , Vol. 1; 1941, Yol. 2). The drawings are made from alcoholic specimens, with the aid of a net reticule placed in the ocular of a binocular-dissecting microscope. Magnifications ranging from twenty-five to one hun- dred times are most suitable for study. Detail is more clearly seen when the specimen is viewed dry, and drying will also bring out the pattern of the scutal ornamentation. The keys to genera and species are artificial and based upon several characters to make identification more certain. Future studies based on breeding experiments may change existing views on the characters regarded as diagnostic, but in these keys an at- tempt is made to select those characters which are felt to be reliable and most diagnostic. In the bibliographies the abbreviations n and 1 stand for nymph and larva respectively. Acknowledgments. To Dr. J. C. Bequaert, Curator of Insects, Museum of Comparative Zoology, I owe a great deal for what has gone into this study ; it was at his suggestion that I began this work and by his constant guidance that I was able to reach a better under- standing of the ticks. Dr. F. M. Carpenter, Professor of Zoology, Harvard University, has given me every encouragement and has been most helpful with aid and advice relative to the many problems in morphology and taxonomy. To Mr. G. M. Kohls of the Rocky 3 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Mountain Laboratory, Hamilton, Montana, I am greatly indebted for his kindness in rendering me every assistance and for making my summer at the laboratory such a pleasant experience. Through the kindness of the late Dr. R. R. Parker, Director of the Rocky Mountain Laboratory, I was able to study their large, excellent col- lection and to bring material to Harvard for further study. To my wife Angelica I owe the greatest debt of gratitude for her understanding kindness and assistance during the preparation of this work. The publication of this work in its present form was made pos- sible by financial assistance from two sources : the Miami University Alumni Association, Oxford, Ohio ; and the Society of the Sigma Xi. These grants-in-aid, herewith gratefully acknowledged, defrayed the major part of the cost of reproduction and printing of the illustrations. Morphology The identification of genera in the ticks is fairly simple, and in many cases specimens can be sorted to genus without need of magnification. The identification of species is more difficult, for within a genus the species are often uniformly monotonous in struc- ture and lack good characters for separation. Even the very few characters present vary within limits. This range of variation is not known for most ticks, and it can only be determined by con- trolled breeding experiments wherein the F-2 generation of selected extreme females from an P-1 generation of one female is carefully studied. The developmental stages and the behavior patterns are known to vary in those ticks having wide distribution, and these factors must also be taken into consideration when defining the limits of a species. Geographical distribution alone can not be used in separating species, for many species of ticks have been trans- ported into new localities with domestic animals in fairly recent times. Since all the East Indian specimens examined were preserved, it was necessary to rely upon external, morphological characters exclu- sively. Data regarding biology and host relationships have been helpful to a certain degree. The dissections I made of numerous living ticks of other species, in an attempt to find characters which might be helpful in separating these forms, proved inconclusive. As with the external features, the internal anatomy appears to be very uniform. A study of the mouthparts also failed to show any- 4 ENTOMOLOGICA AMERICANA CAPITULUM TROCHANTER COXA I EXTERNAL SPUR INTERNAL SPUR coxa n GENITAL OPENING COXA IH GENITAL GROOVE COXA 12 SPIRACLE ANAL OPENING ANAL GROOVE ADANAL PLATE ANAL PLATE FESTOON Fig. 1. External morphology of Bhipicephalus haerriaphysa- loides Supino: A, capitulum of female in dorsal view; B, same in ventral view; C, male in dorsal view; D, spiracle of female; E, female in dorsal view ; F, male in ventral view. 5 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 thing significant; a few differences noticed do not warrant the destruction of valuable specimens. Not too much value should be attached to the pattern of the scutal ornamentation, for it varies considerably wherever present. For a detailed treatment of tick morphology in relation to tax- onomy and for definitions of terms used, the reader is referred to Nuttall, Warburton, Cooper and Robinson, 1911-1926, Ticks: A Monograph of the Ixodoidea. Most of the descriptive terms used in the present work are explained in Fig. 1. General Range of Species The ticks of Indonesia are still far from being completely known and extensive collecting remains to be done on many islands in this area. Excluding from consideration those ticks found on domestic animals, certain generalizations can be drawn from the known reli- able records. Those species occurring on the islands near Asia are generally identical with the mainland fauna. Proceeding eastward along the East Indies, the species common to Asia become fewer and fewer, and nearly all are absent from New Guinea which has a fauna nearly all its own. A very few species occurring on New Guinea, however, are found in the more western East Indies. Sumatra, Java and their adjacent islands have the greatest num- ber of species. Though their faunas are similar and chiefly of Asiatic origin, many species are restricted to this group. The Lesser Sunda Islands, from Bali to Banda, have a smaller represen- tation of species than the preceding group ; a few species are re- stricted to this region, whereas others show affinities to the Sumatra- Java fauna or to the New Guinea fauna. Nothing definite can be said for the Celebes and the Halmahera groups at this time; with the exception of one species, Amblyomma babirussae, known only from Celebes, the rest are common ticks of domestic animals intro- duced into this area. Several species have found domestic animals suitable hosts, and the habits of domestic animals have favored their increase in num- bers and their distribution throughout the islands. In this group belong Amblyomma testudinarium, Boophilus microplus, Haema- physalis bispinosa, H. cornigera, H. hystricis, H. papuana, H. wel- lingtoni, Rhipicephalus haemaphysaloides, R. sanguineus and Der- macentor auratus. Several species, taken off wild animals primarily, are definitely of Asiatic origin and have advanced eastward into Indonesia : Am- 6 ENTOMOLOGICA AMERICANA blyomma crenatum, A. helvolum, A. javanense, A. malayanum, Aponomma lucasi, Haemaphysalis honing sb ergeri and Ixodes gra- mdatus. The following species are restricted to this territory and are un- known from other localities in the Oriental Region: Amblyomma babirussae, A. geoemydae, A. robinsoni, Aponomma barbouri, A. homodoense , Haemaphysalis hylobatis, II. toxopei, H. tragidi, Ixo- des praematurus and I. spinicoxalis. Within this group Ambly- omma robinsoni and Aponomma homodoense are known only from Komodo Island. Amblyomma cyprium and Aponomma trimaculatum appear to be of New Guinea origin and for the present are placed in this group. Amblyomma eordiferum has a peculiar distribution being known from three widely separated localities, Siam, Krakatau Is- land and Banda Island. Future records may fill the gaps in this distribution. Distribution According to Islands2 Simeuloee: Amblyomma helvolum • Aponomma trimaculatum ; Hae- maphysalis honing sb ergeri. Babi : Amblyomma helvolum • Aponomma lucasi (?) ; Haemaphy- salis hystricis. Nias : Boophilus microplus. Siberoet : Haemaphysalis hystricis. Sumatra : Amblyomma crenatum • A. geoemydae ; A. helvolum ; A. javanense ; A. malayanum ; A. testudinarium ; Aponomma lu- casi; A. trimaculatum ; Boophilus microplus ; Dermacentor au- ratus; Haemaphysalis bispinosa ; II. cornigera ; H. hylobatis ; II. hystricis ; H. honing sb ergeri; H. papuana ; H. Wellington i; Ixodes granulatus ; I. spinicoxalis ; Rhipicephalus haemaphy- saloides haemaphysaloides ; R. haemaphysaloides pilans; R. sanguineus. Berhala : Amblyomma helvolum ; Haemaphysalis tragidi. Pandang : Amblyomma testudinarium. Bengkalis : Haemaphysalis hystricis. Riouw: Haemaphysalis hystricis. Bintan : Amblyomma testudinarium. Krakatau : Amblyomma eordiferum. 2 Excludes Borneo, Aroe Islands, Amboina, Kai Islands and Netherlands New Guinea. 7 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Java: Amblyomma crenatum ; A. helvolum; A. javanense ; A. testu- dinarium; Aponomma barbouri ; A. lucasi; Boophilus micro- plus; Dermacentor auratus; Haemaphy satis bispinosa; H. cornigera; H. koningsbergeri; II. papuana ; H. traguli ; H. wellingtoni; Ixodes granulatus ; I. spinicoxalis ; Rhipicephalus haemaphy saloides pilaus ; R. sanguineus. Madoera : Boophilus microplus ; Rhipicephalus haemaphy saloides pilans; R. sanguineus. Karimoendjowo : Haemaphy satis cornigera ; H. hystricis ; H. bispi- nosa; Boophilus microplus. Bawean : Haemaphy satis koningsbergeri; Ixodes granulatus. Bali : Amblyomma testudinarium ; Boophilus microplus ; Haemaphy- salis bispinosa; H. cornigera ; H. hystricis; Rhipicephalus hae- maphysaloides pilans ; R. sanguineus. Lombok : Rhipicephalus haemaphy saloides pilans; Boophilus micro- plus; Amblyomma testudinarium. Soembawa : Haemaphy satis bispinosa ; H. cornigera ; II. papuana; Rhipicephalus haemaphy saloides pilans ; R. sanguineus. Komodo : Amblyomma helvolum; A. robinsoni; Aponomma komodo- ense. Flores : Amblyomma cyprium cyprium; A. helvolum; A. te'studina - rium; Boophilus microplus ; Haemaphysalis bispinosa ; Rhipi- cephalus haemaphy saloides pilans. Alor : Haemaphysalis papuana; Rhipicephalus haemaphy saloides pi- lans; R. sanguineus. Banda : Amblyomma cordiferum. Soemba: Boophilus microplus ; Haemaphysalis cornigera ; Rhipi- cephalus haemaphy saloides pilans ; R. sanguineus ; Amblyomma testudinarium. Sawoe : Rhipicephalus haemaphy saloides pilans. Timor: Boophilus microplus ; Haemaphysalis papuana; Ixodes praematurus ; Rhipicephalus haemaphy saloides pilans; R. san- guineus. Roti : Rhipicephalus haemaphy saloides pilans. Tanimbar : Amblyomma helvolum; Boophilus microplus. Boeroe : Amblyomma babirussae (?) ; Haemaphysalis toxopei. Boo : Amblyomma testudinarium. Saparoea : Boophilus microplus ; Haemaphysalis papuana; Rhipi- cephalus sanguineus. Soela : Boophilus microplus ; Haemaphysalis bispinosa ; H. papuana. Ternate : Amblyomma cordiferum (?) ; Aponomma lucasi (?). 8 ENTOMOLOGICA AMERICANA Halmahera : Haemaphysalis papuana ; Boophilus microplus. Celebes : Amblyomma babirussae ; A. cyprium cyprium ; A. testu- dinarium; Boophilus microplus ; Haemaphysalis bispinosa ; H. hystricis ; H. papuana ; Rhipicephalus haemaphysaloides pi- laus; R . sanguineus. Sangihe : Haemaphysalis papuana ; Boophilus microplus. Boetoeng : Boophilus microplus. Distribution According to Hosts 'Reptilia Order Testudinata Family Emydidae Geoemyda spinosa (Gray), Spiny Hill-tortoise. — Amblyomma ge- oemydae. Order Sqnamata Suborder Sauria, or Lacertilia Family Scincidae Mabuya multifasciata (Kuhl), Many-striped Skink. — Amblyomma helvolum. Family Varanidae Varanus komodoensis Ouwens, Komodo Land-monitor. — Ambly- omma robinsoni, A. helvolum, Aponomma komodoense. Varanus Salvator (Laurenti), Common Water-monitor. — Ambly- omma helvolum , A. javanense, Aponomma lucasi, A. trimacula- tum. Varanus indicus (Daudin), Indies Water-monitor. — Amblyomma helvolum. Suborder Serpentes, or Ophidia Family Boidae Python reticulatus Schneider, Reticulated Python. — Amblyomma helvolum, Aponomma barbouri. Python sp., Python. — Amblyomma cordiferum. Family Colubridae Ptyas korros (Schlegel), Rat-snake. — Amblyomma helvolum. Ptyas mucosus (Linnaeus), Greater Rat-snake. — Amblyomma hel- volum. Boiga dendrophila (Boie), Mangrove Snake. — Amblyomma helvo- lum. 9 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Coluber oxycephalus Boie, Ular selenseng (Malayan name). — Am- blyomma helvolum. Family Elapidae Ophiophagus hannah (Cantor), King Cobra, or Hamadryad. — Am- blyomma helvolum. Naja naja leucodira Boulenger, Hooded Cobra. — Amblyomma hel- volum. Aves Family Timaliidae Babbler. — Ixodes praematurus. Family Tnrdidae Geokichla sp., Ground Thrush. — Ixodes praematurus. Family Zosteropidae Zosterops sp., White-eyes. — Ixodes praematurus. Family Phasianidae Gallus g allies (Linneaus), Domestic chicken. — Haemaphy salts Wel- lingtoni. Family Anatidae Anser anser (Linnaeus), Domestic goose. — Haemaphy salts welling- toni. Mammalia Order Insectivora Family Tupaiidae Tupaia gits hypochrysa Thomas, Common Tree Shrew. — Ixodes granulatus. Tupaia javanica occidentals Robinson and Kloss, Common Tree Shrew. — Ixodes granulatus, I. spinicoxalis. Tupaia tana tana Raffles, Large Tree Shrew. — Haemaphy satis cor - nig era (.?). Family Erinaceidae Hylomys suillus suillus Muller, Short-tailed Shrew, or Lesser Gym- nure. — Ixodes granulatus. Family Soricidae Crocidura orientalis Jentink, White-toothed Shrew. — Ixodes granu- latus, I. spinicoxalis. Order Chiroptera Bat. — Haemaphy satis toxopei. 10 ENTOMOLOGICA AMERICANA Order Primates * Family Hominidae Homo sapiens Linnaeus. — Amblyomma testudinarium, Rhipicepha- lus sanguineus. Family Hylobatidae Hylobates syndactylus syndactylus (Raffles), Siamang. — Haema- phy satis hy lob at is. Family Lorisidae Nycticebus coucang (Boddaert), Slow Loris. — Haemaphy satis ho- ning sb erg eri. Order Carnivora Family Mnstelidae Martes flayigula henrici (Schinz), Yellow- throated Marten. — Am- blyomma testudinarium, Haemaphysalis honing sb erg eri, Ixodes spinicoxalis. Helictis orientalis (Horsfield), Ferret Badger —Haemaphysalis ho- ning sb erg eri. Arctonyx collaris hoevenii (Hubrecht), Hog-nosed Badger. — Hae- maphysalis hystricis. Mydaus javanensis j.avanensis (Leschenanlt) , Teledu. — Haemaphy- salis honing sb erg eri, Ixodes spinicoxalis. Family Canidae Canis familiaris Linnaeus, Domestic Dog. — Boophilus microplus, Haemaphysalis bispinosa, H. hystricis, H. papuana, Rhipice- phalus haemaphy saloides haemaphy saloides, R. haemaphy saloi- des pilans, R. sanguineus. Family Yiverridae Paradoxurus hermaphroditus javanicus (Horsfield), Palm Civet, or Musang. — Haemaphysalis honing §b erg eri, Ixodes spinicoxa- lis. Family Felidae Felis tigris Linnaeus, Tiger. — Haemaphysalis honing sb erg eri, Rhi- picephalus haemaphy saloides haemaphy saloides, R. haemaphy- saloides pilans. Felis pardus Linnaeus, Leopard, or Panther. — Haemaphysalis ho- ningsbergeri. Felis bengalensis javanensis Desmarest, Leopard Cat. — Haemaphy- salis honing sb erg eri. 11 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Order Edentata, or Pholidota Family Manidae Manis javanica Desmarest, Pangolin, or Scaly Anteater. — Ambly- omma javanense. Order Rodentia Family Sciuridae Petaurista petaurista petaurista (Pallas), Large Red Flying Squir- rel, or Giant Flying Squirrel. — Haemaphysalis honing sb erg eri. Batufa bicolor bicolor (Sparrmann), Black Giant Squirrel. — Hae- maphysalis honing sb erg eri. Callosciurus notatus notatus (Boddaert), Common Malay Squirrel, or Plantain Squirrel. — Ixodes granulatus. Callosciurus notatus madurae (Thomas), Common Malay Squirrel, or Plantain Squirrel. — Ixodes granulatus. Callosciurus nigrovittatus nigrovittatus (Horsfield), Black-striped Squirrel. — Ixodes granulatus, I. spinicoxalis. Lariscus insignis javanus (Thomas and Wroughton), Striped Ground Squirrel. — Ixodes granulatus. Family Muridae Battus rattus rattus (Linnaeus), Black Rat.— Ixodes granulatus, Haemaphysalis honing sbergeri. Battus rattus diardi (Jentink), Malaysian House Rat. — Ixodes gra- nulatus. Battus rattus brevicaudatus Horst and de Raadt, Ricefield Rat, or Sawah Rat. — Ixodes granulatus. Battus rattus jalorensis (Bonhote), Malaysian Field Rat. — Ixodes granulatus. Battus concolor ephippium (Jentink), Little Burmese Rat. — Ixodes granulatus. Battus maxi Sody, Giant Rat. — Ixodes granulatus. Battus lepturus (Jentink), Long-tailed Rat. — Ixodes granulatus, I. spinicoxalis. Battus fulvescens buhit (Bonhote), Bonhote Rat. — Ixodes spini- coxalis. Battus fulvescens treubi Robinson and Kloss, Bonhote Rat. — Ixodes granulatus. Battus fulvescens temminchi Kloss, Bonhote Rat. — Ixodes granula- tus. Battus bartelsi (Jentink), Bartels Rat. — Ixodes granulatus. 12 ENTOMOLOGICA AMERICANA Pithecheir melanurus melanurus Cuvier, Red Tree Rat. — Ixodes spinicoxalis. Order Perissodactyla ' Family Rhinocerotidae Rhinoceros sondaicus Desmaresf, Javan, or One-horned Rhinoceros. — Amblyomma testudinarium. Family Tapiridae Tapirus indicus Desmarest, Malay' Tapir. — Amblyomma testudina- rium. Family hlquidae Equus caballus Linnaeus, Domestic Horse. — Amblyomma babirus- sae, Boophilus microplus, Haemaphysalis bispinosa, H. hystri- cis, H. papuana, Rhipicephalus haemaphysaloides pilans. Order Artiodactyla Family Suidae Bus cristatus cristatus Wagner, Wild Pig. — Amblyomma testudina- rium, Rhipicephalus haemaphysaloides pilans. Bus cristatus vittatus Boie, Wild Pig. — Amblyomma testudinarium, Rhipicephalus haemaphysaloides pilans. Bus verrucosus Muller and Schlegel, Java Pig. — Amblyomma testu- dinarium. Bus barbatus Muller, Bearded Pig. — Amblyomma testudinarium. Bus celebensis Muller and Schlegel, Celebes Pig. — Amblyomma ba- birussae, A. cyprium, Haemaphysalis hystricis. Bus scrofa Linnaeus, Domestic Pig.— -Amblyomma testudinarium, Boophilus microplus, Dermacentor auratus, Haemaphysalis bi- spinosa, H. hystricis, H. papuana, Rhipicephalus haemaphysa- loides haemaphysaloides. Babirussa babyrussa (Linnaeus), Babirusa. — Amblyomma babi- russae. Family Tragulidae Tragulus kanchil kanchil (Raffles), Smaller Mouse Deer, or Pelan- dok. — Haemaphysalis traguli. Family Cervidae Cervus unicolor equinus Cuvier, Sandbar, or Rusa. — Boophilus mi- croplus. Cervus unicolor russa Muller and Schlegel, Sambar, or Rusa. — Bo- ophilus microplus. 13 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Cervus sp. — Amblyomma testudinarium. Deer. — Boophilus microplus, Haemaphy sails bispinosa. Family Bovidae Bos sondaicus sondaicus Schlegel and Muller, Banteng, or Wild Ox. — Amblyomma testudinarium, Haemaphy sails cornigera, H. pa- puana. Bos bubalis bubalis Linnaeus, Indian, or Water Buffalo. — Ambly- omma babirussae, A. cyprium cyprium, A. testudinarium, Bo- ophilus microplus, Haemaphysalis hystricis, H. papuana, H. wellingtoni, Bhipicephalus haemaphy saloides haemaphy saloi- des, R. haemaphy saloides pilans, R. sanguineus. Bos taurus Linnaeus, Domestic Cattle. — Amblyomma babirussae, A. testudinarium, Boophilus microplus, Haemaphysalis bispinosa, H. cornigera, H. hystricis, H. papuana, Rhipicephalus haema- physaloides haemaphy saloides, R. haemaphy saloides pilans, R. sanguineus. Capra hircus Linnaeus, Domestic Goat. — Amblyomma testudina- rium, Boophilus microplus, Haemaphysalis bispinosa, Rhipi- cephalus haemaphy saloides haemaphy saloides, R. haemaphysa- loides pilans, R. sanguineus. Ovis aries Linnaeus, Domestic Sheep. — Haemaphysalis bispinosa, Rhipicephalus haemaphy saloides pilans. Key to Genera Males 1. Ventral side mostly covered by seven hardened, non-salient plates, the anal plate limited by a distinct groove curved in front of the anus and extended to the hind margin. Scutum in- ornate, without festoons. Palps elongate. No eyes. Ixodes Posterior groove on ventral side either curved behind the anus or indistinct. Ventral side either without extensive plates or partly covered by salient plates 2 2. Eyes absent. Scutum with festoons. Ventral side without plates , „. 3 Eyes present 4 3. Palps short and broad, at most as long as basis capituli ; second segment more or less angular at sides. Scutum inornate. Haemaphysalis 14 ENTOMOLOGICA AMERICANA Palps long and slender; second segment not angular at sides. Scutum ornate or inornate Aponomma A Palps long and slender; second segment not angular at sides. Ventral side without extensive or salient plates. Basis ca- pituli not angular at sides. Scutum with festoons, ornate or inornate Amblyomma Palps short and broad, at most as long as basis capituli 5 5. Basis capituli rectangular, not angular at sides. Ventral side without salient plates. Scutum with festoons, ornate in East Indian species Dermacentor Basis capituli hexagonal, with angular sides. Ventral side partly covered by four plates, often salient at the hind margins. Scutum inornate 6 6. Scutum with festoons. Post-anal groove distinct. Coxae I with two long spurs Bhipicephalus Scutum without festoons. Post-anal groove indistinct. Coxae I with two very short spurs Boophilus Females 1. Ventral side posteriorly with a distinct groove curved in front of the anus. Scutum inornate. Palps elongate. No eyes. Ixodes Ventral side posteriorly with a groove curved behind the anus or the groove indistinct 2 2. Eyes absent 3 Eyes present 4 3. Palps short and broad, at most as long as basis capituli ; second segment angular or produced at sides. Scutum inornate. Haemaphy salis Palps long and slender; second segment not angular at sides. Scutum ornate or inornate Aponomma 4. Palps long and slender. Basis capituli not angular at sides. Scutum ornate or inornate Amblyomma Palps short and broad 5 5. Basis capituli rectangular, not angular at sides. Scutum ornate in East Indian species Dermacentor Basis capituli hexagonal, with angular sides. Scutum in- ornate 6 6. Post-anal groove distinct. Second and third segments of palps not unusually short and not angular at outer base. Coxae I with two long spurs Rhipicephalus 15 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Post-anal groove indistinct. Second and third segments of palps very short, slightly angular at outer base. Coxae I with two very short, blunt spurs Boophilus Genus Ixodes Latreille, 1795 Oudemans (1925, Ent. Ber. Nederl. Ent. Ver., 7, pt. 146, p. 32) described a single male from Amboina, Moluccas, as E sckatocephalus ropsteini. Schulze (1935, Zeitschr. Morph. Oekol. Tiere, 30, pt. 1, pp. 27, 28, fig. 28) corrected the name to kopsteini, regarded it as an Ixodes and created the new subgenus Lepidixodes for it. The palps are club-shaped and primitive; the hypostome is devoid of teeth ; the spiracles are laterally placed and near the middle of the body length; body, capitulum and legs are covered with scale-like structures; the claws are very thick; and the body is very hairy. All these characters are primitive and unlike those of the ticks; this form probably represents a special type of mite, and I see no reason for regarding it as a tick, let alone an Ixodes. Key to Species of Ixodes 3 Males Coxa I with two short spurs, internal slightly longer than external. Punctations medium and uniformly distributed. Cornua short and blunt I. granulatus Coxa I with two unequal spurs, internal very long, several times longer than external. Punctations large and restricted to lateral borders, region behind pseudoscutum and posterior border. Cornua absent I. spinicoxalis Females Hypostome with 3 : 3 dentition. Cornua absent. Auriculae small. Coxa I with two short spurs, internal twice as long as ex- ternal , I. granulatus H}^postome with 4 : 4 dentition at tip, then 3 : 3. Cornua short and blunt. Large auriculae as flat retrograde prominences. Coxa I with two unequal spurs, internal very long, several times as long as external I. spinicoxalis Ixodes praematurus Schulze. Ixodes praematurus Schulze, 1935, Zeitschr. Morph. Oekol. Tiere, 30, pt. 1, pp. 6, 8, 34, fig. 9 (n, Timor, off Zosterops 3 1. praematurus is known only from the nymphal stage. 16 ENTOMOLOGICA AMERICANA sp., a Timaliide, Geocirla [sic] = Geokichla ; holotype Z. M. Berl.) ; 1941, Op. cit., 37, pt. 3, pp. 497, 499, fig. 4. Schulze described this species from several nymphs taken bn Moetis Mountains in Timor by G. Stein in 1932. He states that he saw a tiny plate at the hypothetical point of union of the genital grooves, and since this structure lies in the position of the future genital orifice, he believes that it might be the primitive genital slit which later gives rise to the typical form found in the sexually ma- ture adult. I. praematurus differs from 7. ricinus in possessing yel- lowish-brown legs, an acuminate tarsus I and a more strongly nar- rowed scutum. I did not see this species, and I do not know of any other species with which this nymph can be correlated. Further specimens may afford a better understanding of its affinities. Ixodes granulatus Supino. Ixodes granulatus Supino, 1897, Atti Soc. Veneto-Trent. Sci. Nat., (2) 3, pt. 1, p. 232 2, Bhamo, Mooleyis, Tarra- waddy [Lower Burma], Terinzo [Upper Burma], off “ Sci- urus gordoni” = Callosciurus erythraeus gordoni, Felis tigris, “ Sciurus rufigenis” = Dr emomys rufigenis, “Sci- urus striatus” = Callosciurus quinquestriatus ; cotypes Ge- noa M.) ; 1897, Op. cit., (2) 3, pt. 2, pp. 248-249, PI. 12, figs. 5-6. Neumann, 1899, Mem. Soc. Zool. France, 12, pp. 164-165 ; 1902, Arch, de Parasit., 6, pt. 1, p. 125 (supple- mentary description based on types; 4 2, male of Supino is unengorged female). Nuttall and Warburton, 1911, Ticks, 1, pt. 2, Ixodes, pp. 285, 291-292 (status doubtful). Nuttall, 1916, Parasitology, 8, pt. 3, pp. 315-316, fig. 17 (examined types; 3 2> Pashok, Darjeeling District, India, off <(Epimys rufescens” = Rattus rufescens) . Warburton, 1926, Treubia, 8, pts. 3M:, p. 280 (2, Muaro Sako, Su- matra’s west coast). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 145, map 5; 1932, Vee- artsenijk. Meded., No. 79, pp. 30-31, 36, figs. 47-48, map 5 (2 .<$, Java, off Rattus concolor ephippium, Rattus rattus diardi; 2, Java and Bawean Island, off “ Mus rattus ” = Rattus rattus, Rattus concolor ephippium ) ; 1933, Zeitschr. f. Parasitenk., 5, pt. 2, pp. 410-411, fig. 3 (2 J1, 2, Moun- tain near Tjibodas, Java, off Rattus concolor ephippium and Rattus rattus diardi). Schulze, 1934, Op. cit., 7, pt. 2, pp. 169-170 ($, 2, n> Java, off ((Rattus rattus ro- ll ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 quei ” = Rattus rattus jalorensis, Rattus rattus brevicauda- tus, “Rattus bukit treubi” = Rattus fulvescens treubi, “Rattus bukit temmincki” = Rattus fulvescens temmincki, Rattus bartelsi, Rattus maxi, Rattus concolor ephippium, Rattus lepturus, “ Sciurus notatus notatus” = Callosciurus notatus notatus, “ Sciurus nigrivittatus nigrivittatus” = Callosciurus nigrovittatus nigrovittatus, “Sciurus notatus madurae” = Callosciurus notatus madurae, Lariscus insig- nis javanicus [sic], Tupaia javanica occidentalis, Croci- dura orientalis, Hylomys suillus suillus) ; 1939, Op. cit., 10, pt. 6, p. 724 (n, Mt. Victoria, Pakokku-Chin Hills, off Crocidura sp.). Toumanoff, 1944, Les Tiques (Ixodoidea) de Plndochine, p. 22, Pl. 7 (in part: only the male cited from Krijgsman and Ponto, 1932 ; female = Ixodes kempi Nuttall) . Ixodes ( Ixodes ) granulatus Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 11, 20-21 (Java, off “Tu- paia ferruginea” = Tupaia glis hypochrysa) . [non Ixodes granulatus Sharif, 1928, Records Indian Mus., 30, pt. 3, pp. 236-237 (J, Pashok in Darjeeling District, off “Epimys rufescens” = Rattus rufescens ; J, Kohima in Naga Hills District, Assam, off squirrel) = Ixodes kempi Nuttall] . In 1928 Sharif briefly described several female specimens from India as I. granulatus, without figuring them. He also examined the type material of Ixodes kempi Nuttall, 1913, and regarded it as synonymous with Ixodes granulatus. It is likely that Sharif only had specimens of kempi and incorrectly determined them as granu- latus, for his description fits kempi more closely than it does granu- latus. I saw specimens of granulatus from Burma which agree with the descriptions and figures given by Neumann in 1902 and Nuttall in 1916, both of whom saw the type specimens. I also saw six fe- males and one male of 7. kempi, in the Museum of Comparative Zo- ology, from Phong Saly and Muong Yo, Indochina, taken off Cal- losciurus imitator and Ratufa gigantea by R. Wheeler in May, 1929 ; the male was copulating with a female at the time it was collected and is preserved in this condition. After comparing these two spe- cies, I am convinced that kempi is a valid species and that it is dis- tinct from granulatus. Toumanoff recorded the female of granula- tus from Indochina, but his description and figures also resemble kempi more than they do granidatus. 18 ENTOMOLOGICA AMERICANA Fig. 2. Ixodes granulatus Supino, female : A, body in dorsal view; B, coxae I-IY ; C, tarsus I; D, spiracle; E. capitulum in dor- sal view ; F, same in ventral view ; G, tarsus IV. 19 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Specimens not being available from the East Indies, the follow- ing description is based upon females from Burma. Female (Fig. 2A-G). Two specimens respectively 1.8 and 1.7 mm. long and 0.96 and 1.05 mm. wide, exclusive of capitulum. Body elongate oval, much longer than wide. Scutum glossy brown, finely granulated, oval, covering about two-thirds of entire body, widest at mid-length, 1.19 mm. long and 0.77 mm. wide, with very few, scattered hairs ; scapulae very pointed ; punctations large, uni- formly distributed on posterior half ; cervical grooves superficial, scarcely visible, beginning some distance behind border of emargi- nation and diverging posteriorly, not reaching postero-lateral bor- ders of scutum; a lateral ridge present on each side, beginning at inner margin of scapula, but not reaching postero-lateral border; a depressed region between lateral ridge and cervical groove on each side, wide anteriorly and narrow posteriorly. Abdomen yellow, with numerous, long, white hairs; marginal groove complete, deep and conspicuously covered with long, white hairs; genital opening opposite coxa IV ; anal opening opposite posterior border of spi- racles; spiracles nearly circular, with large goblets; genital groove diverging widely, not reaching posterior margin of body; anal groove an inverted U. Legs long, fairly slender, brown, finely granulated, with numerous, long, white hairs; tarsi long, slender, narrowed abruptly before end, then gradually tapering to tip ; pul- villus nearly as long as claws ; coxa I with two short spurs, internal long and sharp, external short and blunt ; coxae II-III with a short, external spur ; coxa IV with an extremely short, inconspicuous spur, scarcely visible. Capitulum 0.728 mm. long; basis nearly triangu- lar, without cornua, posterior margin nearly straight ; porose areas large, oval, shallow, their distance apart equal to about one-half their diameter; ventrally basis broad, rounded posteriorly, with small auriculae ; palps long, slender, widest near mid-length ; slightly flattened on inner lateral border; article one apparent as a small protrusion ; article two slightly longer than three ; hypostome long, slender, pointed ; dentition 3 : 3, with ten to eleven teeth per file. Male. Not seen, but described by Krijgsman and Ponto (1933). Nymph and Larva. Described by Schulze (1934). Distribution and Hosts. Known from Burma, India and East Indies. No specimens were seen from Indonesia, but there are rec- ords from Java (Neumann, 1911 ; Krijgsman and Ponto, 1932 ; Schulze, 1934), Sumatra (Warburton, 1926) and Bawean Island (Krijgsman and Ponto, 1932). Specimens were seen at the Rocky 20 ENTOMOLOGICA AMERICANA Mountain Laboratory from the Philippine Islands (Mt. Data, Moun- tain Province, Luzon, off Rattus sp.), Assam (Ledo), Manipur (Im- phal), and Burma (Myitkyina), off Callosciurus erythraeus nagarum, Tupaia belangeri versurae, Callosciurus quinquestriatus quinquestriatus, Rattus rattus sladeni, Urocissa erythrorhyncha magnirostris, Rattus manipulus, mongoose and Rattus sp. Ixodes spinicoxalis Neumann. Ixodes spinicoxalis Neumann, 1899, Mem. Soc. Zool. France, 12, pp. Ill, 123-124, fig. 6 (4 J, Sumatra, off u Martes flavi- gula” = Martes flavigula henrici; holotype in Brit. M.). Salmon and Stiles, 1902, 17th Ann. Rept. Bur. Anim. In- dustry, (1901), p. 402. Nuttall and Warburton, 1911, Ticks, 1, pt. 2, Ixodes , pp. 139, 171-172, figs. 165-166 (based on types). Warburton, 1926, Treubia, 8, pts. 3-4, p. 280 (5, Andalos (Tandjung) Sumatra’s west coast, off Martes flavigula he'nrici). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 145, map 5 ; 1932, Veeartsenijk. Meded., No. 79, pp. 29-30, 36, fig. 46, map 5 ($, Java, off Helictis orientalis) . Schulze, 1934, Zeitschr. f. Parasitenk., 7, pt. 2, pp. 170-171, fig. 2 (rf, 5, n, 1, Tji- boeni, Mt. Lawoe, Mt. Tjareme, Pangrango, Java, off Para- doxurus hermaphroditus javanicus, Crocidura orientalis subsp., “ Sciur us nigrivittatus nigrivittatusf> = Callosciu- rus nigrovittatus nigrovittatus, Tupaia javanica occiden- talis, Rattus lepturus, ( ( Pithecheirus melanurus” = Pithe- cheir melanurus melanurus and (( Rattus bukit subsp.” = Rattus fulvescens bukit) ; 1939, Op cit., 10, pt. 6, p. 723. f Ixodes spinicoxalis Yakimov and Kohl-Yakimov, 1910, Arch, de Parasit., 14, p. 417 (Caucasia). Ixodes (Ixodes) spinicoxalis Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 10, 13. Female (Fig. 3A-G). Single, unengorged specimen 2.85 mm. long and 1.9 mm. wide, exclusive of capitulum. Body elongate oval, widest at level of spiracles. Scutum oval, with rounded borders, glossy, brown-maroon, with sides slightly darker than central por- tion; three specimens respectively 1.6, 1.5, and 1.6 mm. long and 1.4, 1.6 and 1.5 mm. wide ; large to medium, deep, closely set punctations restricted to outer portions of scutum; central portion depressed, slightly rugose, with punctations very small or absent; few, thin, white hairs; cervical grooves weak, superficial, scarcely visible, be- 21 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Pig. 3. Ixodes spinicoxalis Neumann, female : A, body in dor- sal view; B, coxae I-IV ; C, tarsus I ; D, tarsus IV ; E, capitulum in dorsal view ; P, same in ventral view ; G, spiracle. 22 ENTOMOLOGICA AMERICANA ginning some distance behind emargination and extending to region of large, posterior pnnctations ; no lateral ridges. Abdomen yellow, with marginal groove deep, conspicuous and complete ; ventral sur- face yellow, with long and uniformly distributed hairs; genital opening opposite coxa IV ; anal opening located very far posteri- orly; spiracles large, white, nearly circular, with numerous, small goblets ; genital grooves deep, divergent, extending to posterior mar- gin of body ; anal groove deep, forming an inverted U. Legs long, slender, reddish-brown, with distal portion of each joint, except last, yellowish-white, giving leg appearance of being banded ; tarsus I very long, slender, tupering abruptly near tip ; pulvillus nearly as long as claws ; coxae long, nearly touching in unengorged specimen ; coxa I with very long, slender, pointed, internal spur, nearly reach- ing posterior border of coxa II, and very short, external spur ; coxae II— III with short, external spur and faint suggestion of an internal spur ; coxa IV with single, short, external spur. Capitulum 0.94 to 1.05 mm. long in three specimens; basis darker than palps, broader than long, sub-triangular, with short, blunt cornua; porose areas large, oval to rounded; palps long, flat, widest at mid-length; ar- ticle one visible as slight protrusion ; article two slightly longer than article three ; ventral auriculae large, in form of flat, retrograde prominence ; hypostome long, narrow, spear-shaped, with 4 : 4 den- tition at anterior tip and 3 : 3 in posterior part. Male , Nymph and Larva. Not seen, but described by Schulze (1934). Distribution and Hosts. Known from Sumatra (Neumann, 1899) and Java (Krijgsman and Ponto, 1932; Schulze, 1934). The record from Caucasia (Yakimov and Kohl-Yakimov, 1910) is prob- ably an error in determination. Three females were seen at the Museum of Comparative Zoology from Tjibodas, Mt. Gede, Java, collected September 12, 1909, by Palmer and Bryant off Mydaus javanensis javanensis. Genus HaemapJtysalis C. L. Koch, 1844 Key to Species of Haemaphysalis Males 1. Coxa IV with long, slender, internal spur and shorter-, external spur. Coxa I with long spur. Scutum fusiform. Puncta- tions numerous, unequal and uniformly distributed. Ar- ticle three with sharp, lateral spur. Article two very salient 23 t ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 laterally H. cornigera Coxa IV with only one spur 2 2. No lateral grooves present 3 Short or long lateral grooves present 4 3. Article two with lobe-like projection on antero-internal border. Article three with distinct, dorsal, retrograde spur. Hy- postome with 4 : 4 dentition. Scutum elongate-oval. Pune- tations medium and irregularly distributed II. hystricis Article two without lobe-like projection. Article three with only slight indication of dorsal spur. Hypostome with 5 : 5 den- tition. Scutum broad-oval. Punctations extremely fine and uniformly distributed II. toxopei 4. Lateral groove very short. Cornua strongly developed and ex- tending slightly beyond anterior border of scutum. Scutum elongate-oval. Punctations numerous, superficial and regu- larly distributed. Coxae I-IV with single, lobe-like spur. Article two strongly salient laterally H. traguli Lateral groove long 5 5. Palpal article three with distinct, dorsal, retrograde spur 6 Palpal article three without such spur, but posterior border may protrude slightly 8 6. Article three with distinct, dorsal, retrograde spur at inner angle. Article two very salient laterally. Scutum broadest posteriorly. Punctations few, unequal and sparsely dis- tributed. Coxae I-IV with single, short spur. H. wellingtoni Article three with dorsal, retrograde spur at middle of posterior border T 7 7. Coxa I with short, blunt spur. Palpal article two strongly sali- ent laterally. Scutum elongate-oval. Punctations coarse, uniform and closely set H. hylobatis Coxa I with long, pointed spur. Palpal article two only slightly salient laterally. Scutum broad-oval. Punctations medium and uniformly distributed H. bispinosa 8. Scutum very elongate. Punctations medium, numerous and uni- formly distributed. Article two very strongly salient later- ally, with strong, lobe-like projection on antero-internal border. Coxa I with short, blunt spur H. honing sb erg eri Scutum elongate-oval. Punctations medium, less numerous and irregularly distributed. Several raised tracts present mak- ing surface very irregular. Palpal article two only slightly 24 ENTOMOLOGICA AMERICANA salient laterally, with weak lobe-like projection on antero- internal border. Coxa I with short, sharp spur. H. papuana Females 1. Cervical grooves well-marked, sinuons and reaching posterior margin of scutum. Scutum circular. Punctations numer- ous and shallow. Cornua slight II. toxopei Cervical grooves not reaching posterior margin of scutum 2 2. Palpal article three with distinct, dorsal, retrograde spur, more or less erect 3 Palpal article three without such spur, but posterior border may protrude slightly 6 3. Dorsal spur at inner angle of posterior border. Palpal article two very salient laterally. Basis about three times as broad as long. Scutum slightly longer than broad ... H. wellingtoni Dorsal spur at middle of posterior border 4 4. Coxa I with short, blunt spur. Palpal article two strongly sali- ent laterally, with distinct lobe-like projection at antero- internal border. Basis about three times as broad as long. Cornua weak. Scutum nearly circular. Punctations coarse and uniformly distributed H. hylobatis Coxa I with medium, pointed spur 5 5. Cervical grooves broad, shallow and nearly parallel. Spiracles circular, with distinct process. Punctations unequal and ir- regularly distributed H. hystricis Cervical grooves well-marked, deep and converging, then shal- low and diverging. Spiracles oval, without process. Punc- tations equal and more or less uniformly distributed. H. bispinosa 6. Coxae II-III with broad, ridge-like spurs. Scutum slightly broader than long. Punctations equal and uniformly dis- tributed. Basis about three times as broad as long. Cornua large, broad and blunt. Articles two and three salient later- ally H. traguli Coxae II-III with short spurs, about as wide as long 7 7. Scutum much longer than wide. Punctations equal and uni- formly distributed. Basis about three times as broad as long. Palpal article two very salient laterally, with lobe- like projection on antero-internal border. H. honing sbergeri 25 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Scutum slightly wider than long 8 8. Palpal article two very salient laterally, posterior margin wavy and strong, lobe-like projection on antero-internal border. Basis about three times as broad as long. Coxa I with long, pointed spur H. cornigera Palpal article two only slightly salient laterally, posterior mar- gin nearly straight, weak lobe-like projection on antero-in- ternal border. Basis about three times as broad as long. Coxa I with short, pointed spur H. papuana Haemaphysalis bispinosa Neumann. Haemaphysalis bispinosa Neumann, 1897, Mem. Soc. Zool. France, 10, pp. 341-342, 358, figs. 7-8 (1 §, Ramnad, India; holotype described from E. Simon Coll., probably now at Paris M.). Salmon and Stiles, 1902, 17th Ann. Rept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), pp. 458, 459. Warburton, 1907, Journ. Econ. Biol., 2, pt. 3, p. 94; 1907, Imp. Dept. Agric., India, Bull. No. 6, pp. 1, 7, 8, 9, 11, 12, fig. 9 (Bombay and Poona, off horse and cattle) ; 1908, Proc. Cambridge Philos. Soc., 14, pt. 5, pp. 517-518, figs. 9-10 (J', 2, India and Ceylon). Blanchard, 1909, L’ln- secte et 1 ’Infection, Fasc. 1, Acariens, p. 148. Warburton, 1910, Parasitology, 3, pt. 4, p. 396 (India, off dog, tiger and Macaca) . Patton and Cragg, 1913, Textbook Medical Ent., p. 648, PI. 73, fig. 11, PI. 81, figs. 5-6. Nuttall and War- burton, 1915, Ticks, 1, pt. 3, Haemaphysalis , pp. 354, 360, 361, 426-433, figs. 358-362 (J*, J, n, 1, extensive distribution and host records). Nieschulz, 1924, Nederl. Tijdschr. v. Diergeneesk., 51, p. 195 (Kota-Radja in Atjeh, Sumatra, off cattle). Myers, 1924, New Zealand Dept. Agric., Bull. No. 116, pp. 1-105, figs. 1-17 (life history, biology, distribu- tion, hosts and control). Fielding, 1926, Comm. Austral. Service Publ. (Trop. Div.) No. 9, pp. 60-62, fig. 23. Ogura and Takada, 1927, Journ. College Agric., Hokkaido Imp. Univ., 18, pt. 4, pp. 204—205, PI. 14, figs. 1-6 ( Japan, off horse, cattle, and dog; location of types not given). Warburton, 1908, Proc. Cambridge Philos. Soc. 14, pt. 5, p. 519. Blanchard, 1909, LTnsecte et l’ln- fection, Fasc. 1, Acariens, p. 154. Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 106, 107, 109 (lCf, 5, Japan, Ceylon, off Equus caballus, Bos taurus and Canis familiar is) . ? Haemaphysalis neumanni Galli-Valerio, 1909, Centralbl. f. Bakteriol. u. Parasit., 1 Abt., Orig., 51, pt. 5. p. 539 (Cey- lon, off Cams aureus). Yakimov and Kohl-Yakimov, 1910, Arch, de Parasit., 14, p. 418 (6 $, China, Primorsk Govern- ment, E. Siberia, off cattle). Haemaphysalis birmaniae Neumann, 1911, [in part, nec Su- pino, 1897] Das Tierreich, Lief. 26, Acarina, Ixodidae, p. 109. ? Haemaphysalis bispinosa Olenev, 1931, Zeitschr. f. Parasi- tenk., 4, pt. 1, p. 133 (Vladivostok). Pomerantzev, 1946, Inst. Zool. Acad. Sci. URSS, Tabl. Analyt. Faune URSS, 26, pp. 13, 14 (Russia). Haemaphysalis renschi Schulze, 1933, Arch. f. Hydrobiol., Suppl. Bd. 12, pp. 490, 492 (Flores, in key only) ; 1936, Zeitschr. f. Parasitenk., 8, pt. 5, pp. 52L-526, fig. 6 (2 J1, Rana Mese, Flores; holotype Z. M. Berlin). 27 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Numerous males of bispinosa from Roeteng, Flores Island, in the Kraneveld collection agree in all respects with the illustrations and description of renschi given by Schulze for two males from the same locality. These are definitely the same species, and I regard H. renschi Schulze as a synonym of H. bispinosa Neumann. Male (Fig. 4A-F). Twenty-five specimens range from 2.1 to I. 68 mm. long by 1.4 to 1.26 mm. wide, exclusive of capitulum; av- erage 1.88 mm. long by 1.37 mm. wide. Body long, oval and widest at mid-length, yellow to brown depending on degree of hardening ; some with brown, hardened markings in intervals between festoons ; punctations medium, very numerous and uniformly distributed; cervical grooves short and inconspicuous ; lateral grooves beginning near mid-length and passing posteriorly to region of first festoon; festoons distinct and longer than wide ; body ventrally yellow to brown, with many, minute punctations and with very small, white hairs ; genital opening opposite coxa II ; anal opening opposite pos- terior portion of spiracle ; spiracle oval and slightly narrowed an- teriorly, with numerous, very small goblets. Legs long and thick, with many, long, white hairs ; coxa I with single, long, pointed spur ; coxae II-IV with a very short spur ; tarsi short, tapering gradually ; pulvillus almost as long as claws. Capitulum 0.42 mm. long, longer than wide; basis rectangular, slightly wider than long, with large, distinct cornua; palps sfnall and slightly salient; articles two and three subequal ; sharp, retrograde, dorsal spur and long, sharp, ven- tral spur on middle of posterior border of article three ; hypostome very short ; dentition 4 : 4, with approximately seven to eight teeth per file. Female (Fig. 4G-L). Twenty-five specimens range from 2.25 to 1.68 mm. long by 1.5 to 1.26 mm. wide, exclusive of capitulum; average 2.18 mm. long by 1.48 mm. wide ; fully engorged specimen 7.8 mm. long by 6.0 mm. wide. Scutum 0.90 mm. long by 0.96 min. wide, yellow and nearly circular ; cervical grooves anteriorly fairly deep and slightly convergent, posteriorly shallow and divergent nearly reaching postero-lateral border; punctations medium, uni- formly distributed and very numerous; long marginal groove be- ginning behind scutum and including first festoon; festoons about as wide as long. Abdomen very finely punctate ventrally, with short, white hairs ; genital opening opposite anterior border of coxa III; anal opening opposite posterior border of spiracles; spiracles similar to male ; genital grooves diverging gradually ; anal grooves fusing anteriorly with posterior portion of genital grooves. Coxae 28 ENTOMOLOGICA AMERICANA Fig. 4. Ha&maphysalis bispinosa Neumann. A-F, male : A, scutum; B, coxae I-IV ; C, tarsus IV; D, spiracle; E, capitulum in dorsal view ; F, same in ventral view. G-L, female : G, scutum ; H, coxae I-IV ; I, tarsus IV ; J, spiracle ; K, capitulum in dorsal view; L, same in ventral view. 29 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 similar to male; tarsi long and tapering gradually; pulvillus long and narrow, but slightly shorter than claws. Capitulum 0.54 mm. long; basis much wider than in male; porose areas large, rounded and widely separated ; hypostome short ; dentition 4 : 4, with ap- proximately eight to nine teeth per file. Distribution and Hosts. Known from India, Burma, Andaman Islands, Federated Malay States, Ceylon, Japan, Borneo, Australia, New Zealand, East Indies, Formosa, Indochina, East Africa (in- troduced from India), Siberia (?) and China. In the East Indies it has been recorded from Sumatra (Nieschulz, 1924 ; Krijgsman and Ponto, 1931, 1932), Java, Soembawa, Karimoendjowo Island, Cele- bes (Krijgsman and Ponto, 1931, 1932) and Flores (Schulze, 1936). Numerous specimens were seen in the Kraneveld collection from Sumatra (Lhokseumawe, off cow and goat; Padangsidimpoean, off cow; Koetaradja, off cow), Java (Madioen, off dog), Bali (Singa- radja, off cow, horse and sheep), Flores (Roeteng, off buffalo, cow and horse) and Soela Island (off domestic pig). Specimens were also seen in the Museum of Comparative Zoology from Australia and China and in the Rocky Mountain Laboratory from China, Ja- pan, New Caledonia, Australia, Burma, Assam and India off a great variety of hosts. Haemaphysalis cornigera Neumann. Haemaphysalis cornigera Neumann, 1897, Mem. Soc. Zool. France, 10, pp. 350-352, 359, figs. 16-17 (2 1 §, Singa- pore ; 2 <$, Borneo, off deer ; 1 J1, Sumatra, off Bos bubalis [nec from Judea] ; cotypes described from E. Simon Coll., now probably at Paris M. ; R. Blanchard Coll. ; and Oudemans Coll., now at Leiden M.) ; 1910, Ann. Sci. Nat. Zool., 9, pp. 174^175. Salmon and Stiles, 1902, 17th Ann. Rept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), p. 458. Blanchard, 1909, LTnsecte et l’lnfection, Fasc. 1, Acariens, p. 150, fig. 185. Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 106, 107, 112-113, figs. 57-58. Nuttall and Warburton, 1915, Ticks, 1, pt. 3, Haemaphy- salis, pp. 354, 358, 500-504, figs. 441-445 (J', 5> Federated Malay States, British North Borneo, India, off wild pig, tiger, Sambar deer, water buffalo, buffalo). Larrousse, 1925, Ann. Parasit. Hum. et Comp., 3, pt. 3, pp. 301-302, fig. 1 (4 Yado-Carin Ascinii- Cheba, Carin-Cheba, Burma, off <( Hystrix bengalensis = Hystrix hodgsoni, <(TJrsus torquatus” -Enarctos thibe- tanus thibetanus ; cotypes Genoa M.) ; 1897, Op. cit., (2) 3, pt. 2, p. 252, PI. 13, figs. 19-20. Neumann, 1897, Mem. Soc. Zool. France, 10, pp. 342-343. Warburton, 1908, Proc. Cambridge Philos. Soc., 14, pt. 5, pp. 518-519, fig. 11. Blanchard, 1909, L’Insecte et 1 ’Infection, Fasc. 1, Acariens, p. 152. Warburton, 1910, Parasitology, 3, pt. 4, p. 396 (locality uncertain, possibly Burma, off Geoemyda spinosa). Nuttall and Warburton, 1915, Ticks, 1, pt. 3, Haemaphy salis, pp. 353, 357, 422-426, figs. 354-357 (ex- tensive distribution and host list). Kishida, 1922, Zool. Mag., Tokyo, 34, 408, pp. 854, 855. Warburton, 1926, Treubia, 8, pts. 3-4, pp. 279, 280 (§, Bukit Kaba, Ben- koelen, Sumatra, off Arctonyx collaris hoevenii; Babi Island, off Nias pig). Sharif, 1928, Records Indian Mus., 30, pt. 3, pp. 253-254, fig. 11 (J\ J, Assam, off Felis tigris ; Assam ; J, ? Burma, off Geoemyda spinosa ; 2, Ben- gal). Schulze, 1931, Sitz.-Ber. Abhand. Naturf. Gesellsch. Rostock, (3) 3, p. 54, fig. 5 (n, Tsingtau, China, off hedge- 35 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 hogs). Krijgsman and Ponto, 1931, Zeitschr. f. Para- sitenk., 4, pt. 1, pp. 141, 142, map 3; 1932, Veeartsenijk. Meded., No. 79, pp. 1F-15, 33, 34, figs. 14-15, map 3 ( J1, 5, Sumatra, Bali, Celebes, Karimoendjowo Island, Riouw Island, off cow and buffalo). Kishida, 1935, Lansania, 7, 69, pp. 138-139. Sugimoto, 1935, Taiwan-No-Chikusan, 3, pt. 9, pp. 26-27. Schulze, 1935, Zool. Anz., 112, pts. 9-10, p. 234 (Formosa). Sugimoto, 1936, Journ. Soc. Trop. Agric., Formosa, 8, pt. 4, p. 336 (Formosa) ; 1936, Journ. Cent. Soc. Vet. Med., Tokyo, 49, pt. 7, p. 581. Ogura, 1936, Mem. Fac. Sci. Agric. Taihoku Imp. Univ. Formosa, 19, pt. 2, pp. 81-82, figs. 19-23 (.J1, J, Formosa, off dog and wild boar). Sugimoto, 1937, Journ. Cent. Soc. Vet. Med., Tokyo, 50, pt. 5, pp. 311-313; 1937, Journ. Soc. Trop. Agric., Formosa, 9, pt. 3, p. 287. Schulze, 1939, Zeitschr. f. Parasitenk., 10, pt. 6, p. 728 (Burma). Tou- manoff, 1944, Les Tiques (Ixodoidea) de l’lndochine, pp. 52-55, 85, 90, Pis, 33, 34, 46 §, Haut Chlong, off Sambar deer). Haemaphysalis bispinosa Neumann, 1901, [nec Neumann, 1897], Mem. Soc. Zool. France, 14, pp. 261-262 (1 J1, 4 Macao, China, off “ Paradoxurus larvatus” = Paguma lar- vata ; 2$, Japan). Haemaphysalis semermis Neumann, 1901, Op cit., p. 263 (1 J', Bengkalis Island; type in Paris M.). Haemaphysalis histricis [sm] Neumann, 1902, Arch, de Parasit., 6, pt. 1, p. 128. Haemaphysalis birmaniae Neumann, 1911, [in part, nee Supino, 1897], Das Tierreich, Lief. 26, Acarina, Ixodidae, p. 109. Male (Fig. 6A-E). Three Sumatra specimens respectively 3.0, 2.7, and 2.5 mm. long by 2.1, 2.0, and 2.0 mm. wide, exclusive of capitulum. Body elongate-oval, narrowed slightly anteriorly, wid- est at mid-length. Scutum yellow to light brown; punctations small, shallow, irregularly distributed ; cervical grooves small, form- ing deep pits anteriorly and shallow, diverging grooves posteriorly; dorsal surface irregularly and weakly sculptured with fairly shal- low depressions ; pseudoscutum set off by slight elevation ; lat- eral grooves lacking, or at most present as a very short groove in front of first festoon on each side ; festoons about twice as long as broad. Ventral body surface light yellow, finely punctate, with numerous short, white, inconspicuous hairs ; genital opening oppo- 36 ENTOMOLOGICA AMERICANA Fig. 6. Haemaphysalis hy sir ids Supino. A-E, male: A, scu- tum; B, coxae I-IV ; C, spiracle; D, capitulum in dorsal view; E, same in ventral view. F-K, female : F, scutum ; G, coxae I-IV ; H, tarsus IV ; I, spiracle; J. capitulum in dorsalview; K, same in ventral view. 37 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 site coxa II ; anal opening opposite spiracle ; spiracle comma- shaped with distinct, dorsal process ; genital grooves diverging widely around anus; arms of anal groove almost reaching genital grooves. Legs long, stout and very hairy; coxa I with relatively long, external spur ; coxae II-IY with a shorter spur ; tf bchanter I with a distinct dorsal and ventral retrograde spur ; tarsi sloping fairly rapidly ; pulvillus about two-thirds length of claws. Capitu- lum 0.56 mm. long; basis rectangular, with sides nearly straight; cornua large, about as wide as long ; palps only slightly salient laterally; palpal articles two and three subequal; article two with a lobe-like projection on antero-internal margin ; article three with a distinct dorsal spur on middle of posterior border and a strong, ventral spur ; hypostome short ; 4:4 dentition, with ten to twelve teeth per file. Female (Fig. 6F-K}. Unengorged Sumatra specimen 3.3 mm. long by 2.3 mm. wide, exclusive of capitulum. Body elongate-oval, widest at mid-length. Scutum of five specimens from 1.2 to 0.075 mm. long to 1.5 to 1.0 mm. wide, yellow, with scapulae darker; punctations medium, numerous, regularly distributed ; cervical grooves as deep pits somewhat removed from anterior margin, con- tinuing posteriorly as slightly diverging, shallow grooves not reach- ing margin of scutum. Abdomen with medium and very small punctations; long, narrow, marginal grooves including first two festoons on each side ; festoons slightly broader than long ; abdo- men ventrally lighter in color, with numerous, fine, inconspicuous punctations; spiracles nearly circular, narrowed slightly dorsally; genital opening opposite coxa II ; anal opening opposite spiracles ; genital grooves diverging gradually, but more widely in region of anal opening ; arms of anal groove reaching genital grooves. Legs and coxae as in male. Capitulum 0.64 mm. long ; basis rectangular, about three times as wide as long; porose areas large, oval, deep, far apart, with a small pit between them; cornua strong, large, blunt ; palps about twice as long as wide, slightly salient laterally ; article two longer, with lobe-like protuberance on antero-internal margin ; article three with a distinct dorsal spur, and a longer ven- tral spur ; hypostome short ; 4:4 dentition, with ten to twelve teeth per file. Distribution and Hosts. Known from Burma, India, Assam, China, Federated Malay States, Borneo, East Indies, Ceylon, For- mosa and Indochina. In the East Indies it has been recorded from Bengkalis (Neumann, 1901), Sumatra (Nuttall and Warburton, 38 ENTOMOLOGICA AMERICANA 1915; Warburton, 1926; Krijgsman and Ponto, 1931, 1932), Babi Island (Warburton, 1926), Celebes (Nnttall and Warburton, 1915; Krijgsman and Ponto, 1931, 1932), Bali, Karimoendjowo Island, Siberoet, and Riouw Island (Krijgsman and Ponto, 1931, 1932). Specimens were seen from Sumatra (Lhokseumawe, one female, off buffalo ; Medan, one male, off domestic pig ; Padangsidimpoean, one male, off horse; Benkoelen, one female, off dog; Tandjoengkarang, one female, off dog; all in Kranevehl collection; between Takin- genn and Bireuen, one female, in Museum of Comparative Zoology) and Celebes (Makale, one female, off buffalo, in Kraneveld collec- tion). One female taken off Geoemyda spinosa in India and three specimens collected by H. S. Puller in 1944 in Burma (Shadnznp, male, off man; Tingkawk Sakan, female, off man; Shingbwiyang, male, off man) were seen in the Museum of Comparative Zoology. In addition specimens were seen in the Rocky Mountain Laboratory from Burma, Assam, India and Formosa taken off a wide variety of hosts. Biology . This species is primarily a parasite of mammals, but it has been recorded off the tortoise, Geoemyda spinosa (Nuttall and Warburton, 1915; Sharif, 1928). In the Museum of Compara- tive Zoology was seen one engorged female from India also taken off Geoemyda spinosa, and from these records it would appear that it is occasionally found on tortoises. i Haemaphy satis honing sbergeri Nuttall and Warburton. ? Haemaphy satis leachi var. australis Neumann, 1905, Arch, de Parasit., 9, pt. 2, p. 238 (2 J', Sumatra, off Felis tigris ; 7 J, New South Wales, off horse). Warburton, 1910, Parasitology, 3, pt. 4, p. 396 (off Felis tigris). Haemaphysalis honing sb erg eri Warburton and Nuttall, 1909, Parasitology, 2, pts. 1-2, pp. 65-66, figs. 11-12 (2 2 J, Java, off Felis pardus ; cotypes in Brit. Mus.). Schulze, 1934, Zeitschr. f. Parasitenk., 7, pt. 2, pp. 168-169 (larger dark form, 10 J', 3 2, Tjiboeni, Java, off Faradoxurus her- maphroditus javanicus ; 3 J, Pangrango, Java, off Felis bengalensis javanensis ; 2 J', 1 J, Pangrango, Java, off Helictis orient alis; 2 n, Tjiboeni, Java, off Ratufa bicolor bicolor; smaller light form, 4 J', 1 J, 7 n, Tjiboeni, Java, off Petaurista petaurista petaurista ; 1 J', 2 J, Mt. Karang, Java, off Petaurista petaurista petaurista ) ; 1938, Zeitschr. Morph. Oekol. Tiere, 34, pt. 1, p. 140, fig. 45 (morpholog- ical). 39 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Haemaphy satis leachi Nuttall and Warburton, 1915, Ticks, 1, 3, Haemaphy salis, pp. 355, 357, 460-476, figs. 409-410 (in part: 5, Sumatra, off tiger; Java, off Felis pardus; Upper Sarawak, Borneo, off dog; Pahang, Fed. Malay States, off tiger, Paradoxurus, Viverra and Felis chans ; Uln Gombak, Selangor, Fed. Malay States, off “TJrsus ma- layanus” = Helarctos malayanus ; Kampong Batu, Negri Sembilan, Fed. Malay States, off Felis pardus). War- burton, 1926, Treubia, 8, pts. 3-4, pp. 279, 280 $, Sinabang, Simeuloee, off Paradoxurus hermaphroditus ; Andalos (Tandjung), Sumatra’s west coast, off Maries fla- vigula henrici, <(Nycticebus buhu” = Nycticebus coucang). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 143, map 3 ; 1932, Veeartsenijk, Meded., No. 79, pp. 16-17, 33, 34, figs. 18-19, map 3 (lCJ, 5, n, Java, Sumatra, Borneo, Simeuloee, Bawean, off ((Mus rattus” = Battus rattus, Helictis orient alis and panther). Not of Audouin, 1827. Haemaphysalis bartelsi Schulze, 1938, Zeitschr. Morph. Oekol. Tiere, 34, pt. 1, p. 140, fig. 5 a (lighter, smaller form of H. honing sbergeri; type probably in P. Schulze Coll.). I have seen three males and one female from Tjibodas, Java, which I regard as honing sbergeri. These were compared with nu- merous specimens of leachii from Africa, and though there is a su- perficial resemblance, they differ enough to be regarded as a dis- tinct species and not as a variety of leachii. The main difference between true leachii and honing sbergeri, in both sexes, lies in the structure of the capitulum. In honing sbergeri the two palps are bell-shaped with the external lateral margin slightly concave ; ar- ticle two is longer, more erect and its posterior border does not terminate in a distinct dorsal spur; article three is as long as wide; and the basis is wider. In leachii the palps are compressed, with the external lateral margin straight or slightly convex ; article two is compressed, shorter, and has a distinct dorsal spur on the pos- terior border; article three is wider than long; and the basis is narrower. In 1934 Schulze stated that honing sbergeri appeared in two dif- ferent forms, a larger, darker one found on many hosts and a smaller, lighter one found only on Petaurista. In 1938 he decided that the smaller forms represented a new species which he called bartelsi. Size alone is not a good character for separating species, 40 ENTOMOLOGICA AMERICANA A-F, male : A, scutum ; B, coxae I-IV ; C, tarsus IV ; D, spiracle ; E, capitulum in dorsal view; F, same in ventral view. G-L, fe- male :-G, scutum; H, coxae I-IV; I, tarsus IV; J, spiracle; K, capitulum in dorsal view ; L, same in ventral view. 41 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 for the amount of nymphal feeding has a direct relation to adult size, and large and small forms are known to occur in many species. I regard bartelsi as a synonym of honing st> ergeri. Neumann in 1905 said that leachii was strictly an African spe- cies, and he placed specimens from Sumatra and Australia in a new variety, australis. It is probable that the material Neumann had from Sumatra is identical with honing si) ergeri, but there is no way at present of being certain. If found to be the same as honings- t) ergeri, australis will have priority as the earlier name. Male (Fig. 7A-F). Three Java specimens respectively 2.5, 2.4 and 2.4 mm. long by 1.26, 1.32, and 1.25 mm. wide, exclusive of capitulum. Body much longer than wide, widest at posterior end, yellow-brown; punctations medium, deep, partially confluent; cer- vical groove as a deep anterior pit continuing posteriorly as a very shallow groove ; lateral grooves shallow, very inconspicuous, includ- ing first festoon on each side and extending only a very short dis- tance anteriorly; festoons little over twice as long as wide, with grooves separating them distinct and deep. Abdomen ventrally lighter, finely striate; genital opening opposite coxa II, with a dis- tinct cover having a posterior fringe of many, short hairs; anal opening opposite spiracles; spiracles large, oval and with a blunt, dorsal continuation ; genital grooves diverging gradually till reach- ing region of anal opening and then more divergent ; arms of anal groove not reaching genital grooves. Legs normal ; tarsi relatively short, narrow abruptly at end and not ending in a spur; pulvillus nearly as long as claws ; coxa I with a short, blunt spur ; coxae II- IV with a blunt spur, slightly shorter than that of coxa I ; numer- ous, long, conspicuous hairs on coxal plates. Capitulum 0.49 mm. long by 0.728 mm. wide; basis wider than long, widest anteriorly; large punctations grouped together near center; cornua long and sharp ; palps slightly bell-shaped, with external lateral margin con- cave; article two longer than article three, very salient laterally; internal lateral margin of article two with a prominent lobe-like extension; article three triangular, slightly wider than long and with a long ventral spur ; hypostome short ; dentition, 4 : 4, with ap- proximately eight teeth per file. Female (Fig. 7G-L). Single, engorged specimen 6.9 mm. long by 4.5 mm. wide, exclusive of capitulum. Scutum elongate-oval, pointed posteriorly, 1.2 mm. long by 1.05 mm. wide, reddish-brown in anterior two-thirds and remainder yellow; punctations numer- ous, medium, deep, uniformly distributed ; cervical grooves deep 42 ENTOMOLOGICA AMERICANA anteriorly, converging toward center, becoming shallow posteriorly and then diverging again. Abdomen uniformly covered with me- dium punctations; festoons in specimen distorted and not clearly seen; ventral surface lighter, finely striate; genital opening oppo- site coxa II ; anal opening posterior to spiracle ; spiracle oval and pointed dorsally. Tarsi slightly longer and not tapering as sud- denly as those of male ; coxae as in male. Capitulum 0.64 mm. long by 0.91 mm. wide ; basis rectangular, widest anteriorly being about two and a half times as wide as long; porose areas deep and elon- gate-oval ; palps similar to male ; hypostome small ; dentition 4 : 4, with approximately eight teeth per file. Distribution and Hosts. So many of the records are uncertain that the complete distribution can not be given. It is definitely known in the East Indies from Java (Warburton and Nuttall, 1909, 1915; Schulze, 1934; Krijgsman and Ponto, 1931, 1932), Su- matra (Neumann, 1905 Nuttall and Warburton, 1915; Warbur- ton, 1926; Krijgsman and Ponto, 1931, 1932), Simeuloee (Warbur- ton, 1926; Krijgsman and Ponto, 1931, 1932) and Bawean Island (Krijgsman and Ponto, 1931, 1932). Three males and one female were seen in the Museum of Comparative Zoology from Tjibodas, Mt. Gede, Java, taken off My dans javanensis javanensis by Palmer and Bryant. It is probable that honing sbergeri is also found in Australia (Neumann, 1905). I doubt if it occurs as far west as Burma, for I examined many specimens from Myitkyina in the Rocky Moun- tain Laboratory and found them to be very close to typical leachii. Specimens from Calcutta, India, in the Museum of Comparative Zoology are also very close to typical leachii. Haemaphysalis papuana Thorell. Haemaphysalis papuana Thorell, 1882, Ann. Mus. Civ. Genova, 18, pp. 62-66, 69 [pp. 46-50, 53 in reprint] , PL 6, figs. 40-45 (1,J\ 1 J, Ramoi, New Guinea; type in Genoa M.). Neumann, 1897, Mem. Soc. Zool. France, 10, pp. 336-337, 358, 359. Salmon and Stiles, 1902, 17th Ann. Rept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), pp. 458, 459. Warburton, 1908, Proc. Cambridge Philos. Soc., 14, pt. 5, pp. 514-516, fig. 7 J, n, Sarawak, Borneo). Blanchard, 1909, LTnsecte et lTnfection, Fasc. 1, Acariens, p. 154. Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodi- dae, pp. 106, 107, 108. Nuttall and Warburton, 1915, Ticks, 1, pt. 3, Haemaphysalis, pp. 355, 359, 360, 361, 404- 43 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 407, figs. 338-341 (, 20 larvae, Berhala 47 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Island, off Tragulus kanchil; types now in Leiden ,M.). Schulze, 1934, Zeitschr. f. Parasitenk., 8, pt. 2, pp. 168, 169, fig. 1 (1 J, Wijnkoops Bay, Java, off “Tragulus kan- chil pelandoc” = Tragulus kanchil kanchil). Haemaphysalis monospinosa Krijgsman and Ponto, 1932, Vee- artsenijk. Meded. No. 79, pp. 18-19, 33, fig. 22, map 3 (J1, Berhala Island, off Tragulus kanchil ; type in Buitenzorg Zool. M.) ; 1933, Zeitschr. f. Parasitenk., 5, pt. 2, pp. 409- 410, fig. 2. Toumanoff, 1944, Les Tiqnes (Ixodoidea) de l’Indochine, pp. 80, 86, PI. 62. This species was not seen, and the description of the male is from Ondemans, 1928, and Krijgsman and Ponto, 1932, and that of the female is from Ondemans, 1928, and Schulze, 1934. Male. Scutum of Ondemans’ specimen 1.865 mm. long by 1.065 mm. wide and in Krijgsman ’s specimen 1.7 mm. long by 1.0 mm. wide ; cervical grooves very short, crescent-shaped ; marginal grooves very short, distinct; punctations superficial, numerous, regularly distributed. Coxa I much longer than broad, nearly diamond- shaped ; coxae II-IV with a short, broad, lobe-like spur ; trochantal spurs well developed. Basis capituli broader than long; cornua very strongly developed, extending slightly beyond anterior border of scutum; contours of palpal article two very salient laterally; article three with a very long, ventral, retrograde spur ; hypostome with 4 : 4 dentition. Female. Scutum slightly broader than long, in Oudemans’ two specimens 0.854 mm. long by 1.0 mm. wide and 0.808 mm. long by 0.881 mm. wide; punctations large, uniformly distributed. Coxa I with two broad spurs ; coxae II-IV with a broad, ridge-like spur. Basis capituli about three times as broad as long; porose areas small, oval, widely separated ; cornua large, broad and blunt; palpal articles two and three salient laterally ; hypostome with 4 : 4 dentition; palpal article three ventrally with a very long, retro- grade spur. Haemaphysalis wellingtoni Nuttall and Warburton. Haemaphysalis wellingtoni Nuttall and Warburton, 1908, Proc. Cambridge Philos. Soc., 14, pt. 4, pp. 397-398, figs. 9-11 (1 2 $, Sarawak, Borneo, off domestic fowl; cotypes in Brit. Mus.). Blanchard, 1909, LTnsecte et 1 ’Infection, Fasc. 1, Acariens, p. 157. Nuttall and Warburton, 1915, Ticks, 1, pt. 3, Haemaphysalis, pp. 356, 358, 359, 361, 479- 48 ENTOMOLOGICA AMERICANA A-F, male : A, scutum ; B, coxae I-IV ; C, tarsus IV ; D, spiracle ; ,E, capitulum in dorsal view; F, same in ventral view. G-L, fe- male : G, scutum ; H, coxae I-IV ; I, tarsus IV ; J, spiracle ; K, ca- pitulum in dorsal view ; L, same in ventral view. 49 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 482, figs. 416-420 ( J', J n, 1; Borneo, Sumatra, Fed. Malay States, Siam, Andaman Islands, off domestic fowl, Bos bu- balis bubalis, goose, buffalo, dog, turkey). Sharif, 1928, Records Indian Mus., 30, pt. 3, pp. 267-268, fig. 17 (J1, J, Andaman Islands; J, Assam, off cock). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 142- 143, map 3; 1932, Veeartsenijk. Meded., No. 79, pp. 15-16, 33, figs. 16-17, map 3 (J1, J, Java, New Guinea, off hens). Schulze, 1939, Zeitschr. f. Parasitenk., 10, pt. 6, pp. 725- 726 (J1, 5, Burma, off Gallus gallus). Toumanoff, 1944, Les Tiques (Ixodoidea) de PIndochine, pp. 70-72, 88, 91, Pis. 53-54 (J*, J, Krong-Pha and Djiring in So. Annam, and Cambodge, off wild fowl). Male (Fig. 9A-F). Seven partially engorged, Sumatra speci- mens from 1.34 to 1.42 mm. long by 1.2 to 1.12 mm. wide, exclusive of capitulum. Scutum elongate-oval, narrowed anteriorly, yellow- brown to brown, very glossy; punctations small, slightly unequal, fairly numerous, larger, and less numerous on anterior third of scutum; cervical grooves deep and short; lateral groove on each side including first festoon and extending to region corresponding to pseudoscutum ; festoons longer than wide, with a few puncta- tions on anterior half. Body ventrally lighter, and finely striate; spiracle oval, with dorsal surface decidedly narrowed ; genital open- ing opposite coxa II; anal opening opposite posterior border of spiracles. Legs strong, with a double row of long, white hairs ven- trally; coxa I with a single, short spur; coxae II-IV with a single spur slightly shorter than that of coxa I ; tarsi short, tapering gradually, not terminating in spurs; pulvillus nearly as long as claws. Capitulum 0.35 mm. long; basis punctate, broader than long; cornua short and blunt; palpal articles nearly equal; article two strongly salient laterally; article three with a dorsal, retro- grade spur at inner angle and a strong, ventral spur; hypostome short ; dentition 4 : 4, with seven to nine teeth per file. Female (Fig. 9G-L). Unengorged, Sumatra specimen 1.75 mm. long by 1.26 mm. wide, exclusive of capitulum. Body oval, widest at mid-length. Scutum yellow-brown, oval, and slightly longer than wide, in twenty specimens from 0.77 to 0.91 mm. long by 0.71 to 0.81 mm. wide; punctations medium, numerous, and more numerous in posterior half of scutum; cervical grooves long, deep and parallel. Abdomen dark and uniformly covered with punctations; marginal groove distinct and long, including first 50 ENTOMOLOGICA AMERICANA three festoons on each side; festoons nearly as wide as long. Ven- tral body surface lighter, with numerous, medium punctations and a few, short, white hairs posteriorly; genital opening opposite coxae II and III ; anal opening opposite spiracles ; spiracle nearly circular. Coxa I with a long, slender spur ; coxae II and III with a shorter, broader spur ; coxa IV with a short, narrow spur. Ca- pitulum 0.49 mm. long; basis nearly three times as wide as long; posterior margin concave; porose areas oval, large, widely sep- arated, with a depression between them ; palps in general resemble those of male, but article two is slightly longer; hypostome short, widest at mid-length ; dentition 4 : 4, with approximately eleven teeth per file. Distribution and Hosts. Known from Borneo, Fed. Malay States, Siam, Andaman Islands, India, New Guinea, Burma, East Indies and Indochina. It has been recorded in the East Indies from Sumatra (Nuttall and Warburton, 1915) and Java (Krijgs- man and Ponto, 1931, 1932). Numerous males and females were seen from Sumatra (Lhokseumawe and Koetaradja, off domestic fowl, in Kraneveld collection). Specimens were also seen in the Rocky Mountain Laboratory from Myitkyina, Burma, taken off a great variety of birds and mammals. Genus Dermacentor C. L. Koch, 1844 Schulze established the genus Indocentor in 1933 for the species limited to India, China and the East Indies. He placed great stress on the shape of the male scutum in separating this genus, but in a series of twenty-three males in the Museum of Comparative Zoology, collected off one Jiost at Doi Angha, Siam, by H. Coolidge, Jr., the shape of the scutum varies from that characteristic of Indo- centor to that characteristic of Dermacentor. Some specimens in the series are intermediate between these two, and it is difficult to place them definitely in either group. The few differences that do exist are of specific rank only ; I, therefore, regard Indocentor Schulze as a synonym of Dermacentor. Dermacentor auratus Supino. Dermacentor auratus Supino, 1897, Atti Soc. Veneto-Trent. Sci. Nat., (2) 3, pt. 1, p. 235 ©, Carin Cheba, Mooleyit (Tenasserim), off “U rsus torquatus” = Euarctos thibet- anus thibetanus, Sus cristatus ; type in Genoa M.) ; 1897, Op. cit., (2) 3, pt. 2, pp. 25 0-25 i, PI. 13, fig. 14. 51 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 52 ENTOMOLOGICA AMERICANA After studying numerous specimens from India, Burma, Ma- laya, Siam, Sumatra, Java, New Guinea and the Philippine Is- lands, I can recognize with certainty only D. auratus Supino. The status of the other species and. subspecies described by Neumann and Schulze can not be settled definitely at this time; not only are their descriptions meager and insufficient, but illustrations are lacking or inadequate in the essential characters necessary for ac- curate determination. Due to the known, great variation in this genus, the only reliable characters to which some significance can be attached are the spurs on coxa I, and it is possible that this structure varies also. Only detailed breeding experiments will settle this problem of variation and definitely decide the number of valid species in this area. The following descriptions are based upon specimens in the col- lections studied. It was felt advisable to include the description and figures of males (Fig. 10A-F) collected near the type locality in Burma, and these are regarded as typical auratus. The single male (Fig. 11A-F) from Medan, Sumatra, probably represents a variation of auratus. Only one female (Fig. 10G-M) from Medan, Sumatra, was found to agree closely with specimens collected near the type locality, and it is regarded as typical auratus. Three females (Fig. 11G-M) from Sumatra and one female (Fig. 12A-G) from Java probably also represent variations of auratus. Male (Fig. 10A-F)i Scutum of two Burma specimens respec- tively 4.2 and 4.5 mm. long by 3.6 and 4.2 mm. wide, oval, nar- rowed anteriorly, widest at mid-length; brown markings all more or less raised ; pseudoscutum delimited by a series of fused, raised, brown stripes ; lateral margins partially set off by ocular stripe and paramedian stripes fused to form a long, narrow stripe ; posterior margin set off by a transverse stripe and two antero-accessory stripes set at right angles to it ; para-median stripes fused with an- tero-accessory stripes to complete lateral border; antero-accessory stripes extending a short distance behind transverse stripe ; a long, median, longitudinal stripe beginning near anterior margin of Fig. 10. Dermacentor auratus Supino; typical. A-F, male from Burma : A, scutum ; B, coxae I-IV ; C, tarsus IV ; D, capitu- lum in dorsal view; E, same in ventral view; F, spiracle. G-M, female from Sumatra : G, scutum ; H, coxae I-IV ; I, tarsus I ; J, tarsus IV ; K, capitulum in dorsal view ; L, same in ventral view ; M, spiracle. 53 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 scutum, passes through transverse stripe and continues posteriorly a short distance ; postero-median stripe short, nearly reaching me- dian, longitudinal stripe; postero-accessory stripes beginning in vicinity of third and fourth festoons, extended anteriorly a short distance and curving toward postero-median stripe; lateral spots very weak, indistinct; a dark brown, irregularly thickened band, beginning in region of eye, outlines margin of scutum and extends posteriorly to first festoon ; punctations numerous and unequal ; larger ones circular, deep and uniformly distributed; smaller ones numerous, but inconspicuous ; punctations lacking on all dark brown, raised areas; cervical grooves deep and comma-shaped an- teriorly, continued as shallow depressions posteriorly ; lateral grooves as a single row of large punctations beginning a short dis- tance behind eyes; festoons longer than broad and separated by a deep, wide groove ; festoons with varying amounts of white orna- mentation, third and fourth festoons on each side with least amount. Ventral body surface lighter and with a few, fine hairs; genital opening opposite coxa II ; anal opening opposite spiracles ; genital grooves nearly parallel, but widely divergent around anal opening ; spiracles comma-shaped and small. Legs large, strong, thick, or- nate and punctate ; coxa I with two widely separated, short, blunt spurs, with internal spur slightly wider ; coxae II and III with two subequal, widely separated, short, blunt spurs; coxa IV nearly twice as large as other coxae, with one external spur, and one to four internal spurs; tarsus I with a long, proximal part and a short, distal part ; tarsus IV tap.ering gradually, with three ventral spurs ; pulvillus long. Capitulum 1.0 mm. long by 0.8 mm. wide ; basis and palps ornate dorsally and finely punctate; basis rectan- gular and about twice as wide as long; cornua short and broad; palpal articles two and three nearly equal; article one only barely visible dorsally, being concealed by a retroverted prominence of article two ; ventrally article I with a large, triangular plate bear- ing several hairs ; hypostome short ; 3 : 3 dentition, with about nine to twelve teeth per file. Female (Fig. 10Gr-M). A single, Sumatra specimen 4.95 mm. long by 4.6 mm. wide, exclusive of capitulum, partially engorged, nearly oval, narrows slightly posterior to eyes and is widest at mid- length. Scutum 2.1 mm. long by 2.5 mm. wide, slightly wider than long and widest at anterior third ; all brown markings without punc- tations; a broad, dark brown margin, beginning as an ocular spot, continues completely around posterior margin of scutum; median 54 ENTOMOLOGICA AMERICANA stripe long, extending nearly to top margin of scutnm; cervical stripes on each side of median stripe bow-shaped, distinct and not reaching lateral margin of scntum; larger pnnctations numerous and deep ; smaller punctations in scapular and posterior regions ; very fine punctations uniformly distributed over scutum; cervical grooves deep and convergent anteriorly, but less so posteriorly; scutum with pronounced depression in region of cervical grooves; eyes flat and dark. Dorsal body surface with many, large puncta- tions from each of which arises a thick, white, appressed hair ; mar- ginal groove long, distinct, including three festoons and a portion of fourth on each side ; postero-median groove nearly reaching scu- tum ; para-median grooves about one-half length of median groove and obliquely placed; festoons distinct and slightly longer than wide ; white hairs in or near each groove separating festoons. Ab- domen ventrally lighter in color, with a few short, white hairs in region of genital aperture ; genital opening opposite coxa II ; anal opening opposite spiracles; genital grooves nearly parallel, but di- verging widely anterior to anal opening; spiracle triangular, with blunt angles. Legs brown, strong and thick, ornate, with white markings on dorsal surface of third and fourth pairs more pro- nounced ; coxa I with two, short, obtuse, well separated spurs ; in- ternal about twice as wide as external ; coxae II and III with two, shorter spurs, more widely separated than those of coxa I ; coxa IV with two short, sharp, nearly equal, and widely separated spurs having a very tiny spur between them ; tarsi II-IV tapering grad- ually and with three distinct, ventral spurs; pulvillus long. Ca- pitulum 1.3 mm. long and 0.9 mm. wide; enamel marking on basis and palps ; basis rectangular, with lateral margins slightly convex ; cornua distinct; porose areas oval, deep and with interval between them slightly less than their diameter ; palps relatively long ; article two longer than article three ; article one very small, partly fused with article two, only partially visible dorso-laterally ; ventrally with a large, prominent, triangular plate bearing three, long hairs on inner margin; hypostome short, lateral margins nearly parallel and dentition 3 : 3, with approximately fourteen to fifteen teeth per file. Male, variation (Fig. 11A-F). Unengorged Sumatra specimen, exclusive of capitulum, 3.9 mm. long by 3.0 mm. wide, oval, nar- rowed anteriorly and widest just anterior to spiracles, blackish- brown with pale, enamel markings; pseudoscutum slightly raised 55 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Fig. 11. Dermacentor auratus Supino; variation from Su- matra. A-F, male: A, scutum; B, coxae I-IV ; C, tarsus IV ; D, capitulum in dorsal view; B, same in ventral view; F, spiracle. G-M, female: G, scutum; H, coxae I-IV ; I, tarsus I; tarsus IV; K, capitulum in dorsal view ; L, same in ventral view ; M, spiracle. 56 ENTOMOLOGICA AMERICANA above scutum proper; punctations numerous and unequal, with finer punctations in scapular regions; cervical grooves as short, deep punctures; eyes small and dark yellow; lateral groove as a series of linearly arranged punctations, not including any festoons ; first two festoons on each side partially covered with white enamel, other festoons without enamel. Ventral body surface dark; pos- terior portion with many large, closely set punctations and long, white hairs ; genital opening opposite coxa II ; anal opening oppo- site posterior border of spiracles; spiracles large and comma- shaped; genital groove widely divergent around anal opening. Legs, dark, strong, enameled and finely punctate ; coxa I with two relatively long spurs, external spur narrower than internal; coxae II and III with two short, separated and subequal spurs; coxa IV not much larger than others and with two nearly equal, separated and triangular spurs. Capitulum 1.05 mm. long; palps short; article two slightly longer than article three ; hypostome short ; den- tition 3 : 3, with approximately twelve teeth per file. Female, variation (Fig. 11G-M). Scutum of three engorged, Sumatran specimens respectively 1.96, 2.1 and 2.17 mm. long by 2.24, 2.31 and 2.59 mm. wide, reddish-brown, with white enamel markings, antero-lateral margin excavated to receive legs, and pos- terior margin slightly sinuous; larger punctations deep and con- spicuous; smaller punctations more numerous in scapular angles; cervical grooves short, deep and convex externally ; eyes large, yel- low and oval. Abdomen uniformly covered with large, deep punc- tations; marginal groove distinct, deep, including first three fes- toons on each side ; festoons slightly longer than wide, with a few punctations on anterior half. Abdomen ventrally punctate and with short, white hairs ; genital opening opposite coxae II ; anal opening opposite spiracles ; spiracles triangular, with blunt, rounded angles. Tarsus I long, broad and swollen, ending in a single, short spur ; tarsi II-IV slender, tapering gradually, with three dis- tinct ventral spurs; coxa I with two very long, closely set spurs, internal broader; coxae II— III with two short spurs, the internal ridge-like ; coxa IV with two short, pointed spurs. Capitulum 1.28 mm. long ; basis rectangular ; cornua distinct ; porose areas large and divergent anteriorly ; hypostome slender ; dentition 3 : 3, with approximately thirteen teeth per file. Female, variation (Fig. 12A-G). Single, engorged, Java speci- men 7.5 mm. long by 6.0 mm. wide, exclusive of capitulum. Body 57 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 dark brown, oval and widest at region of spiracles. Scutum slightly broader than long, 2.1 mm. long by 2.55 mm. wide, reddish- brown, with extensive yellow-white enamel; larger more numerous punctations deep and circular; finer punctations in scapular re- gions, with very few in posterior portion; punctations not present on brown markings; cervical grooves as deep punctures anteriorly; eyes long, flat and dark brown. Abdomen punctate, with a long, white, conspicuous hair in each punctation; marginal groove nar- F Fig. 12. Dermacentor auratus Supino, female; variation from Java: A, scutum; B, coxae I-IV ; C, tarsus I; D, tarsus IV; E, capitulum in dorsal view ; F, same in ventral view ; G, spiracle. row ; festoons prominent and longer than wide ; genital opening opposite' coxa two ; anal opening opposite posterior border of spi- racles; spiracles triangular, with rounded angles. Legs strong, dark brown and covered with enamel; coxa I with two long spurs, external long, narrow and tapering, internal broader and quadran- gular ; coxae II, III, and IV each with two short, subequal, widely separated spurs ; tarsus I with a short, ventral spur; tarsi I I-IV long, tapering gradually, with three ventral spurs ; pulvillus about one-half as long as claws. Capitulum 1.26 mm. long by 0.72 mm. 58 ENTOMOLOGICA AMERICANA wide ; basis rectangular, with rounded sides ; cornua distinct ; porose areas large, oval and slightly divergent; article two longer than article three ; hypostome with 3 : 3 dentition, with approximately fifteen to seventeen teeth per file. Biology. The United States Army Typhus Commission col- lected many nymphs of auratus in Burma and reared them to the adult stage. These were taken off Viverricula indica thai, Rattus rattus, Cervus unicolor , Callosciurus quinquestriatus quinquestria- tus, Callosciurus sladeni midas, Ratufa gigantea gigantea, Gallus gallus gallus, man, guinea pig and monkey. Adults were taken off Arctonyx collaris , Felis tigris, Bus cristatus and Callosciurus quinquestriatus quinquestriatus. This species is also found on wild and domestic pigs. Though larvae have been recorded by Sharif in 1928 off man and deer and assigned to this species, they were neither described nor illustrated. This species normally attacks wild animals, but its occurrence on domestic animals and on man makes it a potential disease vector. Distribution and Hosts. This species is definitely known from India, Burma and Sumatra. Other records of Dermacentor , de- scribed under various specific names, require further study, and for this reason have been omitted from this work. All specimens studied were taken off domestic pigs. Genus Rhipicephalus C. L. Koch, 1844 Key to Species of Rhipicephalus Males 1. Anal plates triangular, with internal margin only slightly concave R. sanguineus Anal plates sickle-shaped, with internal margin very concave; external and posterior margins forming a regular curve 2 2. Scutum smooth. Hairs small and inconspicuous. R. haemaphysaloides haemaphysaloides Scutum rough and irregular. Hairs larger and more conspicu- ous R. haemaphysaloides pilans Females 1. Scutum elongate-oval, longer than wide and usually somewhat produced posteriorly. Punctations on scutum numerous, close set, irregularly distributed. Legs slender. R. sanguineus Scutum about as wide as long. Punctations on scutum unequal ; 59 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 finer ones seen only with higher magnification. Legs stont. ■ 2 2. Punctations on scutum less coarse. Hairs small and incon- spicuous. Abdomen with weak hairs, sparsely distributed. R. haemaphysaloides haemaphysaloides Punctations on scutum coarser. Hairs larger and more con- spicuous. Hairs on abdomen numerous, long thick and appressed R. haemaphysaloides pilans Rhipicephalus haemaphysaloides Supino. This species is divided into two subspecies, Rhipicephalus hae- maphysaloides haemaphysaloides Supino and Rhipicephalus haema- physaloides pilans Schulze, which form two distinct, geographical units. Zumpt in 1943 noted the presence of transitional specimens between the two subspecies on the island of Sumatra ; he decided that these transitional forms belonged to R. haemaphysaloides pilans and named the Straits of Malacca as the boundary dividing the two subspecies. I saw numerous specimens from eleven locali- ties in Sumatra, and I also found transitional specimens. Those from Upper Sumatra are closely allied to the typical form found in Burma, and those from Lower Sumatra are more closely allied to pilans. All of Zumpt ’s specimens probably came from Lower Sumatra. Both subspecies definitely occur in Sumatra and each is restricted to a definite portion of the island. The boundary be- tween the two subspecies is not the Straits of Malacca, but it is somewhere between Upper and Lower Sumatra. Since no speci- mens were collected in Central Sumatra, the complete range of each subspecies in this region can not be determined at this time. On no other island in this area are both forms known to occur. Rhipicephalus haemaphysaloides haemaphysaloides Supino. Rhipicephalus haemaphysaloides niger Supino, 1897, Atti Soc. Veneto-Trent. Sci. Nat., (2) 3, pt. 1, p. 234 (§, Yado, Burma; type in Genoa M.). Neumann, 1911, Das Tier- reieh, Lief. 26, Acarina, Ixodidae, p. 46 ( incertae sedis). Rhipicephalus haemaphysaloides ruber Supino, 1897, Atti Soc. Veneto-Trent. Sci. Nat., (2) 3, pt. 1, pp. 234-235 (d\ §; Mt. Mooleyit, Tenasserim; Meteleo, Thagata, Juva, Cago del Cadu Gianng, N.E. of Bomo, Burma; cotypes in Genoa M.) ; 1897, Op. cit ., (2) 3, pt. 2, p. 250, PI. 12, figs. 10-11 (, Lubok Paku, Sumatra, off tiger). Male (Fig. 13A-G). Thirty Sumatra specimens range from 3.9 to 2.9 mm. long by 2.64 to 1.8 mm. wide, exclusive of capitulum; average 3.38 mm. long by 2.16 mm. wide. Body elongate-oval, about one and a half times as long as wide, tapering toward an- terior end, with marked indentation in region of eyes, widest op- posite spiracular plates, reddish-brown to black; larger puncta- tions few and evenly distributed, with a small, white hair issuing 62 ENTOMOLOGICA AMERICANA Supino. A-G, male : A, scutum ; B, coxae I-IY ; C, capitulum in dorsal view; D, same in ventral view; E, ventral view of scutum j showing anal and adanal plates ; F, spiracle ; G, tarsus IV. H-M, female : H, scutum ; I, coxae I-IV ; J, tarsus IV ; K, spiracle ; L, capitulum in dorsal view ; M, same in ventral view. 63 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1 4 from each; finer punctations very numerous, but discernible only under higher magnification ; cervical grooves short, deep and slightly convergent; lateral grooves as furrows in hind part, with large punctations linearly arranged in each; median groove long and narrow ; paramedian grooves diagonal ; eyes prominent, yellow, flat and elongate-oval ; festoons protruding in engorged specimens ; median festoon as a pronounced, caudal protrusion ; body ventrally with numerous, large, white hairs; spiracular plates large and comma-shaped ; genital opening opposite coxa II ; anal opening op- posite border of spiracular plate ; anal plates sickle-shaped and heavily chitinised; adanal plates small, less chitinised and weakly developed. Legs stout and covered with long, white hairs; coxae large and conspicuous; coxa I with two large spurs, internal about twice as broad as external; coxae II and III with a short, external spur and a broad, ridge-like, scarcely visible, internal spur; coxa IV with internal and external spurs weakly developed; tarsus IV tapering gradually, ending in a terminal and subterminal spur. Basis capituli hexagonal, about twice as wide as long and salient laterally; cornua prominent and strongly projecting; palpal ar- ticles two and three together slightly longer than broad; distinct, lateral edge on palpal article three ; article one with a distinct, ventral plate containing six to nine large, white hairs; article two with a similar fringe, but with less hairs ; article three with a very short, ventral spur ; hypostome small ; 3 : 3 dentition, with approxi- mately nine to ten teeth per file. Female (Fig. 13H-M). Thirty unengorged, Sumatra specimens from 3.96 to 3.12 mm. long by 2.4 to 2.04 mm. wide, exclusive of capitulum; average 3.48 mm. long by 2.24 mm. wide; largest female 19.2 mm. long by 14.1 mm. wide. Body elongate-oval, about one and a half times as long as broad. Scutum slightly longer than wide, reddish-brown to black, posterior margin sinuous ; cer- vical grooves deep and diagonal ; well developed furrows, leading from cervical grooves, bow-shaped and divergent posteriorly ; large punctations conspicuous, more or less linearly arranged, with a short, white hair projecting from each; eyes as in male. Abdomen dorsally with small, thin, white hairs; spiracular plate nearly as wide as long, with a short, curved extension; body ventrally with numerous, short, white hairs and occasional longer ones; genital opening opposite coxa II ; anal opening opposite posterior border of spiracular plate. Legs similar to those of male ; coxae also simi- lar, but internal spur of coxa IV not very distinct. Capitulum 64 ENTOMOLOGICA AMERICANA more than twice as wide as long ; cornna not as strong as in male ; porose areas oval and slightly divergent anteriorly; palpal articles two and three together much longer than wide; article three with blunt angle on outer edge ; ventral surface of capitulum as in male ; hypostome dentition 3 : 3, with approximately nine teeth per file. Distribution and Hosts. Known from India, Burma, Ceylon, Indochina, southern China, Formosa. It is known in the East In- dies from Upper Sumatra (Neumann, 1897, 1904, 1911; Krijgsman and Ponto, 1931, 1932 ; Schulze, 1936). The following records from Sumatra are based on specimens seen in the Kraneveld collection : Koetaradja, off buffalo and cow; Lhokseumawe, off buffalo, cow and dog; Medan, off buffalo, cow, goat, pig and without host stated; Padangsidimpoean, off buffalo, cow, dog and pig ; Goenoengtoea, off buffalo. Numerous specimens from Burma were seen in the Bocky Mountain Laboratory taken off the following hosts : Muntiacus munt- jak vaginalis, Arctonyx collaris, Hystrix sub crist atus sub crist atus, Cervus unicolor, Felis tigris, Felis pardus, Gallus g alius g alius and Orange-breasted Flycatcher. A survey of the native fauna of Upper Sumatra would probably show a similarly wide host range. Biology. This species is indigenous to the Oriental Region and is found on a wide variety o-f native animals. Introduced domestic animals have proven to be suitable hosts, and the conditions of do- mestic life have hastened and facilitated the spread of this tick throughout the Oriental Region. It is second in importance to Boophilus microplus as a pest of livestock. Little is known about disease transmission by this species, but it should be regarded with suspicion as a possible vector. Though there are hundreds of records of this species taken in various localities, the larvae and nymphs have never been observed. In the large Kraneveld collection not one immature stage was found, and in the extensive survey made by the United States Typhus Commission in Myitkyina, Burma, no young forms were collected. It must be assumed that this is either a two-host tick or a three- host tick spending part of its life cycle on other animals, probably small rodents or birds. Rhipicephalus haemaphysaloides pilans Schulze. Rhipicephalus pilans Schulze, 1935, Wissenschaftliche Ergeb- nisse Exped. Karakorum, Zool., Acarina, Ixodoidea, p. 180 (description of male in key) ; 1936, Zeitschr. f. Parasitenk., S, pt. 5, p. 524, figs. 3-5 (2 Rana Mese, Flores; 1, J. 65 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Badjawa, Flores; 1 5, Swela, Lombok; holotype $ in Zool. M. Berlin). Rhipicephalus haemaphysaloides Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 142, map 2 (dis- tribution and hosts in Indonesia) [in part: not Upper Sumatra records] ; 1932, Veeartsenijk. Meded., No. 79, pp. 10, 32, fig. 7, map 2 (,c?\ ?> Lower Sumatra, Java, Ma- doera, Soembawa, Soemba, Sawoe, Timor, Alor, Roti, Bor- neo, off buffalo, cows, goats and dogs). Rhipicephalus haemaphysaloides paulopunctatus Schulze, 1932, Sitz.-Ber. Abhand. Naturf. Gesellsch. Rostock, (3) 3, p. 116, fig. 13 Java). Anastos, 1948, Psyche, 55, pt. 1, pp. 36-37 (5, Java, on another tick, B. microplus). Rhipicephalus haemaphysaloides pilaus Zumpt, 1940, Zeitschr. f. Parasitenk., 11, p. 675; 1943, Op. cit., 13, pt. 1, pp. 5, 6, 22-23, figs. 22-24 J, Greater and Lesser Sunda Islands, off (( Sus vittatus” = Bus cristatus vittatus, “Buf- felus bubalus” = Bos bubalis bubalis, Bos taurus and Felis tigris). Male ' (Fig. 14A-D). Thirty Java specimens average 3.09 mm. long by 2.09 mm. wide, exclusive of capitulum. Darker forms seen more frequently than lighter ones. Readily distinguished from R. haemaphysaloides haemaphysaloides by its rugose and coarsely sculptured scutum ; much deeper median and paramedian grooves ; very typical, thick, appressed, white hairs. In other features agrees very closely with typical form. Female (Fig. 14E-J). Thirty unengorged, Java females from 3.45 to 3.07 mm. long by 2.25 to 2.1 mm. wide, exclusive of capitu- lum ; average 3.19 mm. long by 2.8 mm. wide ; largest, engorged specimen 18 mm. long by 12.5 mm. wide. Sculpturing of scutum similar to male ; conspicuous, white hairs in each, deep punctation ; finer punctations more clearly visible than in typical form. Very characteristic covering of heavy, white, appressed hairs on abdo- men. Resembles otherwise closely female of typical form. Distribution and Hosts. Known from Lower Sumatra, Madoera, Soembawa, Soemba, Sawoe, Roti, Timor, Alor, Celebes and Borneo (Krijgsman and Ponto, 1931, 1932), Flores, Lombok (Schulze, 1936), Java (Krijgsman and Ponto, 1931, 1932; Anastos, 1948). Numerous specimens were seen in the Kraneveld collection from Su- matra (Loeboeklinggau, off buffalo; Palembang, off cow; Ben- koelen, off buffalo, cow and dog; Lahat, off cow; Tandjoengkarang, 66 ENTOMOLOGICA AMERICANA A-D, male : A, scutum ; B, coxae I-IV ; C, capitulum in dorsal view ; D, same in ventral view. E-J, female: E, scutum; F, coxae I-IV ; G, tarsus IV ; H, spiracle; I, capitulum in dorsal view; J, same in ventral view. 67 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 off buffalo, dog and goat), Java (Serang, off sheep; Batavia, off buffalo and cow; Buitenzorg, off buffalo and cow; Tegal, off dog; Madioen, off cow; Bodjonegoro, off buffalo and sheep; Dj ember, off pig), Madoera (Pamekasan, off cow; Soemenep, off buffalo, cow, goat, and horse), Bali (Singaradja, off cow and horse), Soembawa (Soembawabesar, off buffalo), Soemba (Waingapoe, off buffalo, cow and dog), Celebes (Watampone, off buffalo; Sengang, off goat; Parepare, off buffalo, cow, dog and sheep ; Makale, off buffalo and dog). Numerous specimens were seen from the Philippine Islands in the collection of the Rocky Mountain Laboratory. Most of the records are from domestic animals, with a few from wild animals. It is likely that the host list for this subspecies will pfove to be as extensive as that of the typical form. Rhipicephalus sanguineus (Latreille). Ixodes sanguineus Latreille, 1806, Gen. Crust. Ins., 1, p. 157 (no sex, France, no host). Rhipicephalus sanguineus C. L. Koch, 1844, Arch. f. Natur- gesch., 10, pt. 1, p. 238 (merely lists species). Nieschulz, 1924, Tijdschr. v. Diergeneesk., 51, p. 195 (Sumatra, off native cattle). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, p. 141, map 2 (Java, Soemba, Ma- doera, Sumatra, Timor, Alor Island, Amboina and Sapa- roea, off man, cattle, dog, buffalo) ; 1932, Veeartsenijk. Meded., No. 79, pp. 8-10, 32, figs. 5-6, map 2 (J1, §, hosts, distribution in Indonesia). Rhipicephalus sanguineus sanguineus Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 35-36, figs. 16- 17 (world- wide distribution, including Sumatra and Java) . ? Rhipicephalus rubicundus Frauenfeld, 1867, Verhandl. Zool. Bot. Gesellsch. Wien, 17, p. 462 (J1, taken aboard ship in Sunda Sea). Because of its wide distribution this species has been the subject of much discussion ; only references to the original description and to the East Indies are included here. Male (Fig. 15A-G). Twenty Java specimens from 2.31 to 2.25 mm. long by 1.36 to 1.35 mm. wide, exclusive of capitulum; average 2.28 mm. long by 1.35 mm. wide. Body elongate-oval, widest near region of spiracles, reddish-brown ; punctations more numerous an- teriorly, with a very short, inconspicuous hair arising from each; 68 ENTOMOLOGICA AMERICANA cervical grooves short, deep and convex externally; median and two paramedian grooves not very deep; a small pit slightly an- terior to each paramedian groove; lateral grooves beginning pos- terior to eyes and including first festoon on each side; eyes oval and slightly convex; festoons about as long as wide, with a few punctatio'ns. Ventral body surface lighter than dorsal; spiracles very long and comma-shaped ; genital opening opposite coxa II ; anal opening opposite spiracles ; anal plates triangular and mildly concave on internal border; adanal plates short, sharp points. Legs short, thick and hirsute ; coxa I with two long, closely-set spurs, internal broader ; coxae II and III with a very short, external spur, inner corner salient ; coxa IV with very short, internal and external spurs ; tarsus I long, without ventral spurs and sloping gradually ; tarsi II— IV shorter, with a strong, ventral, terminal spur and a subterminal spur. Capitulum 0.51 mm. long; basis hexagonal, lat- eral edges salient, posterior margin nearly straight ; cornua distinct ; article I ventrally with a distinct plate ; long hairs on articles I and II ; hypostome short ; dentition 3 : 3, with seven to eight teeth per file. Female (Fig. 15H-M). Ten unengorged, Java specimens from 2.55 to 2.1 mm. long by 1.65 to 1.35 mm. wide, exclusive of capitu- lum; largest, engorged specimen 10.2 mm. long by 6.6 mm. wide. Reddish-brown scutum 1.2 mm. long by 1.2 mm. wide ; punctations large, with a short, white hair in each ; cervical grooves long, deep and bow-shaped; eyes large and oval. Abdomen with long, white hairs; marginal grooves distinct, including first three festoons on each side ; median and paramedian grooves long and narrow ; abdo- men ventrally lighter in color and with long hairs ; genital opening opposite coxa II ; anal opening opposite spiracles ; spiracle shorter than in male. Legs long and thin; tarsi and coxae as in male. Capitulum 0.63 mm. long; basis hexagonal, broader than in male; cornua distinct ; porose areas small, oval and slightly divergent an- teriorly ; hypostome dentition 3 : 3, with approximately ten teeth per file. Distribution and Hosts. This is probably the most widely distrib- uted of all ticks, being known throughout the world. Th^ known hosts are numerous and include both domestic and wild animals. It is known in the East Indies from Sumatra (Neumann, 1911 ; Nieschulz, 1924; Krijgsman and Ponto, 1931, 1932), Java (Neu- mann, 1911; Krijgsman and Ponto, 1931, 1932), Soemba, Madoera, Timor, Alor, Amboina and Saparoea (Krijgsman and Ponto, 1931, 1932). Numerous specimens were seen in the Kraneveld collection 69 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 70 ENTOMOLOGICA AMERICANA from Sumatra (Lhokseumawe, off goat; Padangsidimpoean, off pig), Java (Tegal, off dog; Semarang, off dog; Toeban, off dog; Madioen, off dog; Bodjonegoro, off dog; Blitar, off dog; Djember, off dog; Bondowoso, off dog), Madoera (Pamekasan, off cow), Bali (Singaradja, off cow, and dog), Soembawa (Soembawabesar, off dog), Celebes (Manado, off cow). Biology. All stages in the life cycle are known. This tick is implicated in disease transmission both experimentally and nat- urally. It is capable of transmitting Rocky Mountain spotted fever to experimental animals, and.it is a known vector in nature of canine babesiasis and bovine anaplasmosis. Genus Boophilus Curtice, 1891 In 1934 Minning established the subgenera Uroloophilus and Palpohoophilus. These are based on tenuous characters, and I re- gard them as synonyms of Boophilus. Boophilus microplus (Canestrini) . Haemophysalis micropla [sic] Canestrini, 1888, Atti Soc. Veneto-Trent. Sci. Nat., 11, pt. 1, (1887), pp. 104-105, 110, PL 9, figs. 3, 3a-d and 5, 5a-b (Chaco australe, Paraguay; type in Genoa M.). Rhipicephalus annulatus Neumann, 1897, [ nec Say, 1821] Mem. Soc. Zool. France, 10, p. 412 [in part : only records from Timor, off “ Husa [sic] equina” = Cervus unicolor equinus ; Sumatra] . Rhipicephalus australis Fuller, 1899, Queensland Agric. Journ., 4, pt. 5, pp. 389-392, figures on pp. 391, 392 (Australia, off horses and cattle). Koningsberger, 1900, Teysmannia, 11, pt. 1, pp. 59-61, figs. 1-6 (J\ 5, n, 1, Indonesia). Boophilus australis Stiles and Hassall, 1901, Circ. 34, Bur. Anim. Industry, U. S. Dept. Agric., pp. 2-3. Salmon and Stiles, 1902, 17th Ann. Rept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), pp. 417, 420, 422, 426-433, figs. MO- 151, 153d, 154c, PL 80, figs. 114-139. Nieschulz, 1924, Fig. 15. Rhipicephalus sanguineus (Latreille). A-G, male: A, scutum ; B, coxae I-IY ; C, abdomen in ventral view, showing anal and adanal plates; D, tarsus IV ; E, capitulum in dorsal view; F, same in ventral view ; G, spiracle. H-M, female : H, scutum ; I, coxae I-IV ; J, tarsus IV ; K, spiracle ; L, capitulum in dorsal view ; M, same in ventral view. 71 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Tijdschr. v. Diergeneesk., 51, p. 195. Sharif, 1928, Records Indian Mus., 30, pt. 3, pp. 284-289, figs. 27-30. Bhipicephalus annidatus var. australis Neumann, 1901, Mem. Soc. Zool. France, 14, p. 280 ( J1, Borneo, off deer ; Su- matra, off buffalo ; 2, Sumatra ; 4 J1, 2 n, New South Wales, off cattle ; 5, Singapore, doubtfully) . Boophilus microplus Bequaert, 1926, Med. Report H. Rice 7th Exped. Amazon (1924-1925), pp. 168-171, figs. 1-2. .Anastos, 1948, Psyche, 55, pt. 1, pp. 36-37 (Batavia, Java, off cow). Boophilus annulatus [ nec Say, 1821] Doeve, 1923, Nederl. Ind. Bladen Diergeneesk., 34, pt. 1, pp. 4^18, figs. 1-2 (Watam- pone, Celebes). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 140, 141, map 1; 1932, Veeartse- nijk. Meded., No. 79, pp. 6-7, 32, figs. 3-4, map 1 (<$, 2, wide distribution in Indonesia, off buffalo, cow, horse, goat, deer, Cervus unicolor russa) . Boophilus ( TJ roboophilus ) krijgsmani Minning, 1934, Zeitschr. f. Parasitenk., 7, pt. 1, pp. 9, 11, 29-30, 40, figs. 23-24 (J', 2, Celebes, Soemba, Java, Borneo, Timor, Sangihe, Suma- tra, Soela Island, Bali, New Guinea, off cattle ; holotype 2 in Z. M. Berlin). Boophilus {TJ roboophilus) rot undiscut at us Minning, 1934, Op. cit., 7, pt. 1, pp. 9, 10, 30-31, 40, figs. 25-26 ( pt. 5, pp. 28-29 (5, Tandjoeng Keling, Sumatra). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 143, map 4 (J', 5, Java, off Varanus salvator and Mabuya multifasciata) ; 1932, Yee- artsenijk. Meded., No. 79, pp. 23, 34, 35, figs. 29-30 (J\ J, Java). Schulze, 1933, Arch. f. Hydrobiol., Suppl.-Bd. 12, 96 ENTOMOLOGICA AMERICANA pp. 495-496, 497, 498, figs. 3-4 (figures from type; 1 5, Leyte, P. I. ; 7 J1, Komodo, off Varanus komodoensis ; 1 Endeh, Flores, off Varanus salvator). Toumanoff, 1944, Les Tiques (Ixodoidea) de l’Indochine, pp. 115, 118, 120, Pis. 82, 84. Amblyomma decoratum C. L. Koch, 1844, Arch. f. Naturgesch., 10, pt. 1, pp. 230-231, No. 40 ( Philippine Is. ; type in Berlin M.) ; 1847, Uebersicht des Arachnidensystems, 4, pp. 19, 94-95, PI. 18, fig. 67. Neumann, 1899, Mem. Soc. Zool. France, 12, pp. 203, 245 (5 J1, Philippine Is.) ; 1901, Op. cit ., 14, p. 304. Salmon and Stiles, 1902, 17th Ann. Kept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), p. 473. Neumann, 1910, Ann. Sci. Nat. Zool., 9, pp. 175-176 (4 J1, Manila, Luzon; 1 J1, off Varanus salvator , locality unknown; 1 §, Padang, Sumatra, off Varanus salvator). Warburton, 1910, Parasitology, 3, pt. 4, p. 396 (Zool. Gar- dens, Calcutta, off Geoemyda grandis ; Nicobar Islands, off Varanus salvator). Neumann, 1911, Das Tierrelch, Lief. 26, Acarina, Ixodidae, pp. 56, 57, 64, 67, 87 (J1, 5, Philip- pine Islands, Java, Sumatra, off “Zamenis mucosus” = Ptyas mucosus, Varanus salvator, Python reticulatus) . Amblyomma fimbriatum C. L. Koch, 1844, Arch. f. Natur- gesch., 10, pt. 1, p. 231, No. 42 (J\ Manila) ; 1847, Ueber- sicht des Arachnidensystems, 4, pp. 20, 95, PI. 18, fig. 68. Amblyomma quadrimaculatum Neumann, 1899, Mem. Soc. Zool. France, 12, pp. 202, 245-246 (1 (Cf, Java, off “ Trigly - phedon dendrophilum” = Boiga dendrophila ; type de- scribed from Oudemans Coll., now at Leiden M.) ; 1901, Op. cit., 14, p. 304 (4 J' and 2 mutilated f, Java, off Python ' reticulatus ). Salmon and Stiles, 1902, 17th Ann. Kept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), p. 473. Schulze, 1933, Arch. f. Hydrobiol., Suppl.-Bd. 12, pp. 492, 495, 497, 498, figs. 5-6 (5 5, Ranau, Sumatra, off Varanus salvator; 2 Java off ((Varanus bivitt at us” = Varanus salvator, 1 5, Edi, Sumatra; 3'^, Java, off e< Dipsadomor- phus dendrophilus” = Boiga dendrophila) ; 1943, Zeitschr. Morph. Oekol. Tiere, 39, pt. 2, p. 355, fig. 38 (Sumatra; morphological). Buitendijk, 1945, Zoolog. Meded. Leiden, 24, pts. 3-4, p. 285 (type; locality here given as Pulau Berhala). Amblyomma furcosum Neumann, 1901, Mem. Soc. Zool. France, 97 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 14, pp. 299-300 (2 Java, off Python reticulatus ; type in Berlin M.) . Amblyomma ( Aponomma ) tenimberense S. and L. Hirst, 1910, Ann. Mag. Nat. Hist., (8), 6, pp. 307-308, fig. 9 ( Java, off Manis javanica). Toumanoff, 1944, Les Tiques (Ixodoidea) de lTndochine, pp. 104-105, 118, 120, Pis. 72-74 (lCf, 2? Cambodge and Cochin China, off bear, Sambar deer and wild pig). Aponomma latum, Neumann [nec Koch, 1844] 1901, Mem. Soc. Zool. Prance, 14, pp. 291-292, 344 (4 1 2? India, off serpent) [in part: equals Aponomma politum Neumann, 1899, syn.]. Howard, 1908, Ann. Transvaal Mus., 1, pt. 2, p. 153 (after Neumann, 1901). Neumann, 1911, Das Tier- reich, Lief. 26, Acarina, Ixodidae, p. 97 (uncertain spe- cies, Port Natal (Natal), India, off Python molurus) [in part : equals Aponomma politum Neumann, 1899, syn., ex errore, fide G. Theiler, 1945, The Onderstepoort Journ. Vet. Sci„ 20, pt. 2, p. 182]. Amblyomma badium Neumann, 1901, Mem. Soc. Zool. France, 14, pp. 300-301 (3 Sumatra; 6 lCf, off Manis sp. without locality; 19 6 2? 6 n, Java, off Manis javanica ; cotypes in Z. M. Berlin and in Oudemans Coll, now in Leiden M.). Aponomma javanense Cooper and Robinson, 1908, Proc. Cam- bridge Philos. Soc., 14, pp. 464-466, figs. 15-17 (3 2> Biserat, Siam, off Varanus salvator). Amblyomma javanense Schulze, 1934, Zeitschr. f. Parasitenk., 7, pt. 2, p. 167 (6 J1, 1 2, 1 n, Pangrango, Java, off Manis javanica) ; 1936, Zeitschr. Morph. Oekol. Tiere, 32, pt. 2, p. 206, fig. 23c (morphological) ; 1938, Op. cit., 34, pt. 1, p. 139, figs. 3, 7 (2, Hainan) ; 1939, Zeitschr. f. Parasitenk., 10, pt. 6, p. 728 (Burma). 103 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Fig. 22. Amblyomma javanense (Supino). A-F, male:, A, scutum B, coxae I-IV ; C, capitulum; D, spiracle; E, tarsus I; F, tarsus IV. G-L, female : G, scutum ; H, coxae I-IV ; I, capitulum in dorsal view ; J, spiracle ; K, tarsus I ; L , tarsus IV. 104 ENTOMOLOGICA AMERICANA Amblyomma javanense javanense Schulze, 1937, Op. cit., 9, pt. 6, pp. 690-692, fig. 2. Amblyomma compressum javanense Schulze, 1941, Zeitschr. Morph. Oekol. Tiere, 37, pt. 3, p. 514 (morphological). In 1937 Schulze believed that a relationship existed between the ticks found on the ant-eaters of Africa and of South Asia, and he regarded them as subspecies. Amblyomma javanense is the tick found on the Asiatic ant-eaters and Amblyomma cuneatum occurs on African ant-eaters. Schulze used the name A. javanense java- nense (Supino) for the Asiatic form and A. javanense cuneatum Neumann for the African form. In 1941 to further complicate matters he decided that A. compressum (Macalister) was an older name for A. cuneatum Neumann. In giving the former name priority he changed A. javanense javanense to A. compressum java- nense. I examined specimens of A. cuneatum in the Museum of Comparative Zoology from the Gold Coast, West Africa; it is prob- able that cuneatum and javanense had a common origin, but they are so distinct now that they must be regarded as separate species. Since no specimens are available from Indonesia, specimens from the Philippine Islands are used for the description of the male and the female. „ Male (Fig. 22A-F). Seven specimens from 5.4 to 4.8 mm. long to 4.8 to 4.05 mm. wide, exclusive of capitulum; average 5.2 mm. long by 4.52 mm. wide. Scutum inornate, maroon-brown in periph- eral regions and yellow-brown in central region; punctations un- equal, sparsely distributed in periphery and almost entirely absent in anterior and median portion ; lateral groove as linearly arranged punctations extending two-thirds distance to eyes ; cervical grooves very small and inverted comma-shaped ; few, short, white hairs in region of festoons; eyes small and indistinct; festoons large and distinct. Ventral body surface yellow, finely wrinkled, with nu- merous, fine punctations, and very short, indistinct hairs; genital opening opposite coxa two; anal opening opposite posterior border of spiracle; anal groove very distinct; spiracular plate long and comma-shaped. Legs brown-maroon, strong and relatively short; coxa I with two short, broad and widely separated spurs ; coxae II- IV with a single very broad and flat spur ; tarsi short, terminating abruptly and with prominent terminal and sub-terminal spurs; claws very long, nearly three times as long as pulvillus. Capitulum relatively short; basis rectangular, with lateral edge slightly con- vex ; palps short and club-shaped ; article two about one and a half 105 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 times as long as article three ; hypostome short and broad ; dentition 3 : 3, with six to eight teeth per file. Female (Fig. 22G-L). Largest of four slightly engorged speci- mens, exclusive of capitulum, 6.0 mm. long by 4.8 mm. wide, small- est 5.0 mm. long by 4.2 mm. wide. Body nearly circular, nar- rowed slightly in region of eyes. Scutum dark brown, inornate, from 2.4 to 2.1 mm. long by 3.0 to 2.5 mm. wide, average 2.19 mm. long by 2.8 mm. wide, cordiform with rounded angles, slightly wider than long and with postero-lateral margins very slightly concave ; punctations numerous, shallow, small to medium and evenly distributed; cervical grooves small and inverted comma- shaped; marginal groove distinct, ending near scutum; eyes small and often indistinct. Abdomen with scattered, medium puncta- tions and white hairs; festoons prominent, slightly longer than wide. Body ventrally with numerous, medium punctations and with short, white hairs on posterior two-thirds ; genital opening op- posite coxae II— III ; anal opening opposite posterior margin of spiracular plate; spiracular plate comma-shaped, slightly broader than that of male. Legs more slender than those of male; coxal spurs as in male. Basis wide, with sides slightly convex; porose areas deep, oval, small and divergent anteriorly; palps short and club-shaped ; hypostome short ; dentition 3 : 3, with eight to ten teeth per file. Distribution , Hosts and Biology. This species is widely distrib- uted in the Orient, and is known from Siam,' China, Tenasserim, Burma, Ceylon, Indochina, Fed. Malay States, India and the East Indies. It has been recorded from Sumatra (Neumann, 1901) and Java (Neumann, 1901; Krijgsman and Ponto, 1931, 1932; Schulze, 1934). No specimens were seen from Indonesia, but I saw five lots of many males, females, nymphs and larvae from the Philippine Islands in the Rocky Mountain Laboratory. In addition I saw in the Museum of Comparative Zoology two females taken off Manis sp. in China. Most of the known host records for this tick are from species of Manis, but it occasionally attacks reptiles. Neumann in 1899 recorded a male off Python molurus ; Cooper and Robinson in 1908 recorded it off Varanus salvator ; and Sharif in 1928 recorded it off Geoemyda tricarinata in India. Robinson in 1926 recorded a nymph off Hyaena hyaena in India. All stages are found on Manis, and A. javanense is apparently a one host tick. 106 ENTOMOLOGICA AMERICANA Amblyomma malayanum Neumann. Amblyomma malayanum Neumann, 1908 (March 20), Arch, de Parasit., 12, pp. 14-16, 26, figs. 9-10 (1 J1, 2 J, Buki- tima, Singapore; types in British M.) ; 1908, Notes Leyden Mus., 30, p. 86 (1 <£, 1 Malayan Peninsula, off tortoise). Blanchard, 1909, L’Insecte et 1 ’Infection, Fasc. 1, Aca- riens, p. 132. Neumann, 1911, Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 56, 59, 64, 68, 89. Amblyomma caelaturum Cooper and Robinson, 1908 (July 16), Proc. Cambridge Philos. Soc., 14, pt. 5, pp. 460-462, figs. 6-9 (2 1 J, Kwala Aring, Java [sic], off Geoemyda spinosa; cotypes in Brit. Mus.). Robinson, 1926, Ticks, 2, pt. 4, Amblyomma , pp. 17, 21, 227-230, figs. 111-112 (type locality corrected to Kuala Aring, Federated Malay States; 2, Malay Peninsula, off tortoise). Krijgsman and Ponto, 1931, Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 144, map 4; 1932, Veeartsenijk. Meded., No. 79, pp. 23, 35, figs. 31-32, map 4 (Sumatra, off reptiles). Schulze, 1932, Zeitschr. Morph. Oekol. Tiere, 25, pt. 2-3, pp. 510, 511, fig. 2 (1) (morphological) ; 1936, Meddel. Zool. Mus., Oslo, No. 13, p. 160 (morphological). Toumanoff, 1944, Les Tiques (Ixodoidea) de l’lndochine, pp. 116, 119, Pis. 82, 84. Amblyomma caelaturum caelaturum Schulze, 1932, Zeitschr. Morph. Oekol. Tiere, 25, pts. 2-3, p. 512. Amblyomma caelaturum perfectum Schulze, 1932, Zeitschr. f. Parasitenk., 4, pt. 3, pp. 469-470, fig. 6 (4 ^ , Soekaranda, Sumatra) ; 1932, Zeitschr. Morph. Oekol. Tiere, 25, pts. 2-3, pp. 512, 520. ? Amblyomma cyprium Toumanoff, 1944 [in part : nec Neu- mann, 1899] Les Tiques (Ixodoidea) de l’lndochine, pp. 106-111, PI. 78, figs, a, b, PI. 79 [only description and figures of female, nec male] . Neumann described A. malayanum in 1908 from one male and two females from Bukitima, Singapore, in the British Museum. This description was published March 20, 1908. He also recorded in April, 1908, another male and female from the Malayan Penin- sula collected by H. E. Durham. Cooper and Robinson described this same species in 1908 as Amblyomma caelaturum from a male and a female collected at 107 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Kuala Aring, Federated Malay States. The date of publication for this name was July 16, 1908. Robinson in his monograph of the genus in 1926 incorrectly gave priority to A. caelaturum. The name proposed by Neumann is the ’oldest name, and it has priority over that proposed by Cooper and Robinson. In 1932 Schulze divided this species into two subspecies, A. caelaturum caelaturum and A. caelaturum perfectum. His new subspecies is based upon four males from Soekaranda, Sumatra, and it differs from the typ- ical form in the relation between musculature and scutum. The male of perfectum is supposed to have porose areas like the female. These characters are tenuous, and the claim that the male of this species has the porose areas, which are attributes of female ticks only, is incorrect. I examined specimens in my possession, and though I found the posterior portion of the basis capituli to be finely punctate, in no way does it bear any resemblance to the porose areas. Similar punctation also occurs on the basis of A. cyprium. Toumanoff in 1944 has figured as the female of A. cyprium a female resembling A. malayanum; however, it differs in the shape of the scutum, the porose areas, the spiracle and the spur of coxa IY. Since specimens are not available for study from Indonesia, material from the Philippine Islands is used for the description of the male and female. Male (Fig. 23A-F). Two unengorged specimens respectively 5.55 and 6.66 mm. long by 4.95 and 6.15 mm. wide. Scutum nearly circular, slightly longer than wide and narrowed anteriorly in re- gion of the eyes; pseudoscutum well defined and lighter than scu- tum; punctations medium, numerous, deep and slightly confluent; pale, enamel markings anterior to the eyes, and a very small, pale marking in posterior angle of pseudoscutum ; cervical grooves short, deep and extending as elongated, shallow, wide depressions pos- teriorly; a median, narrow, longitudinal elevation beginning near median festoon and continuing anteriorly toward pseudoscutum ; a narrow, smooth elevation beginning opposite second and third fes- toons on each side and continuing obliquely toward median eleva- tion; three elevations anterior to these oblique elevations on each side ; all elevations free of punctations ; lateral groove composed of numerous linearly arranged punctations; eyes brown, small, flat, and indistinct; festoons longer than wide. Ventral body surface with many, fine wrinkles and numerous, small punctations; short, white hairs restricted to posterior third; spiracular plates very 108 ENTOMOLOGICA AMERICANA Fig. 23. Amblyomma malayanum Neumann. A-F, male : A, scutum; B, coxae I— TV ; C, tarsus IV; D, capitulum; E, spiracle; F, tarsus I. G-K, female: G, scutum; H, coxae I-IV ; I, capitulum in dorsal view ; J, dorsal foveae ; K, spiracle. 109 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 large and broad; genital opening opposite coxa II; anal opening opposite posterior portion of spiracles. Legs fairly slender, long, and lighter in color than body ; pale markings dorsally on distal end of femur, tibia and protarsus ; coxae small ; coxa I with two short, nearly equal spurs ; remaining coxae with a single, short spur ; tarsi II-IV long and with prominent terminal and sub-terminal spurs. Capitulum relatively short; basis rectangular, with pos- terior margins slightly salient ; hypostome short and slightly spatu- late ; dentition 4:4; palps short and club-shaped. Female (Fig. 23G-K). Largest of two engorged specimens 12 mm. long, 9.9 mm. wide and 6 mm. thick. Triangular scutum from 2.7 to 2.7 mm. long by 3.6 to 3.48 mm. wide, angles rounded, with posterior angle very broad, sides nearly straight, antero-lateral mar- gins prominent; brownish-maroon, a small pale marking in each scapular angle, and a rounded spot in posterior angle ; punctations numerous, deep, unequal and uniformly distributed, with those in region of eyes slightly larger ; cervical grooves deep, prominent anteriorly, enlarging into wide, shallow depressions posteriorly; eyes small, pale and not very prominent. Body reddish-brown, with many, small punctations and short, thick, white hairs; dorsal foveae very large, ovoid, slightly elongate transversely, 0.30 mm. long by 0.35 mm. wide, with surface hard, shining and dotted with very fine punctures ; festoons about as long as broad, with posterior half free of hairs. Ventral body surface similar to that of male; spiracles large and broad. Legs and coxae as in male. Gapitulum broader than long, with posterior border slightly concave; porose areas large, deep, oval, slightly divergent in front, with interval between porose areas slightly wider than their diameter ; hypostome long, with 4 : 4 dentition ; palps longer than those of male ; article two slightly over twice as long as article three. Distribution and Hosts. This species, which is rare in collections, is known from Bukitima (Singapore )r Kuala Aring (Federated Malay States), and Soekaranda (Sumatra). Two males and two females were seen in the Museum of Comparative Zoology. These were taken off a tortoise in the Philippine Islands, and where host records are known, all are from tortoises. Amblyomma robinsoni Warburton. Amblyomma robinsoni Warburton, 1927, Parasitology, 19, pt. 4, pp. 408-409, PI. 27, figs. 1-2 (8 Sumatra, Bali, Lombok, Soemba, Java, Borneo, off buffalo, cow, man, and “Sus vittatus ” = Sus cristatus vittatus). Sugimoto, 1935, Taiwan-No-Chikusan, 3, pt. 114 ENTOMOLOGICA AMERICANA 9, p. 7 ; 1936, Op. cit., 4, pt. 2, pp. 4—7, Pl. 6, figs. 1-7, PI. 8, figs. 1—4 ; 1936, Journ. Cent. Soc. Vet. Med., Tokyo, 49, pt. 7, p. 584, pl. 3, figs. 1-7 ; 1936, Journ. Soc. Trop. Agric., Tokyo, 8, pt. 4, p. 337 (Formosa). Ognra, 1936, Mem. Fac. Sci. Agric., Taihoku Imp. Univ., Formosa, 19, pt. 2, pp. 79-81, figs. 13-18 (,J\ 5> Formosa, off cattle, water buffalo, pig and wild boar). Sngimoto, 1937, Trans. Nat. Hist. Soc. Formosa, 27, No. 160, pp. 1-6, Pl. 1, figs. 1-6, Pl. 2, figs. 1-5 (nymph and larva from Formosa) ; 1937, Journ. Jap. Soc. Vet. Sci., 16, pt. 1, pp. 11-16, Pl. 5, figs. 1- 6, Pl. 6, figs. 1-5; 1937, Journ. Cent. Soc. Vet. Med., Tokyo, 50, pt. 5, pp. 318-322 (synonymy, distribution, hosts, Formosa) ; 1937, Journ. Soc. Trop. Agric., Tokyo, 9, pt. 3, p. 287. Sharif, 1938, Indian Journ. Vet. Sci. and Anim. Husb., 8, pt. 4, p. 363 (disease transmission). Tou- manoff, 1944, Les Tiques (Ixodoidea) de lTndochine, pp. 102-104, 119, 120, Pis. 69-71, 75 (,J\ 5, Indochina, off Sambar deer, tiger cat, wild pig, dog, horse ; 1 n,. off wild cock). Amblyomma infestum C. L. Koch, 1844, Arch. f. Naturgesch., 10, pt. 1, p. 226, No. 12 (J', 5, Bintan Island; types in Berlin M.) ; 1847, Uebersicht des Arachnidensystems, 4, pp. 17, 68-70, Pl. 12, figs. 41-42. Schulze, 1935, Zool. Anz., 112, pts. 9-10, p. 234; 1939, Zeitschr. f. Parasitenk., 10, pt. 6, p. 728 (Burma). Ixodes auriscutellatus Konigsberger, 1901, Teysmannia, 11, pt. 1, pp. 61-62, figs. 7-8 (J\ J, Bandong and Buitenzorg, Java, off buffalo; location of types unknown). Amblyomma compactum Neumann, 1901, Mem. Soc. Zool. France, 14, pp. 296-297, 342 (2 J, Sumatra; cotypes in Berlin M.). Amblyomma infestum testudinarium Schulze, 1932, Zeitschr. f. Parasitenk., 4, pt. 3, pp. 467-469 (4 .J', 4 5, Soekaranda, Sumatra) ; 1932, Zeitschr. Morph. Oekol. Tiere, 25, pts. 2- 3 pp. 528-529 (morphological) ; 1941, Op. cit., 37, pt. 3, p. 514 (morphological). Amblyomma infestum infestum Schulze, 1932, Zeitschr. f. Para- sitenk., 4, pt. 3, pp. 468, 469 (based on Koch’s types from Bintan Island near Singapore). Bequaert, 1933, Psyche, 40, pt. 4, pp. 140-142 (1 $ 3 J Bernam River, Selangor, Fed. Malay States, off Rhinoceros sumatrensis) . Schulze, 1936, Zeitschr. Morph. Oekol. Tiere, 32, pt. 2, pp. 204, 206, 207, fig. 19 b (morphological). 115 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Amblyomma fallax Schulze, 1932, Zeitschr. f. Parasitenk., 4, pt. 3, p. 468 ( nomen novum for A. testudinarium as de- scribed and figured by Robinson, 1926, pp. 253-257). Be- quaert, 1933, Psyche, 40, pt. 4, p. 141. Amblyomma infestum taivanicum Schulze, 1935, Zool. Anz., 112, pts. 9-10, pp. 234, 236-237, fig. 2 ( J1, $, Kosempo, Formosa, off dog; holotype in D. Ent. Inst. Berlin- Dahlem). Amblyomma infestum borneense Schulze, 1936, Zeitschr. f. Parasitenk., 8, pt. 6, pp. 623-624, fig. 4 (3 3 5, Borneo, off Sus verrucosus ; holotype in Z. M. Berlin). Haemalastor infestum Oudemans, 1936, Krit. Hist. Overzicht Acarologie, 3, Bd. B, pp. 497-499, fig. 202. Haemalastor infestum var. testudinarium Oudemans, 1936, Op. cit., pp. 544-545, fig. 225. In 1844 C. L. Koch described A. infestum from Bintan Island and A. testudinarium from Java in the same publication as distinct species. Neumann, as first reviser, made A. infestum a synonym of A. testudinarium even though A. infestum had line priority. Schulze in 1932 reversed the names making A. testudinarium a synonym of A. infestum. Neumann gave no reasons for choosing the name A. testudinarium over A. infestum , but since he was the first reviser, the choice of which name to use was up to him. No case can be made for line priority in this situation, and since the name is now firmly established in the literature, no good purpose would be served by reviving the name A. infestum. In 1932 Schulze stated that the male described and figured by Robinson in 1926 on page 254 as A. testudinarium is not that of Koch ; he regarded it as a new species, and he gave it the name Amblyomma fallax. He said that it differed from A. testudinarium in that ventral muscle scutes were present and that the peltae were obliquely or diagonally placed. Schulze, however, has misinter- preted the drawing in regard to the peltae. Robinson in figure 125 has not depicted the salience of the peltae at the postero-internal angle, because it is impossible to see the peltae when the specimen is viewed from the ventral position ; it is necessary to raise the pos- terior end of the specimen and to look down on the posterior margin in order to see the salience of the peltae. In Robinson’s figure the peltae run parallel with the festoons, and this is shown by the solid lines which represent both the lateral margin of the festoons and the peltae. Robinson merely shows the dark brown, ventral scutes of which the internal, lateral border is oblique. The drawing is misleading, however, for the solid lines representing a portion of 116 ENTOMOLOGICA AMERICANA the border of the muscle scutes could easily be mistaken for oblique peltae. Specimens were seen where the ventral muscle scutes are missing entirely and others where they are only slightly developed, but the majority of those examined show well developed ones. For these reasons I believe that the name A. fallax Schulze is unneces- sary and that it should be regarded as a synonym of A. testudi- narium. Schulze attempted to separate A. infestum and A. testudinarium as distinct races in 1932, but I can not agree with this division. The characters used by Schulze to separate these races are color, pattern of the ornamentation and size. All of these characters are known to vary widely and have little, if any, taxonomic value. Numerous specimens were seen from Burma, Assam, Formosa, the Philippine Islands, India, China, Siam, the Federated Malay States and the East Indies, and I am convinced that they are all the same species. I compared specimens from Formosa with the description and figures given by Schulze for A. infestum taivanicum, and I am con- vinced that this is the same as A. testudinarium. I saw no material from Borneo, but the description and figures given by Schulze for A. infestum borneense show that it is also A. testudinarium. Both these subspecies should be considered as synonyms of A. testudi- narium. Male (Fig. 25A-G). Numerous, unengorged specimens from Sumatra, Java, Celebes, Flores from 6.42 to 5.4 mm. long by 6.0 to 5.1 mm. wide, exclusive of capitulum. Body broad-oval, and nar- rowed anteriorly. Scutum yellow-orange and light-brown, some more ornate than others; punctations numerous, large and irregu- larly distributed, with a slight confluency in region immediately anterior to festoons ; elevations present generally free of puncta- tions; cervical grooves short, deep and comma-shaped; eyes large, yellow and flat. Festoons distinct and longer than wide, with inter- val between them wide. Ventral body surface yellow, finely wrinkled and with numerous, small punctations; genital opening opposite coxa II ; anal opening opposite posterior margin of spiracu- lar plate; spiracular plate comma-shaped and obliquely placed; muscle scutes in region of festoons dark brown; peltae parallel to festoons and postero-internal angle slightly salient. Legs long and stout, with a faint, circular marking at distal end of each joint; coxa I with two long, subequal spurs, external spur slightly longer ; coxae II and III each with a broad, single, rounded spur ; coxa IV with a single spur about twice as long as wide ; tarsi short, ending in a distinct terminal and sub-terminal spur; pulvillus about one- 117 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 half as long as claws. Capitulum long ; basis rectangular, with sides slightly convex and posterior margin slightly concave ; palps long, with article two a little more than twice as long as article three; Fig. 25. Amblyomma testudinarium C. L. Koch. A-G, male : A, scutum ; B, coxae I-IV • C, capitulum in dorsal view ; D, spiracle ; E, tarsus IV ; F, tarsus I ; G, ventral muscle scutes, or muscle at- tachments. H-M, female : H, scutum ; I, coxae I-IV ; J, capitulum in dorsal view ; K, spiracle ; L, tarsus IV ; M, tarsus I. hypostome long and slender ; dentition 4 : 4, but internal row on each side beginning about one-fourth distance from anterior tip; eight to nine teeth per file. 118 ENTOMOLOGICA AMERICANA Female (Fig. 25H-M). Fully engorged specimen 25 mm. long by 18 mm. wide by 15 mm. thick. Body of unengorged female nearly circular. Scutum triangular, wider than long, posterior angle narrow, 3.15 mm. long by 3.9 mm. wide, amount and pattern of ornamentation variable; punctations unequal, irregularly dis- tributed, with larger ones in lateral margins near eyes; cervical grooves small and comma-shaped ; eyes yellow, large and conspicu- ous. Abdomen dorsally with numerous, large punctations and short, white hairs ; dorsal foveae small, dark and conspicuous. Abdomen ventrally with numerous, fine wrinkles and many, small puncta- tions ; genital opening opposite coxa II ; anal opening opposite pos- terior border of spiracular plate; spiracular plate large and tri- angular, with rounded corners. Coxae I— III with spurs similar to those of male • coxa IV with spur much shorter than that of male ; tarsi longer than those of male. Capitulum long ; basis rectangular, with sides slightly convex; in some specimens posterior margins slightly concave, in others straight; porose areas large, oval and parallel ; hypostome long and slender ; dentition 4:4; inner row with smaller teeth than other rows. Distribution and Hosts. This species is common and is widely distributed throughout the Oriental Region; it is known from Burma, Indochina, Annam, Ceylon, Borneo, India, Federated Malay States, Formosa, Japan ( ?), Cochin China and the East Indies. In the East Indies it has been recorded from Java (Koch, 1844; Neu- mann, 1899, 1901, 1911 ; Koningsberger, 1900 ; Robinson, 1926 ; Krijgsman and Ponto, 1931, 1932), Sumatra (Neumann, 1901; Robinson, 1926; Krijgsman and Ponto, 1931, 1932; Schulze, 1932), Bintan Island (Koch, 1844; Schulze, 1932), Andalos, Pandang and Boo Island (Warburton, 1926), Bali (Robinson, 1926; Krijgsman and Ponto, 1931, 1932), Lombak and, Soemba (Krijgsman and Ponto, 1931, 1932). Numerous specimens were seen from the following localities : Sumatra (Lhokseumawe, off buffalo and goat; Medan, off buffalo; Padangsidimpoean, off buffalo and goat ; Lahat, off buffalo and cow ; all in Kraneveld collection), Java (Blitar, off buffalo; Dj ember, nymph, off pig, in Kraneveld collection ; two females, off Sus verrucosus, in Museum of Comparative Zoology; West Java, two males, four females, off Bos sondaicus sondaicus collected by F. C. Kraneveld, in Museum of Comparative Zoology), Celebes (two fe- males, without host; four males, off Sus barbatus, in Museum, of Comparative Zoology), Flores (Roeteng, one male and one nymph, off cow, in Kraneveld collection). 119 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Biology. All stages in the life cycle are known. The only host record for the larval stage is that of two larvae, taken off Tapaia belangeri versurae, in the Rocky Mountain Laboratory collection. These ticks occur on a wide variety of wild as well as domestic animals, and they must be regarded with suspicion as possible dis- ease vectors. Krijgsman and Ponto stated in 1932 that this species transmits piroplasmosis and anaplasmosis, and Sharif in 1938 con- sidered this species as a possible disease vector. Genus Aponomma Neumann, 1899 Key to Species of Aponomma Males 1. Hypostome with 4 : 4 dentition. Coxa I with single, blunt spur, much broader than long. Scutum nearly oval and ornate. Cornua absent. Punctations unequal and restricted more to margins and ornamented regions. Median area less punctate A. trimaculatum Hypostome with 3 : 3 dentition 2 2. Scutum inornate and brown in color. Punctations unequal and uniformly distributed. Coxa I with two distinct and sepa- rated spurs ; external about twice as long as internal. Spi- racles about twice as long as wide A. barbouri Scutum ornate 3 3. Coxa I with two subequal spurs ; internal slightly broader than external. Scutum decidedly broader than long. Puncta- tions numerous, unequal and more or less uniformly dis- tributed. Spiracles very long and narrow. Cornua very short and blunt A. komodoense Coxa I with two distinct, short, separated spurs ; external slightly longer than internal. Scutum nearly as broad as long. Punctations few, unequal and sparsely distributed. Spiracles large and almost as wide as long. Cornua small and blunt ^ A. lucasi Females5 1. Hypostome with 4 : 4 dentition. Coxa I with single, broad spur. Punctations numerous, small to medium and uniformly distributed. Scutum cordiform and ornate. A. trimaculatum Hypostome with 3 : 3 dentition. Coxa I with two distinct and separated spurs 2 5 Female of A. barbouri unknown. 120 ENTOMOLOGICA AMERICANA 2. External spur of coxa I slightly longer than internal, but nar- rower. Spiracles about as long as broad. Scutum slightly wider than long. Punctations unequal with smaller ones more numerous. Porose areas medium, fairly deep and slightly divergent anteriorly. Basis capituli sub-triangular. A. lucasi Internal spur of coxa I broader and longer than external. Spira- cles long and narrow. Scutum slightly wider than long. Punctations unequal with larger ones more numerous, deeper and more or less regularly distributed. Porose areas medium, circular and deep. Basis capituli triangular. A. komodoense Aponomma barbouri, new species. This is the only inornate. Aponomma known from Indonesia ; in addition, the arrangement and number of punctations, the shape of the spiracular plate and the spurs On coxa I are distinctive. Male (Pig. 26A-E). Three specimens respectively 3.0, 2.55 and 2.25 mm. long by 2.85, 2.55 and 2.55 mm. wide, exclusive of capitu- lum. Scutum light brown, inornate, about as broad as long and slightly narrowed at region opposite cervical grooves; punctations numerous, small to large and uniformly distributed ; cervical grooves medium, deep and inverted comma-shaped ; no depressions, elevations or marginal grooves present ; few, small, insignificant hairs in region of festoons ; festoons short and inconspicuous. Ven- Pig. 26. Aponomma barbouri, new species, male : A, scutum ; B, coxae I-IY ; C, capitulum in dorsal view; D, spiracle; E, tarsus IV. 121 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 tral body surface yellow with long, white hairs; spiracular plate long and narrow, with antero-dorsal edge heavily chitinised ; genital opening opposite coxa II ; anal opening opposite mid-portion of spiracular plate. Legs medium; coxa I with two spurs, external nearly twice as long as internal; coxae II-IV each with a distinct, median, triangular spur ; tarsus I short, tapering abruptly and end- ing in a distinct, ventral, terminal spur ; tarsus IV short, tapering gradually and with a small, sub-terminal spur and a large, terminal spur ; claws about twice as long as pulvillus. Capitulum 0.84 mm. long; basis with lateral edges slightly convex and posterior border slightly concave ; cornua distinct ; palps short and rugose ; article two less than twice as long as article three ; hypostome long and slender ; dentition 3 : 3, with approximately six teeth per file ; toothed area extends posteriorly less than one-half length of hypo- stome. The specimens are badly broken and in a very fragile condition. The holotype and paratypes, in the Museum of Comparative Zool- ogy, were taken on Python reticulatus at Buitenzorg, Java, by the late Dr. Thomas Barbour, in December, 1906. Aponomma komodoense Oudemans. Aponomma komodoense A. C. Oudemans, 1929, Zool. Meded. Leyden, 11, pt. 4, (1928), pp. 227-231, PL 11, figs. 1-6, PI. 12, figs. 7-14 (35 J1, off Varanus komodoensis ; cotypes at Zool. M. Amsterdam). Whittick, 1939, Parasitology, 31, pt. 4, pp. 434-436, figs. 1-2 (23 J1, 9 J, 2 n, Zool. Gardens, London, off Varanus komodoensis) . Aponomma draconis Warburton, 1933, Parasitology, 24, pt. 4, pp. 564-565, fig. 7 (3 6 §, Zool. Gardens, London, off Varanus komodoensis from Komodo Island; types in Brit. Mus.). The cover of part 4 of volume 11 of the “Zoologische Mededeelin- gen” is dated December 31, 1928, but according to a MS note on a reprint sent to Dr. J. C. Bequaert by Dr. A. C. Oudemans, the paper actually appeared February, 1929. The description then dates from 1929. Whittick in 1939 pointed out that the figure of coxa I given by Oudemans is inaccurate, because there are two blunt spurs present on this coxa. Dr. J. C. Bequaert in unpublished note states that he saw a male cotype from the Zoological Museum, Amsterdam, and that it does have a bifid spur. Several specimens seen from the American Museum of Natural History confirm the findings of Whittick and Bequaert. 122 ENTOMOLOGICA AMERICANA Male (Fig. 27A-E). Scutum broader than long and broadest posteriorly in region of spiracles ; four specimens, exclusive of the capitulum, from 2.9 to 2.7 mm. long by 3.15 mm. wide; yellow- green to reddish-brown with variable ornamentation ; in general, a small, irregular marking on each side of cervical grooves, a small marking in center of scutum, a variable, lateral stripe on each side of scutum, and in two specimens very pale, insignificant markings on festoons. Punctations more numerous, deeper and restricted to posterior half; smaller punctations less conspicuous, shallow and more uniformly distributed ; cervical grooves teardrop-shaped, short, deep and wide ; no lateral grooves ; festoons slightly longer than wide, punctate on anterior portion, posterior half lighter in color. Ventral body surface yellow, with numerous, short, white hairs; genital opening opposite coxa II ; anal opening opposite posterior border of spiracles; spiracles unusually long and narrow; genital groove widely divergent; anal groove distinct. Legs medium and thick ; some specimens with a pale marking on dorsal, distal end of each femur ; tarsus I with two dorsal processes ; tarsi II-IV with only one dorsal process; claws very long and slender, about three times as long as pulvillus; coxa I with two. spurs, internal slightly broader and longer than external ; coxae II— III with a single, small spur; coxa IV with a single spur, but slightly larger than that of coxae II— III. Capitulum 1.68 mm. long; basis triangular; cornua very short and blunt ; palps long, slender and hirsute ; article two nearly twice as long as article three ; hypostome long and spatulate ; dentition 3 : 3, with approximately nine small teeth per file. Female (Fig. 27F-K). Single, partially engorged female nearly circular in shape, 3.3 mm. long by 3.3 mm. wide, exclusive of capi- tulum. Scutum light brown, cordate, wider than long, posterior angle rounded, 1.8 mm. long by 2.4 mm. wide ; an irregular, orna- mented spot beside each cervical groove, and a smaller spot in center of scutum ; punctations and cervical grooves similar to those of male. Abdomen rugose and with large punctations ; dorsal foveae small, circular and distinct ; no marginal groove ; festoons similar to those of male. Abdomen ventrally punctate, with numerous, short, white hairs ; genital opening opposite coxae II-III ; anal opening opposite spiracles; spiracles slightly broader than those of male; genital grooves widely divergent; anal groove distinct. Legs and coxae similar to those of male. Capitulum 1.05 mm. long; porose areas medium, circular, deep, with interval between them narrow ; cornua short and blunt ; hypostome spatulate ; 3 : 3 dentition, with approxi- mately eight, large teeth per file. 123 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Distribution and Hosts. This species is restricted to Komodo Island and apparently is found only on reptiles. Pour males and one female from Komodo Island were seen in the collection of the American Museum of Natural History. They were found in associ- ation with specimens of Amblyomma helvolum and Amblyomma robinsoni, without indication of host. According to J. C. Bequaert’s Pig. 27. Aponomma komodoense Oudemans. A-E, male : A, scutum ; B, coxae I-IV • C, capitulum in dorsal view ; D, spiracle ; E, tarsus IV. F-K, female : F, scutum ; G, coxae I-IV ; H, tarsus I ; I, capitulum in dorsal view ; J, spiracle • K, tarsus IV. unpublished notes, one male and four females taken off Varanus komodoensis , on Komodo Island, are found in the Chicago Natural History Museum. Aponomma lucasi Warburton. Aponomma gervaisi yar. lucasi Warburton, 1910, Parasitology, 3, pt. 4, pp. 396, 404, 405, 406-407, fig. 10 (,'J1, J, off various 124 ENTOMOLOGICA AMERICANA reptiles in Zoological Gardens, Calcutta ; type in Ind. Mns. Calcutta). Sharif, 1928, Records Indian Mns., 30, pt. 3, pp. 337-340, fig. 49 (.J1, 2, n, 1, Siam, Lower Burma, Nar- condam Island, Nicobar Islands, India and Ceylon, off many hosts) ; 1932, Op. cit., 32, pt. 2, p. 112. Aponomma gervaisi lucasi Schulze, 1932, Zeitschr. f. Parasi- tenk., 4, pt. 3, p. 471. Aponomma lucasi Schulze, 1933, Arch. f. Hydrobiol., Suppl. Bd. 12, pp. 498-500 (12 J', 3 2, Ranau, Sumatra, off Varanus salvator) ; 1934, Zeitschr. f. Parasitenk., 7, pt. 2, p. 167 (2 2; Tjiboeni, Java, off a coluberide) ; 1935, Zool. Anz., 112, pts. 11-12, p. 330; 1936, Zeitschr. f. Parasitenk., 8, pt. 6, pp. 631-633, 634 (India to Indonesia) ; 1939, Op. ■cit., 10, pt. 6, p. 724 (2 4 2, Maymyo, Burma, off Va- ranus) ; 1941, Zeitschr. Morph. Oekol. Tiere, 37, pt. 3, p. 518, fig. 30 (morphological). Aponomma gervaisi Krijgsman and Ponto, 1931, [nec Lucas, 1847] Zeitschr. f. Parasitenk., 4, pt. 1, pp. 141, 144, map 5; 1932, [nec Lucas, 1847] Veeartsenijk. Meded., No. 79, p. 25, figs. 39-40, map 5 (J\ 2? Java, off Varanus). Tou- manoff, 1944, [in part: nec Lucas, 1847] Les Tiques de l’Indochine, pp. 122-124, PI. 85. f Aponomma gervaisi Neumann, 1911 [in part: nec Lucas, 1847] Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 93-94 [only the Java record]. Yitzthum, 1931, Resultats Sci- entif. Voyage Indes Orient. Neerland. Prince Leopold, 3, pt. 5, p. 30 (Ternate, off Varanus indicus and Lophura amboinensis ; Babi, off Varanus sp.). Warburton gave good characters for separating lucasi and gervaisii, but subsequent workers have confused them. Though similar in facies, these two ticks differ enough to be regarded as distinct species. Schulze in 1932 first pointed out that these sup- posed varieties might be distinct species ; and in a series of papers from 1933 to 1939 he regarded them as such. In the males of gervaisii the lateral stripe on each side is elon- gate and enlarged anteriorly; this enlargement may be separated from the stripe to form a distinct spot which is on a level with the cervical grooves and projects more or less into the scapular region; in some cases the lateral stripe is reduced or obsolete, but the de- tached enlargement still persists in the scapular region. In the males of lucasi the lateral stripe does not reach to the level of the cervical grooves, and the ornamentation is completely absent in the 125 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 scapular region. The punctations of gervaisii are more equal in size and more uniformly distributed over the scutum, while those in lucasi are markedly unequal and less uniformly distributed. Coxa I of gervaisii may have a very short, blunt spur or else may have a second, smaller, inconspicuous spur internal to it ; they are so near each other that they appear as a single, blunt, spur, but on careful examination there can be seen, in most cases, two distinct points, with the external slightly longer. In lucasi the coxal spurs are definitely separated, and they appear as two distinct spurs with the external slightly longer. The spurs of coxae II-IV in gervaisii are more pointed than in lucasi. The cornua of lucasi are very short and blunt, whereas gervaisii lacks cornua. The females also differ in several important respects. The coxal spurs of gervaisii are similar to those of the male ; in some cases the internal spur is so reduced that it is inconspicuous or obsolete, but in no cases were they seen to be distinctly separated. The coxal spurs of lucasi are also similar to those of the male ; the spurs are distinct and decidedly separated. The spiracular plate of lucasi is wider and larger than that of gervaisii. The scutum of gervaisii has larger metallic spots and larger punctations than that of lucasi. The geographical distribution of these two species also gives evi- dence to their distinctness. Aponomma gervaisii probably does not occur in the East Indies. Schulze stated in 1935 that lucasi is more widely distributed than gervaisii ; he found lucasi to range from India to Indonesia, whereas he found gervaisii only in India and Ceylon. Material I have seen from numerous localities in the Ori- ental Region substantiates these findings. The published records of gervaisii from Indonesia are all ques- tionable ; when a figure is given it is always seen to be lucasi and not gervaisii. Neumann confused several forms under this name in 1911, so his record from Java must be regarded as doubtful. Fig- ures given by Krijgsman and Ponto in 1932 for gervaisii are defi- nitely lucasi. Vitzthum who recorded it from the Netherlands East Indies in 1931 as gervaisii gave no figure or description, so it is impossible to know what he had ; from the known distribution, how- ever, it can only be lucasi. Toumanoff ’s figure of the female is mis- leading, for he shows the spur of coxa I to be very long ; either he has a new species or his drawing is inaccurate. Toumanoff claims to have found all gradations of the spurs from gervaisii to lucasi ; he further claims to have found males of one variety copulating with females of the other variety and vice versa. It is doubtful that Tou- manoff had both species in his possession; he probably had lucasi, 126 ENTOMOLOGICA AMERICANA / where some variation occurs in the form of coxa I, and he is probably referring to this variation within the species under the mistaken im- pression that he has both species. Male (Fig. 28A-E). This species varies from 2.9 to 2.18 mm. long by 2.85 to 2.14 mm. wide, exclusive of capitulum. Scutum nearly as broad as long, posterior margin nearly straight, reddish- brown and with five metallic-green spots of variable thickness, shape and intensity; two lateral elongate spots not reaching cervical grooves; a single spot in median area; two posterior spots sub- Fig. 28. Aponomma lucasi Warburton. A-E, male : A, scu- tum; B, coxae I-IV ; C, capitulum in dorsal view; D, spiracle; E, tarsus IV. F-K, female : F, scutum ; G, coxae I-IV ; H, tarsus IV ; I, capitulum in dorsal view ; J, spiracle ; K, tarsus I. triangular and close together; lateral margin completely outlined by a continuous, light brown stripe ; punctations small to large, with some specimens more punctate than others ; cervical grooves short, deep and convex externally ; no lateral grooves ; festoons set off by 127 ENTOMOLOGICA AMERICANA / Vol. XXX, Nos. 1-4 a light brown stripe, broader than long and without punctations. Abdomen ventrally pale yellow, with short, thick, white hairs ; large, genital opening opposite coxa II ; anal opening opposite spiracnlar plate ; spiracular plate large and sub-rectangular ; genital groove widely divergent; anal groove deep and distinct. Legs yellow- brown and fairly thick; tarsus I short, with three dorsal and one ventral prominence, tapering abruptly without ending in a spur; tarsi II— IY humped in middle, tapering gradually and terminating in a strong, terminal and sub-terminal spur; pulvillus small, one- third length of claws ; coxa I with two short, distinct and separated spurs; external spur slightly longer than internal; coxae II-IV with a single, blunt spur about as wide as long. Capitulum 0.9 mm. long; basis sub-triangular; cornua small and blunt; palpal article two about twice as long as article three ; long, white hairs on palps ; hypostome spatulate ; dentition 3 : 3, with approximately nine teeth per file. Female (Fig. 28F-K). Largest of six engorged specimens 5.4 mm. long by 4.35 mm. wide, exclusive of capitulum. Scutum from 1.8 to 1.5 mm. long by 2.11 to 1.95 mm. wide, cordiform, slightly wider than long, posterior angle rounded ; reddish-brown and with three greenish, metallic spots of variable size, shape and intensity; punctations unequal, with smaller ones more numerous; cervical grooves similar to those of male. Abdomen lighter than scutum and with a few, short, white hairs; postero-median groove and two postero-lateral grooves present ; festoons about as wide as long and punctate ; genital opening opposite coxa II; anal opening opposite spiracles ; spiracles broader than those of male, but similar in shape. Legs and coxae similar to those of male. Capitulum 1.12 mm. long ; basis sub-triangular; porose areas medium, fairly deep and close together; palps similar to those of male; cornua short and blunt; hypostome spatulate ; 3:3 dentition, with approximately eight to nine teeth per file. Distribution and Hosts. This species is distributed from India to the East Indies. It is definitely known from India, Siam, Burma, Narcondam Island, Nicobar Islands, Ceylon and the East Indies. It was recorded from Java (Neumann, 1911 ; Krijgsman and Ponto, 1931, 1932; Schulze, 1934), Sumatra (Schulze, 1933) and Ternate and Babi (Vitzthum, 1931). Specimens were seen from the following localities in the collec- tions studied: Java (Buitenzorg, eight males, four females collected by Palmer and Bryant in March, 1909 ; six males, one female, two nymphs off Varanus collected by Palmer and Bryant on March 17, 128 ENTOMOLOGICA AMERICANA 1909 ; thirteen males, one female off V ar anus collected by Palmer and Bryant on March 28, 1909 ; four males, four females off V ar anus salvator collected by T. Barbour; all in Museum of Comparative Zoology) ; Malay States (Singapore, four males, two females off Naia tripudians in New York Zoological Society, collected by T. Barbour, in Museum of Comparative Zoology) ; India (two males, one female off king cobra, collected by G. Wiley, in Museum of Comparative Zoology) ; Burma (Myitkyina, three males, three fe- males off Varanus monitor , collected by G. E. Davis, May 28, 1945, in Rocky Mountain Laboratory ; Maymyo, three males, two nymphs off Varanus salvator , in Museum of Comparative Zoology). Aponomma trimaculatum (Lucas). Ixodes trimacidatus Lucas, 1878, Ann. Soc. Ent. France, (5) 8, Bull., pp. lxxvii-lxxviii ('§, New Guinea, off Varanus chloro stigma; type in Paris Museum). Aponomma trimaculatum Neumann, 1899, Mem. Soc. Zool. France, 12, pp. 187-188, fig. 41 (type material; 5 J, New Guinea, off Varanus leuco stigma; 1 5, Padang, Sumatra, off Varanus salvator). Salmon and Stiles, 1902, 17th Ann. Rept. Bur. Anim. Industry, U. S. Dept. Agric., (1901), p. 470. Neumann, 1911 Das Tierreich, Lief. 26, Acarina, Ixodidae, pp. 93, 94 [in part: only New Guinea distribu- tion]. Warburton, 1926, Treubia, 8, pt. 3-4, p. 279 (<£, Sinabang, Simeuloee, off Varanus salvator). Krijgsman and Ponto, 1932, Veeartsenijk. Meded., No. 79, p. 25, map 5. Amblyomma ( Aponomma ) primaculatum [sic] Warburton, 1925, Spolia Zeylanica, 13, pt. 2, p. 256 (Colombo, Ceylon, off Varanus salvator). AmMyomma ( Aponomma ) trimaculatum Fielding, 1926, Comm. Austral. Service Publ. (Trop. Div.) No. 9, p. 92, fig. 36 (after Neumann). Aponomma undatum Schulze [nec Fabricius, 1775], 1932, Zeitschr. Morph. Oekol. Tiere, 25, pt. 2-3, pp. 515, 516, 527, figs. 6c, 20, 21 (morphological) ; 1933, Arch. f. Hydrobiol., Suppl.-Bd. 12, pp. 500-501, figs. 11-12, (1^), Ranau, Sumatra, off Varanus salvator • 5, n, Gasmata, New Britain, off Varanus) ; 1941, Zeitschr. Morph. Oekol. Tiere, 37, pt. 3, pp. 518, 524 (morphological). Aponomma trabeatum Schulze, 1933, Zool. Anz., 104, pts. 11- 12, pp. 322-323, fig. 7 (3 4 'J, Wasior, New Guinea, off Varanus ; holotype $ in Z. M. Berlin). 129 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 Aponomma fimbriatum trabeatum Schulze, 1935, Zool. Anz., 112, pts. 11-12, p. 330 (J*, Wangil Wammer, Aroe Island, off Varanus ; 1 2 J, Langemak Bay, New Guinea, off Varanus). ? Aponomma trimaculatum Neumann, 1901 Mem. Soc. Zool. France, 14, p. 291 (1 J, New South Wales, off Bos taurus). Roberts, 1934, Queensland Agric. Journ., 42, pt. 2, p. 123 (Townsville, Australia, off horse). ? Aponomma gervaisi Oudemans [nec Lucas, 1847], 1927, Zool. . Meded. Leyden, 10, pt. 4, pp. 222-223 (<£, , Amboina, off Varanus indicus). Records of trimaculatum from Australia (Neumann, 1901 and Roberts, 1934) and from Ceylon (Warburton, 1925) are doubtful because the distribution and hosts are not normal; neither figures nor descriptions were given for these records. It is possible that it was mistaken for A. decorosum (Koch), an Australian species superficially resembling trimaculatum. Schulze in 1932 and 1933 considered A. trimaculatum (Lucas) as a synonym of Acarus undatus Fabricius, but the evidence for this is inconclusive. Fabricius described undatus in 1775 from specimens collected by Banks on Captain Cook’s voyage to Australia, and he gave the locality as Nova Hollandia. Schulze may have been mislead because of the supposed occurrence of trimaculatum in Australia. Oudemans in 1929 stated that the description of undatus was based on the male of Ixodes decorosus Koch, placed in the genus Aponomma by Neumann, and I am inclined to agree with him. Schulze described trabeatum in 1933 from three males and four females collected in North New Guinea. I regard it as a synonym of -trimaculatum, for these two species resemble each other in all essen- tial details. At first I thought that these two forms could be sepa- rated on the basis of the observed differences In the ornamentation. However, I saw somewhat later a lot of six males and one female from New Britain in which the males are intermediate in regard to the ornamentation, and one specimen is unique in showing on the left side of the body the pattern typical of trimaculatum and On the right side that typical of trabeatum (see Fig. 29B-E). The varia- tion in the ornamentation is so great that it has no value in separat- ing these forms. An occasional specimen of each form is often more punctate, but this is also an individual variation, for it can not be correlated with any other character. Since material is not available from the East Indies, specimens 130 / ENTOMOLOGICA AMERICANA in the Museum of Comparative Zoology from Munda, New Georgia, are used for the descriptions. Fig. 29. Aponomma trimaculatum (Lucas). A-I, male: A, scutum ; B, C, D, E, variations of pattern of scutum ; F, coxae I— IV ; G, capitulum in dorsal view; H, spiracle; I, tarsus IV. J-O, fe- male : J, scutum ; K, coxae I-IV ; L, tarsus IV ; M, capitulum in dorsal view ; N, spiracle ; 0, tarsus I. Male (Fig. 29 A-I). Three specimens, exclusive of capitulum, respectively 2.16, 2.1 and 2.04 mm. long by 2.1, 1.98 and 1.95 mm. 131 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 wide. Scutum reddish-brown and ornate, with five metallic-green spots of variable size and shape; median spot nearly circular and slightly behind cervical grooves; two elongate, lateral spots begin at level of median spot and extend a short distance posteriorly to level of fourth pair of legs; posterior spots on each side as two ir- regular lobes joined by a narrow band; punctations unequal and more restricted to margins and to ornamented regions, making median region less punctate ; cervical grooves short, deep and slightly convex externally; festoons broader than long, being set off by a light, marginal band. Ventral body surface wrinkled, punctate, lighter in color and with a few, very short, white hairs ; spiracular plates nearly rectangular; genital opening relatively large and opposite coxae II and III ; anal opening opposite spiracu- lar plate ; genital groove widely divergent ; anal groove distinct. Legs light brown and relatively short; tarsus I slightly swollen, with two prominent, dorsal protuberances, tapering abruptly with- out terminating in a spur; tarsi II-IV with a single, very large, dorsal protuberance slightly beyond mid-length ; tapering abruptly from this point and terminating in a very short, inconspicuous spur ; very small pulvillus, about one-third as long as claws ; coxa I with a single, blunt spur, broader than long ; coxae II-IV with a single spur. Capitulum 0.84 mm. long; palps long and narrow; article two slightly over twice as long as article three ; basis sub-rectangular and bordered in black; lateral margins slightly convex and pos- terior margin slightly concave ; hypostome dentition 4 : 4, with ap- proximately twelve teeth per file. Female (Fig. 29J-0). Scutum of two specimens 1.65 and 1.71 mm. long by 1.86 mm. wide, cordiform, slightly broader than long, lateral angles not well marked, posterior angle not very broad ; red- dish-brown and ornate, with three metallic-green spots ; lateral spots extending into scapular angles, and median spot behind cervical grooves; punctations numerous, fine, unequal and close together; cervical grooves short and deep anteriorly, prolonged posteriorly as a short, shallow groove concave externally. Abdomen lighter in color than scutum, punctate, and with a few, very short hairs; no marginal groove ; festoons about as wide as long, with grooves sepa- rating them narrow and deep. Ventral body surface finely wrin- kled, punctate and with very short, white hairs in posterior half ; genital opening opposite coxae II — III ; anal opening opposite pos- terior border of spiracles ; spiracles broad and tapering toward top ; genital groove distinct. Legs long and light brown ; tarsus I long, with three dorsal protuberances and tapering abruptly from last 132 ENTOMOLOGICA AMERICANA protuberance ; tarsi II-IV long, with a distinct, dorsal protuberance and tapering from this point ; tarsi without spurs ; pulvillus about one-third as long as claws ; coxa I with a single spur broader than long; coxae II-IV with a single, blunt, short spur, slightly longer than that of coxa I. Capitulum normal, 0.98 mm. long; basis broad and outlined, in black; lateral margins rounded, posterior margin slightly concave ; palps long and narrow ; article two about twice as long as article three ; hypostome dentition 4 : 4, with ap- proximately twelve teeth per file. Distribution and Hosts. This species is recorded from the East Indies, New Guinea, New Britain, Australia (?), and Ceylon (?). In the East Indies it has been recorded from Sumatra (Neumann, 1899; Schulze, 1933), Simeuloee . (Warburton, 1926), Aroe Island (Schulze, 1935) and Amboina (Oudemans, 1927). Specimens were seen from the following localities in the Orien- tal Regions: New Guinea (Toem, five males off Varanus indicus, in Museum of Comparative Zoology; Liki Island, two males off Varanus indicus, in Museum of Comparative Zoology; one male, off Varanus collected by W. E. Stickel, August 18, 1944, in Rocky Mountain Laboratory; New Guinea, two males off Varanus, in American Museum of Natural History), New Britain (six males off Varanus collected January, 1933 by W. F. Coultas, in American Museum of Natural History), New Georgia (Munda, one male, one female off large spotted lizard collected by W. G. Downs, Septem- ber, 1943, in Rocky Mountain Laboratory ; and same locality, three males, two females, one nymph off Corucia zebrata, collected by R. W. Brubaker, December 8, 1944, in Museum of Comparative Zoology). Species of Doubtful Occurrence in the East Indies Amblyomma petersi Karsch, 1878. Neumann determined a female from Java as this species which is strictly African in dis- tribution, and in 1908 he stated that it was A. testudina- rium and not A. petersi. Amblyomma serpentinum Schulze, 1936. He described and figured a male and female taken in the Hamburg Zoo off Python molurus bivittatus which occurs in the Oriental Region. The tick resembles A. marmoreum Koch, 1844, and it is probably this species. More than likely it is an African tick gone astray in the zoo. Haemaphysalis leporis-palustris (Packard, 1869). Neumann in 1897 determined a nymph from Timor off Paradoxurus. 133 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 This determination is erroneous, for this species only oc- curs in the New World. Haemaphysalis parmata Neumann, 1905. Galli-Valerio in 1909 stated that Dr. Narbel found this species on a monkey in Sumatra. This determination is erroneous, for parmata is not known to occur outside Africa. Hyalomma aegyptium (Linnaeus, 1758) . This species was reported from Java by^de Blieck in 1913 and 1914; it was probably introduced with domestic livestock from India. There are no recent records, and it may have died out. Rhipicephalus bursa Canestrini and Fanzago, 1877-1878. This is an African species, and Neumann in 1897 recorded one female from Timor taken off Cervus unicolor equinus. This is an error in determination. 134 Index Valid genera and species in Roman; synonyms in Italics ; new names of species and main page references in bold face ; # names of animals, other than ticks. #abramus, Vesperugo, 103 Acarus, 130 aegyptium, Hyalomma, 134 aeratipes, Amblyomma cyprium, 91, 94, 95 Amblyomma, 2, 3, 6, 7, 8, 9, 11, 12, 13, 14, 15, 77, 78, 79, 80, 81, 82, 83, 84, 85, 86, 87, 89, 90, 91, 92, 94, 95, 96, 97, 98, 99, 100, 102, 103, 104, 105, 106, 107, 109, 110, 111, 113, 114, 115, 116, 117, 118, 124, 129, 133 *amboinensis, Lophnra, 125 ^Anaplasma, 77 #Anatidae, 10 annulatus, Boophilns, 72, 73 Boophilus, 72 Rhipicephalus, 71 *Anser, 10 #anser, Anser, 10 Aponomma, 7, 8, 9, 15, 102, 103, 120, 121, 122, 124, 125, 126, 127, 129, 130, 131 aquilae, Ixodes, 98 - #Arctonyx, 11, 35, 59, 65 ardena, Harpactes, 95 *aries, Ovis, 14 *aristotelis, Cervus, 27, 44 *Artiodactyla, 13 *Ateles, 90 auratus, Dermacentor, 6, 7, 8, 13, 51, 52, 53, 56, 58, 59. Fig. 10 A-M; fig. 11 A-M; fig. 12 A-G #aureus, Canis, 27 auriscutellatus, Ixodes, 115 *aurita, Manis, 102 australis, Boophilus, 71 Haemaphysalis leachi var., 39, 42 Rhipicephalus, 71 Rhipicephalus annulatus var., 72 *Aves, 10 ^Babesia, 77 *Babirnssa, 13, 80 *babirussa, Bus, 80 babirnssae, Amblyomma, 6, 7, 8, 9, 13, 14, 79, 80, 82, 91. Fig. 17 A-L *babyrussa, Babirussa, 13, 80 badium, Amblyomma, 103 *barbatus, Sns, 13, 114, 119 barbouri, Aponomma, 7, 8, 9, 120, 121, 122. Fig. 26 A-E bartelsi, Haemaphysalis, 40, 42 *Rattus, 12, 18 *belangeri, Tnpaia, 21, 34, 120 135 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 *bengalensis, Hystrix , 35 *Felis, 11, 39 *bicolor, Ratufa, 12, 39 #Ratufa bicolor, 12, 39 #bicornis, Rhinoceros, 85 *bigemina, Babesia, 77 birmaniae, Haemaphysalis, 27, 36 bispinosa, Haemaphysalis, 6, 7, 8, 9, 11, 13, 14, 24, 25, 26, 27, 28, 29. Fig. 4 - A-L Haemaphysalis, 36 #bivittatus, Python molurus, 133 *Varanus, 97 #Boidae, 9 *Boiga, 9, 96, 97, 98 Boophilus, 2, 6, 7, 8, 9, 11, 13,-14, 15, 16, 65, 66, 71, 72, 73, 74, 77 borneense, Amblyomma infes- tum, 116, 117 *Bos, 14, 27, 30, 31, 34, 47, 50, 61, 62, 66, 114, 119, 130 *Bovidae, 14 * b r e vi c au d at us , Rattns rattns, 12, 18 #bubalis, Bos, 14, 30, 31, 50, 62, 66, 114 *Bos bubalis, 14, 50, 62, 66, 114 *bubalus, Buffelus, 66 *Buffelus, 62, 66, 114 *bukit, Battus, 18, 21 #Rattus fulvescens, 12, 21 *buku, Nycticebus, 40 *bungarus, Naja, 96 bursa, Rhipicephalns, 134 *caballus, Equus, 13, 27 caelaturum, Amblyomma, 107, 108 Amblyomma caelaturum, 107, 108 *Callosciurus, 12, 17, 18, 21, 34, 59 *Canidae, 11 *Canis, 11, 27, 62 # Capra, 14 # Carnivora, 11 #celebensis, Sus, 13, 80, 90 *Cervidae, 13 #Cervrdus, 30, 114 #Cervus, 13, 27, 34, 44, 59, 65, 71, 72, 134 *chaus, Felis, 40 *Chiroptera, 10 #chlorostigma, Varanus, 129 ^collaris, Arctonyx, 11, 35, 59, 65 # Coluber, 10, 96 #Colubridae, 9 compactum, Amblyomma, 115 compressum, Amblyomma, 105 #concolor, Rattus, 12, 17, 18 cordiferum, Amblyomma, 7, 8, 9, 77, 79, 83, 84. Fig. 18 A-F cornigera, Haemaphysalis, 6, 7, 8, 10, 14, 24, 26, 30, 31, 33. Fig. 5 A-L Haemaphysalis, 31 #Corucia, 133 136 ENTOMOLOGICA AMERICANA #coucang, Nycticebus, 11, 40 crenatum, Amblyomma, 7, 8, 78, 79, 85, 86, 87, 89. Fig. 19 A-M *cristatus, Sus, 13, 51, 59, 62, 66, 114 #Sus cristatus, 13 ^Crocidura, 10, 18, 21 cuneatum, Amblyomma, 105 Amblyomma javanense, 105 cyprium, Amblyomma, 7, 8, 9, 13, 14, 79, 80, 89, 90, 91, 92, 94, 95. Fig. 20 A-L Amblyomma, 80, 89, 94, 107, 108 Amblyomma cyprium, 8, 9, 14, 79, 80, 89, 91, 92. Fig. 20 A-L dammermani, Amblyomma, 90, 91 decoloratus, Boophilus, 72, 73 decoratum, Amblyomma, 97, 98, 99 decorosum, Aponomma, 130 decorosus, Ixodes, 130 #dendropbila, Boiga, 9, 96, 97, 98 *dendrophilum, Triglyphedon, 97 *dendrophilus, Dipsadomorpkus, 96, 97, 98 Dermacentor, 6, 7, 8, 13, 15, 51, 52, 53, 56, 58, 59, 102 *diardi, Rattus rattus, 12, 17 *Dipsadomorphus, 96, 97, 98 #dolmanni, Manis, 114 *Manis pentadactyla, 102 draconis, Aponomma, 122 *Dremomys, 17 * Edentata, 12 •Elapidae, 10 *Emydidae, 9 #ephippium, Rattus concolor, 12, 17, 18 *EpHmys, 17, 18 *Equidae, 13 * equina, Rusa, 71 #equinus, Cervus unicolor, 13, 71, 134 *Equus, 13, 27 ^Erinaceidae, 10 #erythraeus, Callosciurus, 17, 21 *erythrorhyncba, Urocissa, 21 E schatocephalus, 16 #Euarctos, 35, 51 *Euprepes, 102 expedita, Rhipicephalus haema- physaloides var., 62 expeditus, Rhipicephalus haema- physaloides, 62 fallax, Amblyomma, 116, 117 Boophilus, 72 *familiaris, Canis, 11, 27, 62 ^Felidae, 11 *Felis, 11, 17, 35, 39, 40, 44, 59, 61, 65, 66, 114 *ferruginea, Tupaia, 18 feuerborni, Amblyomma, 98, 99 fimbriatum, Amblyomma, 97, 98, 99 Aponomma, 130 137 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 #flavigula, Martes, 11, 21, 40, 114 * Maries, 21 ^fulvescens, Rattus, 12, 18, 21 fur cosum, Amblyomma, 97, 98, 99 *Gallus, 10, 34, 50, 59, 65 ^gallus, Gallus, 10, 34, 50, 59, 65 # Gallus gallus, 34, 59, 65 *Geocirla, 17 #Geoemyda, 9, 35, 39, 95, 97, 103, 106, 107 geoemydae, Amblyomma, 7, 9, 77, 80, 95 Haemalastor, 95 Ixodes, 95 #Geokichla, 10, 17 geomydae, Amblyomma, 95 gervaisi, Aponomma, 124, 125, 130 gervaisii, Aponomma, 125, 126 ^gigantea, Ratufa, 18, 59 ^Ratufa gigantea, 59 #glis, Tupaia, 10, 18 *gordoni, Callosciurus erythra^ eus, 17 *Seiurus, 17 #grandis, Geoemyda, 97 granulatus, Ixodes, 7, 8, 10, 12, 16, 17, 18, 19. Fig. 2 A-G Haemalastor, 95, 98, 116 Haemaphysalis, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 23, 24, 25, 26, 27, 28, 29, 30, 31, 33, 34, 35, 36, 37, 39, 40, 41, 42, 43, 45, 47, 48, 49, 71, 133, 134 haemaphysaloides, Rhipicepha- lus, 3, 5, 6, 7, 8, 9, 11, 13, 14, 59; 60, 61, 62, 63, 65, 66, 67. Fig. 1 A-F; fig. 13 A-M; fig. 14 A-J Rhipicephalus haemaphysa- loides, 7, 11, 13, 14, 59, 60, 62, 63, 66. Fig. 13 A-M Haemophy salis, 74 *hannah, Ophiophagus, 10, 85, 96 *Harpactes, 95 #Helarctos, 40, 44 *Helictis, 11, 21, 39, 40 helvolum, Amblyomma, 7, 8, 9, 10, 78, 79, 96, 98, 99, 100, 102, 124. Fig. 21 A-L Haemalastor, 98 *henrici, Martes flavigula, 11, 40, 114 #hermaphroditus, Paradoxurus, 11, 21, 39, 40 hircus, Capra, 14 histricis, Haemaphysalis, 36 *hodgsoni, Hystrix, 35 ^hoevenii, Arctonyx collaris, 11, 35 ^Hominidae, 11 #Homo, 11 ^Hyaena, 106 ^hyaena, Hyaena, 106 Hyalomma, 134 Hylobates, 11, 34 138 ENTOMOLOGICA AMERICANA *Hylobatidae, 11 hylobatis, Haemaphysalis, 7, 11, 24, 25, 34 #Hylomys, 10, 18 hypochrysa, Tupaia glis, 10, 18 *hypoleucus, Pomatorrhinus, 34 ^Pomatorrhinus hypoleucus, 34 hystricis, Haemaphysalis, 6, 7, 8, 9, 11, 13, 14, 24, 25, 35, 37. Pig. 6 A-K #Hystrix, 35, 65 *Iguana, 102 ^imitator, Callosciurus, 18 *indica, Viverricula, 59 *indicus, Buffelus, 62, 114 Dermacentor, 102 #Tapirus, 13, 114 #Varanus, 9, 98, 125, 130, 133 Indocentor , 51 infestum, Amblyomma, 115, 116, 117 Amblyomma infestum, 115 Haemalastor, 116 Hnsectivora, 10 *insignis, Lariscus, 12, 18 Ixodes, 7, 8, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 21, 22, 67, 95, 98, 115, 129, 130 *jalorensis, Pattus rattus, 12, 18 javanense, Amblyomma, 7, 8, 9, 12, 78, 79, 102, 103, 104, 105, 106. Fig. 22 A-L Amblyomma compressum, 105 Amblyomma javanense, 105 Aponomma, 103 #javanensis, Felis bengalensis, 11, 39 *Mydaus, 11, 23, 43 *Mydaus javanensis, 11, 23, 43 Rhipicephalus, 102, 103 *javanica, Manis, 12, 102, 103 *Tupaia, 10, 18, 21 *javaniciis, Paradoxurus herma- phroditns, 11, 21, 39 #javanus, Lariscus insignis, 12, 18 *kanchil, Tragulus, 13, 48 *Tragulus, 48 #Tragulus kanchil, 13, 48 kempi, Ixodes, 18 kinneari, Haemaphysalis, 47 komodoense, Aponomma ,7, 8, 9, 120,121,122,124. Fig. 27 A-K *komodoensis, Yaranus, 9, 97, 110, 111, 112, 113, 122, 124 koningsbergeri, Haemaphysalis, 7, 8, 11, 12, 24, 25, 39, 40, 41, 42, 43. Fig. 7 A-L kopsteini, Ixodes, 16 #korros, Ptyas, 9, 102 krijgsmani, Boophilus, 72, 73, 75 139 v ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 ^Lacertilia, 9 *Lariscus, 12, 18 Harvata, Paguma, 36 Harvatus, Paradoxurus, 36 ^Potamochoerus, 114 latum, Aponomma, 103 leachi, Haemaphy satis, 39, 40, 42 leachii, Haemaphysalis, 40, 42, 43 leporis-palustris, Haemaphy- salis, 133 #lepturus, Rattus, 12, 18, 21 Heucodira, Naja naja, 10, 102 Heucostigma, Yaranus, 129 longiscutatus, Boophilus, 72 #Lophura, 125 #Lorisidae, 11 lucasi, Aponomma, 7, 8, 9, 120, 121,124,126,127. Fig. 28 A-K Aponomma, 125 Aponomma gervaisi, 125 Aponomma gervaisi var., 124 #Mabuya, 9, 96, 102 #Macaca, 26 #madurae, Calloscinrns notatus, 12,-18 *Sciurus notatus, 18 #magnirostris, Urocissa erythro- rhyncha, 21 malayanum, Amblyomma, 7, 78, 80, 81, 91, 107, 108, 109. Fig. 23 A-K *malayanus, Helarctos, 40, 44 *Ursus, 40, 44 ^Mammalia, 10 #Manidae, 12 *manipulus, Rattus, 21 *Manis, 12, 102, 103, 106, 114 #marginale, Anaplasma, 77 *marmorata, Felis, 61 marmorenm, Amblyomma, 133 *Martes, 11, 21, 40, 114 *maxi, Rattus, 12, 18 *melanochir, Ateles, 90 #melanurus, Pithecheir, 13, 21 #Pithecheir melanurus, 13, 21 *Pithecheirus, 21 micropla, H aemophy satis, 71 microplus, Boophilus, 2, 6, 7, 8, 9, 11, 13, 14, 65, 66, 71, 72, 73, 74, 77. Fig. 16 A-M #midas, Callosciurus sladeni, 34, 59 #molurus, Python, 102, 103, 106, 133 ^monitor, Yaranus, 129 monospinosa, Haemaphysalis, 48 ^mucosus, Ptyas, 9, 97, 98 *Zamenis, 97 *multifasciata, Mabuya, 9, 96 #Muntiacus, 30, 34, 65, 114 *muntjac, Cervulus, 30 *muntjah, Cervulus, 114 #Muntiacus, 30, 34, 65, 114 *Muridae, 12 *Mus, 17, 40 ^Mustelidae, 11 *Mydaus, 11, 23, 43 *nagarum, Callosciurus ery- thraeus, 21 140 ENTOMOLOGICA AMERICANA "Naia, 129 "Naja, 10, 96, 102 "naja, Naja, 10, 102 "nebulosus, Varanus, 96 neumanni, Haemaphysalis, 27 "Nicoria, 103 niger, Rhipicephalus haemaphy- saloides, 60 *nigrivittatus, Sciurus, 18, 21 *8 'ciurus nigrivittatus, 18, 21 "nigrovittatus, Callosciurus, 12, 18, 21 * Callosciurus nigrovittatus, 12, 18, 21 "notatus, Callosciurus, 12, 18 " Callosciurus notatus, 12, 18 * Sciurus, 18 " Sciurus notatus, 18 "Nycticebus,, 11, 40 "occidentals, Tupaia javanica, 10, 18, 21 "Ophidia, 9 "Ophiophagus, 10, 85, 96 "orientalis, Crocidura, 10, 18, 21 "Helictis, 11, 21, 39, 40 "Ovis, 14 "oxycephalus, Coluber, 10 "Paguma, 36 papuana, Haemaphysalis, 6, 7, 8, 9, 11, 13, 14, 25, 26, 43, 45. Fig. 8 A-L Haemaphysalis 44 "papuensis, Sus, 94 "Paradoxurus, 11, 21, 36, 39, 40, 133 "pardus, Felis, 11, 39, 40, 44, 65 parmata, Haemaphysalis, 134 paulopunctata, Rhipicephalus haemaphysaloides var., 62 paulopunctatus, Rhipicephalus, 60 Rhipicephalus haemaphysa- loides, 62, 66 *pelandoc, Tragulus kanchil, 48 "pentadactyla, Manis, 102, 103 perfectum, Amblyomma caela- turum , 107, 108 "Perissodactyla, 13 "Petaurista, 12, 39, 40 "petaurista, Petaurista, 12, 39 Petaurista petaurista, 12, 39 petersi, Amblyomma, 133 "Phasianidae, 10 "Pholidota, 12 pilaus, Rhipicephalus, 65 Rhipicephalus haemaphysa- loides, 7, 8, 9, 11, 13, 14, 59, 60, 65, 66, 67. Fig. 14 A-J "Pithecheir, 13, 21 "Pithecheirus, 21 politum, Aponomma, 102, 103 "Pomatorrhinus, 34 "Potamochoerus, 114 praematurus, Ixodes, 7, 8, 10, 16, 17 primaculatum, Amblyomma, 129 "Primates, 11 proxima, Haemaphysalis, 31 "Ptyas, 9, 97, 98, 102 "Python, 9, 97, 98, 102, 103, 106, 122, 133 141 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 quadrimaculatum, Amblyomma, 97, 98, 99 quasicyprium, Amblyomma, 90 #quinquestriatus, Callosciurus, 17, 21, 59 # Callosciurus quinquestria- tus, 21, 59 *radiatus, Coluber, 96 Rattus, 12, 17, 18, 21, 34, 40, 59 * raft us, Mus, 17, 40 Rattus, 12, 17, 18, 21, 34, 40, 59 Rattus rattus, 12 Ratufa, 12, 18, 39, 59 renschi, Haemaphy satis, 27, 28 Reptilia, 9 *reticulatus, Python, 9, 97, 98, 122 Rhinoceros, 13, 62, 85, 89, 114, 115 Rhinocerotidae, 13 Rhipicephalus, 2, 3, 5, 6, 7, 8, 9, 11, 13, 14, 15, 59, 60, 61, 62, 63, 65, 66, 67, 68, 70, 71, 72, 102, 103, 134 ^rhodiosternum, Harpactes ar- dena, 95 ricinus, Ixodes, 17 robinsoni, Amblyomma, 7, 8, 9, 78, 79, 110, 111, 113, 124. Fig. 24 A-L Rodentia, 12 ropsteini, E schatocephalus , 16 *roquei, Rattus rattus, 17, 18 rot undiscut at us, Boophilus, 72, 73, 75 ruber, Rhipicephalus, 62 Rhipicephalus haemaphy sa- loides, 60 rubicundus Rhipicephalus, 68 *rufescens, Epimys, 17, 18 Rattus, 17, 18 ^rufigenis, Dremomys, 17 *Sciurus, 17 Rusa, 62, 71 #russa, Cervus unicolor, 13, 72 ^salvator, Yaranus, 9, 96, 97, 103, 106, 125, 129 sanguineus, Ixodes, 68 Rhipicephalus, 2, 6, 7, 8, 9, 11, 14, 59, 61, 68, 70. Fig. 15 A-M Rhipicephalus sanguineus, 68 ^sapiens, Homo, 11 #Sauria, 9 scaevola, Amblyomma, 90 #Scincidae, 9 #Sciuridae, 12 *Sciurus, 17, 18, 21 *scrofa, Sus, 13 semermis, Haemaphy salis, 36 #Serpentes, 9 serpentinum, Amblyomma, 133 #sladeni, Callosciurus, 34, 59 Rattus rattus, 21, 34 ^sondaicus, Bos, 14, 34, 47, 119 Ros sondaicus, 14, 34, 47, 119 Rhinoceros, 13, 62, 85, 89, 114 Roricidae, 10 142 ENTOMOLOGICA AMERICANA spiniceps, Haemaphy salts, 31 spinicoxalis, Ixodes, 7, 8, 10, 11, 12, 13, 16, 21, 22. Fig. 3 A-G Ixodes, 21 spinigera, Haemaphy satis, 31 *spinosa, Geoemyda, 9, 35, 39, 95, 107 #Squamata, 9 *striatus, Sciurus, 17 #subcristatus, Hystrix, 65 * Hystrix subcristatus, 65 sublaeve , Amblyomma, 102 subluteum, Amblyomma, 85, 86 *Suidae, 13 *suillus, Hylomys, 10, 18 #Hylomys suillus, 10, 18 #sumatrensis, Rhinoceros, 115 supinoi, Amblyomma, 111 *Sus, 13, 51, 59, 62, 66, 80, 90, 94, 114, 116, 119 *syndactylus, Hylobates, 11, 34 ^Hylobates syndactylus, 11 taivanicum Amblyomma infes- tum, 116, 117 taiwana, Haemaphy satis corni- gera var., 31 #Tana, 31 *tana, Tupaia, 10, 31 *Tana, 31 *Tupaia tana, 10, 31 *Tapiridae, 13 *Tapirus, 13, 114 *taurus, Bos, 14, 27, 61, 62, 66, 130 *temmincki, Eattus bukit, 18 #Rattus fulvescens, 12, 18 tenimberense, Amblyomma, 98, 99 testudinarium, Amblyomma, 2, 3, 6, 7, 8, 9, 11, 13, 14, 78, 79, 114, 116, 117, 118, 133. Fig. 25 A-M Amblyomma infestum, 115 Amblyomma infestum var., 116 Haemalastor infestum var., 116 #Testudinata, 9 #thai, Viverricula indica, 59 *thibetanus, Enarctos, 35, 51 #Euarctos thibetanus, 35, 51 *tigris, Felis, 11, 17, 35, 39, 59, 65, 66, 114 *Timaliidae, 10 *torquatus, TJrsus, 35, 51 toxopei, Haemaphysalis, 7, 8, 10, 24, 25, 47 trabeatum, Aponomma, 129, 130 Aponomma fimbriatum, 130 tragnli, Haemaphysalis, 7, 8, 13, 24, 25, 47 ^Tragulidae, 13 *Tragnlus, 13, 48 *treubi, Eattus bukit, 18 #Rattus fulvescens, 12, 18 #tricarinata, Geoemyda, 103, 106 *Nicoria, 103 *Triglyphedon, 97 trimaculatum, Amblyomma, 129 Aponomma, 7, 9, 120, 129, 130, 131. Fig. 29 A-0 trimaculatus, Ixodes, 129 #tripudians, Naia, 129 143 ENTOMOLOGICA AMERICANA Vol. XXX, Nos. 1-4 *Tupaia, 10, 18, 21, 31, 34, 120 #Tupaiidae, 10 *Turdidae, 10 *Turdus, 34 undatum, Aponomma, 129 undatus, Acarus, 130 #unicolor, Cervus, 13, 27, 34, 44, 59, 65, 71, 72, 134 *Rusa, 62 *Urocissa, 21 *Ursus, 35, 40, 44, 51 # vaginalis, Muntiacus mnntjak, 34, 65 *Varanidae, 9 *Varanus, 9, 96, 97, 98, 102, 103, 106, 110, 111, 112, 113, 122, 124, 125, 128, 129, 130, 133 #verrucosns, Sus, 13, 116, 19 ^versurae, Tupaia belangeri, 21, 34, 120 *Vesperugo, 103 $ *i nttatus, Sus, 66, 114 *Sus cristatns, 13, 66, 114 *Viverra, 31, 40 *Viverricula, 59 *Viverridae, 11 wellingtoni, Haemaphysalis, 6, 7, 8, 10, 14, 24, 25, 48, 49. Fig. 9 A-L *Zamenis, 97 #zebrata, Corncia, 133 *Zosteropidae, 10 #Zosterops, 10, 16 144 Americana A Journal of Entomology. Volume XXXI (New Series) 1951 PUBLICATION COMMITTEE JOSEPH C. BEQUAERT, EDITOR GEORGE S. TULLOCH EDWIN WAY TEALE PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY 1951 JAN 1 0 1952 ENTOMOLOGICA AMERICANA YOL. XXXI (N. S.), 1951 CONTENTS PAGE The Tingoidea of New England and their Biology. Norman S. Bailey. December 19, 1951 1 6CTs> VOL. XXXI (New Series) Dll. iNS. U.S. NATL, ms: JAN 1 5 »52 Americana A Journal of Entomology. PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY PUBLICATION COMMITTEE JOSEPH C. BEQUAERT, Editor GEORGE S. TULLOCH E. W. TEALE Published for the Society by the Business Press Inc. N. Queen St. and McGovern Ave., Lancaster, Pa. Subscription, $5.00 per year Date of issue, December 19, 1951 r Notice to Subscribers of “Entomologica Americana” 1. Beginning with Yol. XXIX, “ Entomologica Americana” is no longer published as a quarterly. A yearly volume of approxi- mately 130 to 180 pages is issued instead. 2. The yearly volume will comprise one or more articles, ac- cording to the size of the individual articles. Each article will be issued when printed and bear its own date of publication. 3. Subscription is to the yearly volume. 4. Beginning with Yol. XXYII, every article is provided with its own Index. No other Index will be published for the complete volume, when this comprises more than one article. 5. The title page and contents of each volume are issued and mailed either with the article completing the volume or with the only article, in case the volume comprises one only. 6. Index and title page of Yol. XXVII. Nos. 3-4 of this volume comprise one article, published as a unit with its own index. The title page of this volume and an Index covering parts 1 and 2 only were published in July 1949. They were mailed with Nos. 1 and 2 of Vol. XXIX. Subscribers who lack this title page and Index should communicate with the undersigned Editor. el. Bequaert Editor, “ Entomologica Americana ” Museum of Comparative Zoology Cambridge 38, Mass. November, 1951 Volume XXXI THE TINGOIDEA OF NEW ENGLAND AND THEIR BIOLOGY* 1 By Norman S. Bailey Department of Biology, Boston University, Boston, Massachusetts Contents Page Acknowledgments 2 An Explanation 3 Introduction 3 General Considerations 4 The Tingoidea of New England 8 Family Piesmidae 16 Genus Piesma 16 Family Tingidae 19 Genus Alveotingis 20 Genus Melanorhopala 21 Genus Hesperotingis 22 Genus Physatocheila 24 Genus Leptoypha 27 Genus Dictyonota 30 Genus Acalypta 32 Genus Leptopharsa 35 Genus Galcatus 39 Genus Gargaphia 42 Genus Corythaica 47 1 The material in this paper was included in a thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at Harvard University. 1 ENTOMOLOGICA AMERICANA s. Genus Stephanitis 53 Genus Corythucha 62 Specific Characters in the Male Genitalia of Corythucha 106 Technique for Mounting the Male Genitalia 107 Method of Drawing the Male Genitalia 109 Taxonomic Significance of the Genitalia 110 Appendix 113 Check List of New England Tingoidea 113 Host Plants of New England Tingoidea 114 Conclusions and Suggestions 122 Selected References 123 Acknowledgments In the preparation of this paper much invaluable assistance has been freely given by a number of friends and fellow entomolo- gists. Much of the content of the thesis is directly traceable to their individual contributions. Without their aid, omissions and errors would necessarily be more numerous. During the early period of this study Professor H. M. Parsh- ley and Professor C. J. Drake both kindly checked determinations. Professor Drake has since answered several inquiries concerning the Tingidae and Professor Parshley generously allowed me to go through his extensive collection and select an appreciable number of duplicate Specimens for my own collection. The friendly help of both gentlemen is much appreciated. Through the kindness of the following individuals several col- lections of New England Tingidae were made available to me. Dr. M. E. Smith, Dr. Henry Dietrich, Professor J. G. Conklin, Profes- sor Herbert Knutson, Dr. C. L. Remington, Dr. R. B. Friend, and Dr. J. C. Bequaert allowed me to borrow the collections in their care. Some years ago notes were taken on the specimens in the Boston Society of Natural History collection. To all of these per- sons and the institutions they represent I express my appreciation for the privileges granted. Mr. H. G. Barber kindly sent me a list of the records of New England material in his collection. Dr. R. I. Sailer prepared a long list of the pertinent records and host plant data from the large United States National Museum collection, for which I am greatly indebted. He also assisted me on numerous occasions during the course of this study, and for all of his aid I am duly grateful. Others who have provided me with useful information include, the late Mr. J. R. de la Torre-Bueno, who furnished reprints and 2 Volume XXXI suggestions, Miss Hilda Vilkomerson, who sent me notes on Cory- thaica, and Dr. E. W. Baker who identified some of the mites asso- ciated with the tingids. Furthermore, I am deeply indebted to Mr. F. Y. Cheng of Harvard University for the splendid drawings of a Corythaica nymph (Fig. 2) and of the new species of Corythucha (Fig. 3) de- scribed herein. Mr. F. W. Maynard prepared the fine microscope slides of the cherry leaf sections which are shown in Fig. 4. The excellent photographic work was done by Mr. Frank White and Miss Ruth Dunn of the Harvard Biological Laboratories staff and by Mr. F. B. Burrill of Boston University. An additional word of thanks is due to Mr. F. B. Burrill, Dr. K. A. Christiansen, Mr. A. E. Feldman, Mr. C. A. Frost, Dr. S. K. Harris, Dr. A. G. Humes, Dr. W. L. Nutting, Mr. M. E. Richards, Dr. F. G. Werner, Dr. J. T. Woodland, and others who thought- fully remembered to collect lace bugs for me when they had the opportunity. With several of these colleagues and students, many pleasant hours have been spent collecting tingids. The patient support and encouragement of my wife, Ercolina, and the considerate interest of my family and associates have also contributed in large measure to the gradual completion of this work. Finally, the intelligent suggestions and the technical abilities of the typist greatly eased the arduous task of actual composition. In a very real sense, any merit the paper may have from the stand- point of accuracy is a good index of my debt to those who have so generously rendered these services and to many others who remain nameless but equally appreciated. An Explanation Throughout the paper previously unpublished species records are cited by naming, in parentheses, the collector or the institution in whose collection the specimens were seen, e.g., (Parshley) or (Cornell University). All native plant binomials are those used by the late Professor M. L. Fernald (1950) in his recent edition of Gray’s Manual of Botany and the names of cultivated plants follow Dr. L. H. Bailey’s (1949) Manual of Cultivated Plants. The family name Tingidae is used in accordance with Opinion 143 of the International Commission of Zoological Nomenclature. Introduction The extensive literature dealing with the Tingoidea of North America includes a wealth of information concerning matters taxo- 3 ENTOMOLOGICA AMERICANA nomic but only an occasional careful life history study and scat- tered notes on tingid biology. The relative unimportance of the lace bugs in an economic sense is responsible for this neglect. How- ever, even a cursory consideration of the Tingidae suggests a va- riety of problems biologically both interesting and significant. Before such investigations may be properly initiated, however, a summation of our present knowledge becomes necessary. This paper, therefore, represents an effort to collate, review, and expand our understanding of the Tingidae. Although New England is a restricted portion of the Nearctic faunal area, the species occurring there are sufficiently representative to provide a basis for such work. That the tingid fauna may be considered representative is sup- ported by the following evidence. The excellent review of the taxonomy of American Tingidae entitled “Generic Classification of the North American Tingoidea” (Hurd, 1946) reveals that these heteropterous insects have been most actively studied since 1900, although Fabricius described two American species as early as 1794. By 1886 Uhler could list only eleven genera and twenty-four species from North America. The next thirty years brought an increase to twenty-four genera and seventy-six species from the area north of Mexico (Van Duzee, 1917). Only twenty years later fifty-two genera containing about four hundred and twenty-four species were recorded from the Western Hemisphere (Drake and Poor, 1936). Forty-seven of the genera and over three hundred of the species occur in North Amer- ica. For eastern North America Blatchley (1926) recorded twenty- two of these genera and seventy-three species. This paper notes forty species and four varieties, falling into fourteen genera, for which there are authentic New England records. One of these species is described as new and one is a recent introduction from the Orient here first recorded from North America. Undoubtedly Blatchley 7s 1926 totals are no longer valid. However, if we con- servatively estimate the present number of known tingid species in eastern North America at approximately one hundred, it is prob- ably safe to assume that a third or more of the total number of species and about half of the genera east of the Mississippi are found in this region. Both in variety of genera and in number of species, therefore, the tingid fauna of New England is truly representative. General Considerations Although the main purpose of this investigation is to elucidate tingid biology, as usual in entomological studies, it is first necessary 4 Volume XXXI to determine as accurately as possible the species definitely recorded from the selected geographic area. The over-all problem, there- fore, involves a limited taxonomic review. The details of this re- view will only be included, however, if they are essential to a proper understanding of a particular species. Full bibliographic references will support all taxonomic conclusions and provide ready access to the literature which is available for the New England species of Tingidae. Two exceptions to the procedure just outlined follow. In the course of this study one new species has been encountered in the field and another species, apparently introduced rather recently from the Orient, has been collected. A full description of the new species is given and the oriental species is redescribed since the original description is in Japanese (Matsumura, 1905) and, al- though the description is translated in a later edition (Matsumura, 1930), both references are difficult to obtain and the English ver- sion is not very satisfactory. More than a third of the New England species belong in the wholly American genus Corythucha Stal. Over sixty species of Corythucha (Hurd, 1946) have been described. Fifteen of these occur within the New England area. The species in this genus exhibit considerable intraspecific variation. As a group they ap- pear to be of fairly recent origin and, employing the current taxo- nomic criteria, specific limits are difficult to determine in some cases. For this reason the external genitalia of males in this genus have been studied to learn whether or not their structural features would aid in the separation of the species. The method of prepa- ration and the results of this approach to tingid taxonomy will be included. Hurd (1946) has established a generic order that seems sound in the light of our present knowledge. Accordmgly, her sequence is followed. For the species an alphabetical arrangement is most satisfactory because of the limited representation of species in any one genus within the New England area. Unless all known species in a gexlus are compared, a phylogenetic order can scarcely be ap- proximated. The morphology of the Tingidae presents many clues to evolu- tionary relationships and many points of interest in general. All are of small size, and they are mostly somewhat dorso-ventrally flattened. The term “lace bug” is quite apt, since membranous structures on the dorsum of many species are of decidedly lace-like appearance. The hemielytra and expanded paranota display a re- ENTOMOLOGICA AMERICANA markably delicate areolate condition in some. In others the ner- vnres are more conspicuous and the surface is coarsely or finely punctate. The head is often wholly or partially covered by a more or less inflated pronotal hood. Although the closely related Pies- midae have two ocelli, the Tingidae lack simple eyes. The disc of the pronotum is either uni- or tri-carinate and is prolonged back- ward as a triangle which covers the scutellum in native species. The hemielytra lack both a distinct clavus and membrane and the cells may be hyaline, or in part whitish opaque, or fuscous. Sev- eral genera have marked tumid elevations in the discoidal areas of the hemielytra. Both the antennae and the rostrum are four- jointed. The tarsi have only two joints. In some species both macropterous and brachypterous forms occur. Intermediate hemielytral lengths may also be common. Sometimes macropterous specimens greatly predominate and bra- chypterous individuals are rare. In other species the reverse is true. The degree of hemielytral development may even be a sec- ondary sexual characteristic, and sexual dimorphism occurs in the antennae of some species also. Since we are concerned primarily with the biology of these heteropterous insects, the taxonomic details, although essential, are only of incidental importance. The species included have been thoroughly studied and carefully determined. However, the in- formation necessary for their identification has been reduced to a minimum and is presented chiefly in the form of a key supple- mented by figures which portray the diagnostic features of the Tingidae. This plan provides for a full elaboration of all biological information. Apparently all of the Tingidae are phytophagous. Therefore, a knowledge of their host relationships offers a promising approach to a study of their biology. It soon becomes evident that they show a remarkable diversity in their host preferences. The com- moner British species (Butler, 1923) are usually associated with herbaceous plants or with mosses, but the better known American species are generally thamnophilous or arboreal. The degree of host specificity also reveals an interesting range of variation. Since they are usually rather sedentary and in some species, at least, inclined to be gregarious, they may occasionally be collected in large numbers when the host plants are known. Consequently, all information pertaining to the food habits of the Tingidae will be fully considered. 6 Volume XXXI Although little, if any, information concerning their habitat preferences is available, it is obvious that the extremes are deter- mined by the ecological tolerances of satisfactory host plants. How- ever, it soon becomes apparent that certain species are much more restricted in their distribution than are their known hosts. It is only an exceptional species that almost invariably occurs wherever its food plant grows. Examples of both the extremes and of inter- mediate kinds of distribution patterns will be noted. What the reasons may be for local, disrupted patterns remains undetermined. Full distributional data will be given only for the less well-known species or where the plant-tingid relationship is of particular sig- nificance. A source of food is but one factor in the complex of such plant-insect associations. Probably one of the factors limiting the distribution of some species is the availability of suitable conditions for hibernation in the immediate vicinity. Such conditions may exist only locally within the broad range limits of tolerant host plants. That this may, in fact, be a vital matter is strongly suggested by one plant that itself furnishes favorable winter quarters for its insect para- sites. This host is one of the exceptional few that is nearly always infested regardless of local conditions or of its isolation from other plants of the same species. The problem of overwintering is commonly solved by hiberna- tion of the adults in the duff, under loose bark, or in other suitable shelters. Nymphs have been rarely collected in late winter or early spring and, therefore, some species may hibernate either as adults or as nymphs. One genus, common to North America, Europe, and Asia, overwinters in the egg stage. Species in this genus usually feed on broad-leaved evergreens and the eggs are inserted into the lower surface of leaves that remain on the plants throughout the year. Here again adaptations to the host appear. Seasonal and climatic influences on plant growth are naturally reflected in the reproductive cycles and activities of the Tingidae. This intimate interrelationship between the plants and the lace bugs deserves special emphasis. The available information con- cerning life histories and population trends will be broadly studied. In some instances only the seasonal range of the adults can be suggested from the records. In concluding this general introduction a statement of the methods employed seems pertinent. The information presented was gathered from three main sources. First, a thorough survey 7 ENTOMOLOGICA AMERICANA of the literature laid the foundation. Secondly, an appreciable amount of field work and collecting provided specimens and a num- ber of original observations. Finally the laboratory study of this material and of borrowed collections supplied valuable supplemen- tary information. The first method requires no further comment but a brief explanation of the other two seems desirable. The field work, extending over six seasons, included much gen- eral collecting as well as rather specific observations made regularly for two years at selected stations in the vicinity of Boston. During early September of 1949 a two thousand mile drive was taken for the express purpose of collecting tingids in Vermont, New Hamp- shire, and Maine. Short excursions into south-eastern Quebec and one along the lower St. Johns River in New Brunswick, Canada were also made. Although somewhat late in the season, the trip was satisfactorily productive. Eight collections of New England Tingidae have been carefully examined in addition to my own. Three of these are especially im- portant while some of the others contained individual specimens of note. These collections have been cited in the acknowledgments above. Records have also been made available to me from other sources. The most notable is a long list of the New England speci- mens in the United States National Museum very kindly supplied by Dr. Reece I. Sailer. Without the generous cooperation such as- sistance typifies, this study would be necessarily far less complete. The Tingoidea of New England Superfamily Tingoidea Reuter, 1912 There are but two families in this group. The members of both are plant feeders with walking legs. Their antennae consist of four antennites of which the third is the longest. The hind coxae are rotatory and the tarsus is composed of only two tarsites with arolia beneath the claws. The rostrum lies in a sulcus formed by usually prominent bucculae on the underside of the head which are commonly continuous with more or less raised sternal ridges on the thoracic segments. The bucculae and sternal ridges may be membranous with a single row of areolae. Key to the Families 1. Jugal spines projecting beyond the tylus, their tips free or contiguous. Scutellum exposed. Discoidal area of hemi- 8 Volume XXXI elytron divided by cubital vein. Lunate cavities visible beneath the paranota Piesmidae Piesma cinerea (Say)2 Jugae not surpassing the tylus. Scutellum covered by pro- notum in all native species. Discoidal area undivided. No thoracic cavities visible Tingidae Key to the New England Genera A. Head spines mostly stout and appressed. 1. Paranota usually poorly developed. Pronotum with three costate carinae and its anterior margin collar- like with no more than a suggestion of a hood. a. Hemielytra very convex with only the uniseriate costal area well defined. Small, blackish brown species with third antennite very stout and cov- ered with coarse recurved hairs. Alveotingis grossocerata 0. and D. b. Third antennite longer and slender, the distal end much swollen and fuscous. Large, straw colored species with hemielytral areas distinct. Costal margin uniseriate and flaring erect. Melanorhopala clavata Stal c. Third antennite much shorter and stouter than in (b), gradually increasing in diameter towards the distal end and clothed with recurved hairs. Costal area uni- or biseriate. Shorter, but other- wise somewhat resembling (b). Genus Hesperotingis 2. Paranota prominent but closely reflexed against disc of pronotum. Small hood evident at anterior end of median carina never projects forward beyond occiput. Genus Physatocheila 3. Paranota barely represented by a ridge. Carinae also nearly obsolete. Dorsal surface more nearly punc- tate than areolate. Very small Genus Leptoypha 4. Paranota with three to four rows of areolae, somewhat reflexed and concave above. Pronotal hood trans- verse and small, not covering head. Antennites of 2 If only a single species occurs in this region it will be indi- cated in this key. If more than one is known, refer to appropriate genus in the following keys. 9 ENTOMOLOGICA AMERICANA equal thickness, rough and setose. Costal area as wide as discoidal with two slight irregular rows of areolae Dictyonota tricornis var. americana Parshley B. Head spines short and stout, bluntly rounded and not ap- pressed. Three small, broadly oval and slightly convex species Genus Acalypta C. Head spines slender, long and freely erect. Paranota foliace-. ous, their areolae conspicuous. 1. Hood areolate but small, scarcely covering the occiput. Pronotal carinae sub-equal in height. Paranota nearly erect and narrow with one or two rows of areolae. Costal area also uni- or biseriate. General outline elongate rectangular Genus Leptopharsa 2. Lateral carinae greatly enlarged and swollen laterally, meeting behind the small areolate hood and well above the median carina. Triangular process of pronotum inflated. Areolae of paranota and hemielytra uniquely large and rectangular. Galeatus peckhami (Ashmead) 3. Hood and pronotal carinae as in C 1. Paranota much broader with three to five rows of areolae. Costal area also with three to five rows of areolae opposite apex of discoidal area. Broadly rectangular species with numerous long hairs on the nervures. Genus Gargaphia D. Head completely covered by membranous pronotal hood. 1. Hood elongate, low and flat. Somewhat rounded be- hind, gradually tapering anteriorly and reaching be- yond proximal end of third antennite. Pronotal cari- nae uniseriate and of sub-equal height. Very small. Corythaica bellula Bueno 2. Hood not drawn out anteriorly and not reaching be- yond first antennite. Paranota irregularly triseriate and flaring erect. Median carina high, arched, and much larger than the lateral carinae. Genus Stephanitis 3. Hood more or less globose behind, usually constricted and tapering in front to reach beyond the proximal end of antennite three. Paranota broadly rounded and essentially explanate with the center bullate. Median carina foliaceous and usually prominent. Genus Corythucha 10 Volume XXXI CAUDAL MARGIN GLASPER HYPANDRIUM CRENULATIONS TERGITE 10 -TERGITE 9 PARANOTUM. CARINAE TRIANGULAR PROCESS HOOD BUGCULA ROSTRUM PARANOTUM HYPOCOSTAL RIDGE MEDIAN CARINA DISCOIDAL ELEVATION HEMIELYTRON D. SPINES PARANOTUM CARINAE HEMIELYTRON OSTAL AREA DISCOIDAL SUTURAL SUBCOSTAL OSTIOLE 2ND ABDOMINAL ARANOTUM F. Fig. 1. Taxonomic anatomy of the Tingoidea. A, male genital capsule of Corythucha, dorsal aspect. B, Hesperotingis, dorsal as- pect, after Parshley, 1917a. C, Leptopharsa, dorsal aspect, after Parshley, 1923b. D, Corythucha, ventral aspect. E, Corythucha , lateral aspect, after Parshley, 1923b. F, Piesma, dorsal aspect. 11 ENTOMOLOGICA AMERICANA Keys to the New England Species of Tingidae3 Genus Hesperotingis 1. Third antennite fuscous beyond proximal third, fourth anten- nite sub-conical. Paranota reduced and reflexed. Costal area uniseriate and subcostal area biseriate. H. antennata Parshley ( H . antennata var. borealis Parshley is distinguished only by its slightly shorter and uniformly dark antennae and by haying the subcostal area irregularly triseriate.) 2. Third antennite longer, more arcuate and fuscous only beyond the middle. Fourth antennite fusiform. Costal area rather regularly biseriate and broad. Paranota foliace- ous and vertical H. illinoiensis Drake Genus Physatocheila 1. Rostrum not quite reaching mesocoxae. Costal area rather regularly triseriate P. brevirostris 0. and D. 2. Rostrum extending beyond the mesocoxae but not beyond the metacoxae. Costal area rather regularly biseriate. P. plexa (Say) 3. Rostrum extending beyond the rostral sulcus to the base of the second abdominal sternite. Costal area varies between a biseriate and triseriate condition P. variegata Parshley Genus Leptoypha 1. Costal area biseriate in humeral region and uniseriate posteri- orly. Lateral carinae barely discernible. L. costata Parshley 2. Costal area with only a few distinct cells at constriction before the apex. This species is much smaller and has shorter antennae than the other two L. ilicis Drake 3. Very similar to L. ilicis but with distinguishable areolae along the costal margin from the constriction of the hemielytron to the apex. It is also appreciably larger and has longer antennae L. mutica (Say) Genus Acalypta 1. Pronotal carinae almost parallel, foliaceous and uniseriate, their anterior ends in line. Median carina but slightly higher 3 Except for the species keyed out in the preceding key to the genera. 12 Volume XXXI than laterals. Paranota broad, rounded and mostly bi- seriate. Costal area regularly uniseriate. A. lillianis Bueno 2. Lateral carinae only areolate in front and slightly divergent behind. Median carina slightly higher in front and more prominent. Paranota narrow, biseriate in front and uni- seriate behind, areolae small. Costal area narrow and almost uniseriate A. nyctalis Drake 3. Paranota broad and rectilinear. Lateral carinae short, very low, and strongly divergent posteriorly. Median carina raised in front. Costal area with one, two, or three rows of areolae at different points A. thomsonii Stal Genus Leptopharsa 1. Costal area rather regularly biseriate. Median head spines rela- tively short, stout and blunt, lateral pair reduced and ap- pressed L. clitoriae (Heidemann) 2. Head spines needle-like, tips fuscous. Hood with three rela- tively large areolae on each side of the median nervure from front to back. Costal margin irregularly uniseriate except for biseriate section opposite apex of discoidal area, its nervures mostly fuscous L. heidemanni (0. and D.) 3. Head spines needle-like and whitish. Hood with four rela- tively small areolae on each side of the median nervure. Costal margin more regularly uniseriate than in (2) and a shorter portion biseriate. Costal nervures colorless ex- cept at apex of hemielytron L. oblong a (Say) Genus Gargaphia 1. Lateral head spines reduced, appressed and not visible from above because of hood. Paranota with three rows of areo- lae at widest point. Paranota and pronotal carinae of sub-equal height. Nervures of pronotal area and of prox- imal region of hemielytra covered with long, soft hairs. G. angulata Heidemann 2. Head spines prominent. Paranota conspicuously high and an- gulate, with five rows of areolae at widest point. Hood laterally compressed and sub -triangular in outline, its crest sub-equal in height with paranota. The pronotal carinae much lower and the three of equal height. First and second antennites fuscous as are some of the dorsal 13 ENTOMOLOGICA AMERICANA nervures. Very long hairs on the pronotal and hemielytral nervures G. solani Heidemann 3. Head spines prominent and white. Paranota rounded and with four rows of areolae at widest point. First and second antennites pale stramineous. Disc of pronotum invested with whitish pubescence. Hairs on nervures finer, shorter and much less obvious. Nervures all whitish except for two short sections near the widest part of the discoidal area G. tiliae (Walsh) Genus Stephanitis 1. Hypocostal ridge uniseriate. a. Hood nearly twice as high as crest of the median Ca- rina 8. globulifera (Matsumura) b. Hood and median carina of sub-equal height. 8. pyrioides (Scott) 2. Hypocostal ridge irregularly biseriate 8. rhododendri Horvath Genus Corythucha 1. Hood and median carina of sub-equal height. A. All membranous structures whitish except for a small fuscous spot on the discoidal elevation. Male genital capsule large (Fig. 5E) and lacking crenulations be- fore insertion of claspers. Claspers large with un- tapered, bluntly rounded tips C. ciliata (Say) B. Basal band and sutural area infuscated. Apical band never developed. Male genital capsule slightly smaller than 1A above and with crenulations before insertion of claspers (Fig. 5D). Claspers tapering and sub-acute at tips C. caryae sp. nov. C. Usually with both a basal and an apical hemielytral band present, but the pigmentation is variable and the apical band may be entirely lacking. The male genital capsule is small and fragile (Fig. 5A) and the claspers taper to acute points C. arcuata (Say) D. Both hemielytral bands invariably present. Male genital capsule as in Fig. 6G. The median caudal border of the hypandrium is slightly convex in this species alone and the claspers have unique spatulate tips C. pruni 0. and D. 2. Hood noticeably higher but less than twice as high as the median carina. 14 Volume XXXI A. Male genital capsule lacking crennlations before inser- tion of claspers which taper gradually to bluntly rounded tips (Fig. 5H) C. heidemanni Drake B. Male genital capsule with crenulations. Claspers taper to rather sharp tips (Fig. 6A) C. juglandis (Fitch) 3. Hood at least twice as high as the median carina. A. Both hood and carina very small and low. a. Unique in lacking marginal spines on paranota and hemielytra. Male genitalia as shown in Fig. 6D C. mollicula 0. and 1). b. Hood well rounded behind. Anterior lobe of para- nota with large spot and apical hemielytral band prominent. Male genitalia as in Fig. 6F. C. pergandei Heidemann c. Hood somewhat flattened on top. Paranota with very small spot and apical hemielytral band lack- ing. Male genitalia as in Fig. 6H. C. ulmi (0. and D.) B. Hood high and somewhat laterally compressed. a. Most areolae are whitish opaque and many ner- vures are lightly embrowned. Male genital cap- sule small and the claspers with extremely long slender, sharp tips (Fig. 6B). C. marmorata (Uhler) (C. marmorata var. informis Parshley differs chiefly in having very short hemielytra that are usually broader than long. Their discoidal area extends behind the middle. Male genitalia are almost identical, as shown in Fig. 6C.) C. Hood high and globose. a. Species 3.5 to more than 4.0 mm. long. al. Cells of the hood about four times as large as the paranotal areolae. Hood nervures dark. Male genitalia as in Fig. 5B. C. associata 0. and D. bl. Cells of the hood about twice as large as the the paranotal areolae. Hood nervures pale. Male genitalia as in Fig. 6E. C. pallipes Parshley b. Species 2.8 to 3.5 mm. long. a2. Paranota whitish-opaque and nearly immacu- . 15 ENTOMOLOGICA AMERICANA late. Cells of the hood about four times as large as the paranotal cells. Genital capsule of the male small and the claspers taper to sharp tips (Fig. 5F). Usually less than 3.0 mm. long C. coryli 0. and D. b2. Paranota sub-hyaline and bearing a conspicu- ous brown spot on the antero-lateral face of the tumid area. Cells of the hood about twice as large as the paranotal cells. Geni- tal capsule of the male larger than in C. coryli and claspers with blunter tips (Fig. 5C). This species is usually of intermediate size between a2 and c2 C. bellula Gibson c2. Bullate area of paranota heavily infuscated. Genital capsule of the male larger than that of C. bellula and claspers with sharp tips (Fig. 5G). Usually over 3.0 mm. long and darker than other species in this region. C. cydoniae (Fitch) Family Piesmidae Amyot et Serville, 1843 A single genus represents this family in North America and Europe. In the Nearctic ten extant and one fossil species occur. About a dozen species are known in the Old World (Hurd, 1946). The piesmids show affinities with the Lygaeidae and in the past some students have placed them with the Lygaeidae. Recent hemip- terists, however, have more frequently associated them with the Tingidae. Macropterous piesmids have ocelli and both the clavus and membrane of the hemielytron are distinct. Their external geni- talia are much less prominent, especially in the males, than are those of the tingids. - Genus Piesma Le Peletier et Serville, 1828 Piesma cinerea (Say). Tingis cinerea Say, 1832; in Fitch reprint, 1858, p. 793; Say (in LeConte edition), 1859, p. 349. Piesma cinerea (Say) var. inornata McAtee, 1919a, p. 87. •This species and its variety may be treated together since the latter differs only from the usual form in its entire lack of dark markings and since, according to McAtee (1919a), it may occur 16 Volume XXXI anywhere within the range of the species. Piesma cinerea, and its variety inornata, is the only piesmid known from New England. There are specimens of the variety in the Parshley collection from Rhode Island and from Connecticut. Adults vary in size from 2.75 to 3.25 mm. in length and from 1.3 to 1.4 mm. in width. They may be separated from all other tingoids in this region by the characters given in the family key. This is one of the commonest and most widely distributed spe- cies. P. cinerea ranges from Massachusetts to the state of Wash- ington, south to California and Florida. It occurs in Canada, Mexico, and Argentina (McAtee, 1919a; Hurd, 1946). In New England there are records of its occurrence in Massachusetts, Rhode Island, and Connecticut (Parshley, 1917b and 1923b), but it is apparently unknown from the three northern states. Eggs: The eggs are white to light yellow and are laid on the food plant. Most are placed on the under side of the leaves and close to the larger veins. Unlike tingids, piesmids do not insert the abopercular end of the egg into the leaf tissue. They simply deposit an egg on its side and fasten it lightly with some adhesive substance. In heavy infestations eggs may be laid along the impressed veins of the upper leaf surface, in the leaf axils, or in the dense flower heads of Amaranthus retroflexus L. The eggs are laid individually and in no regular pattern. Twenty or more may be found on a single leaf (Weiss and Lott, 1924b). Barber (1924) has given a good description of the egg but my observations make some modifications necessary. The eggs vary in length from about .68 to .72 mm. and in diameter from .25 to .28 mm. The sub-cylindrical chorion tapers to a rounded point at the abopercular end and bears somewhat irregularly spaced longitudinal ridges. The ridges can be seen rather distinctly on the exposed side of the egg, but when the egg is freed from the plant the attached side is seen to be relatively smooth. The opercular end is only slightly constricted. A low ridge borders the operculum which is a little convex and bears a ring of four to seven minute hemispherical protuberances. These chorial processes are evenly spaced and form a ring about midway between the center and the edge of the cap. Barber (1924) states that the operculum is flat and bears only five processes around the edge. Although his observations were made in the Boston area also, local material now before me differs in the features indicated. The convexity of the operculum probably depends on the age of the embryo to some extent, while it may also be affected by preservation. In the eggs at hand the compound eyes are readily discernible as bright pink 17 ENTOMOLOGICA AMERICANA spots seen through the transparent chorion. This suggests that the embryo was fairly advanced in development when the eggs were pre- served. There is some variation in the number of chorial processes on the opercula of this species. On the few eggs examined (about 10) the number varied from 4 to 7 with 6 occurring most frequently. There are eggs in my collection taken on June 30 in Boston (Hyde Park) on Amaranthus retroflexus L. Weiss and Lott (1924b) found eggs abundant on July 25 on the same host in Moores- town, New Jersey, and observed a few as late as August 30. They also noted adults in copulation on the last date. Therefore, ovipo- sition probably occurs throughout the season from soon after emer- gence from hibernation until the temperature reaches a critical low (as yet undetermined) in late summer or early fall. Nymphs : Barber (1924) has described and figured the five nymphal instars. Since his paper is readily available, and since the nymphs are almost invariably found in association with adults, details will be omitted here. It is desirable to emphasize the total lack of dorsal spines on nymphs of this species, however, since the related tingid nymphs are conspicuously spinose. In their notes for July 25, Weiss and Lott (1924b) point out that all stages of Piesma cinerea (Say) were then present but that adults and last instar nymphs greatly outnumbered the others. On August 30 a fair number of adults were seen, but very few fifth instar nymphs. Adults: According to McAtee (1919a), piesmids are partial to plants of the family Chenopodiaceae. For this reason they are potential pests of such crops as beets and sugar beets. A European species, P. quadrat a (Fieber), is recorded as the only known vector of the sugar beet leaf curl virus on that continent (Wille, 1929). Early reports in this country indicate that P. cinerea (Say) will feed on this host (Bruner, 1891 ; Osborn and Gossard, 1891). How- ever, it has apparently not yet become seriously destructive. Drake (1928a) records Chenopodium album L. as its preferred host. It is said to feed on the young leaves and flower buds of grapes in spring (Walsh and Riley, 1868; Summers, 1891), on the foliage of oak, sycamore, buckeye, and beech, and on Conocarpus erecta L. in Florida (Blatchley, 1926). Weiss and Lott (1924b) found them feeding in the flower heads of Scirpus at ro virens Muhl. on June 7 and on August 9 in New Jersey. Barber (1924) records them from three species of Amaranthus. He states that A. caudatus L. was completely destroyed by them. The leaves curled and dropped and buds of partly grown plants were blighted. A form of A. hybridus 18 Volume XXXI L. and the weedy A. retroflexus L. were also infested. Feeding results not only in a curling under of the leaf margins which shelters the attacking nymphs and adults, but also causes a white mottling of the upper surface. This discoloration of the foliage is so evident that infested plants are readily located. My collections have almost always been from A. retroflexus L. which is a common garden weed. Since the Amaranthaceae are closely related to the Chenopodiaceae, these host selections are not surprising. However, Barber andWeiss (1922) report them as locally common on horse-chestnut in New Jersey. Professor Parshley (1917b) found one in ocean drift. One specimen in my collection came from a Japanese beetle trap. Hibernation is undertaken by adults and they have been found under bark of oak and sycamore and about logs (Blatchley, 1926). To these situations Drake (1928a) adds bark of birch and in fallen leaves. A specimen in the Parshley collection was found hiber- nating in a spider egg case. They probably crawl into any suitable and convenient shelter when cool weather comes. In a long series taken on Amaranthus in West Newbury on July 22, 1950, a single specimen was found with a larval mite attached just under the base of the right hind wing. This is the only direct evidence of parasites, though they may be more frequent than this would indicate. The mites are of such small size that in such a position they would be easily overlooked, and it is only by chance that an occasional one would be noticed. Adults are active locally from June 16 to August 24 at least, since my collections include those dates. Actually, their season of activity probably extends from a few weeks earlier in the spring until somewhat later in the summer. As for so many common species, more accurate information is not available. Records for eggs and nymphs suggest more or less continuous reproduction throughout the summer with perhaps two main broods. Develop- ment from egg to adult may require about six or seven weeks. Veri- fication of the number of annual broods and of the cycle from egg to adult is yet to be done, however. Family Tingidae Laporte, 1832 Members of this family may be as small or somewhat larger than piesmids. Species with the lacy, expanded paranota and hemielytra are rather rectangular in outline, while those in which these structures are more minutely areolate- are usually more nearly oval in shape. In all New England tingids the pronotum has a pos- terior triangular process that covers the scutellum. They never 19 ENTOMOLOGICA AMERICANA have ocelli. The hemielytra lack a clavus and the membrane is undifferentiated. The tylus always exceeds the jugae which are inconspicuous. Their more general characteristics have been pre- viously discussed. Thirteen genera occur in the New England area. Genus Alveotingis Osborn and Drake, 1916 Alveotingis grossocerata Osborn and Drake, 1916a, p. 245, Fig. 9. This genus and species is based on a specimen taken on August 5, 1913, by Professor Osborn in Orono, Maine. A figure of this brachypterous form accompanies the original description. Parshley (1917a) notes that the type is a male and points out that the plate is inaccurate in some respects. The antenniferous tuber- cles are not as shown but developed as in the related genus Melanorhopala Stal. The third antennite is evenly clavate and not fusiform as drawn. Osborn and Drake (1917a) give a photograph of the type and note the same antennal errors in their original figure. Although Professor Parshley also considered the hemielytral areas more distinguishable than the drawing suggests, they are certainly not readily discerned in the brachypterous individuals before me. Undoubtedly there is some variation in this feature. These speci- mens agree with his other criticisms, however. The rostrum reaches the mesocoxae. A. grossocerata is rarely collected and nothing is known of the early stages nor of the habits of this small tingid. Macropterous adults are 3.4 mm. long and 1.4 mm. wide, while the brachypterous forms are 2.0 mm. long and 1.2 mm. wide. The species has been found in these additional localities : Great Barrington, Massachusetts (Bueno, 1931) ; Franconia (Parshley, 1920c), Mt. Washington, and Durham, New Hampshire ; New Haven (two records), North Branford, Westport, and Easton, Connecticut. Of the four specimens in the New Haven Experiment Station col- lection, two are females. One of these is macropterous. This is also true of the Great Barrington specimen reported by Bueno (1931) and of a female from Mt. Washington described by Parshley (1917a). The seasonal range is suggested by two Connecticut records for the year 1920. The earliest was June 12 and the latest August 16. Only three bits of ecological information are available. Bueno (1931) noted that the Great Barrington specimen was found on a rock on the shore of Lake Buel and he took one elsewhere on a thistle (Drake, 1928a) . One of the New Haven specimens bears a pin label stating that it was swept from grass. 20 Volume XXXI In addition to the New England records, the species was found in White Plains and on Long Island in New York (Drake, 1928a) and is also known from the Delaware Water Gap in Pennsylvania (Blatchley, 1926). The genus is restricted to North America. Genus Melanorhopala Stal, 1873 Melanorhopala clavata Stal, 1873, p. 130. Tingis ( Melanorhopala ) lurida Stal, 1873, p. 131. Tingis ( Melanorhopala ) uniformis Stal, 1873, p. 131. Cantacader henshawi Ashmead, 1886, p. 20. Melanorhopala obscura Parshley, 1916b, p. 167. Melanorhopala reflexa Blatchley, 1926, p. 492. In several papers Professor Parshley (1917a, 1919a, 1920c) and Professor Drake (1926, 1930) cleared the confused taxonomy of this species. This confusion arose when Stal described M. clavata, M. lurida, and M. uniformis without noting that certain antennal characteristics were always associated with males and others with females and without making allowances for structural variations associated with the brachypterous or with the macropterous con- dition. Drake (1926) finally obtained drawings of Stal’s types, which confirmed the view that M. lurida Stal is the male of M. clavata Stal and that M. uniformis Stal is a brachypterous female of the same species. Parshley (1919c) had previously decided that his M. ohscura was probably the male of M. clavata Stal and also synonymous with M. lurida Stal. In 1926 Blatchley described M. reflexa from Florida. Drake (1930), having examined Blatchley ’s type, relegated M. reflexa Blatchley to the synonymy of M. clavata Stal also. In his opinion it was simply an extremely long-winged individual with no other indications of specific distinctness. Sexual dimorphism, combined with the common tendency for variation in degree of wing development among the Heteroptera, was largely responsible for the synonymy of this species. M. clavata Stal is widely distributed in North America. It ranges from Manitoba, Wyoming and Colorado to Maine, Long Island (Hurd 1946) and Florida (Drake, 1930). New England records include Maine, New Hampshire, Massachusetts, Rhode Island, and Connecticut. Although several collections contain numerous specimens, little is recorded concerning their habits. A long series collected on weeds near Chicago (Drake, 1930) is mentioned. Blatchley (1926) took specimens while sweeping herbage in dense woodlands and low meadows, and by beating foliage 21 ENTOMOLOGICA AMERICANA of black oak. He also notes that Bueno got them while sweeping rank weedy places near swampy meadows. The few I have taken in Maine and Massachusetts were swept from herbs where Solidago was a common element in the cover. Professor Parshley told me that he took many on Solidago on Long Island. One of the two other species now recognized in this genus (M. infuscata Parshley) was recorded from Liriodendron in Maryland and Virginia. An individual of this species was actually found at sap of this tree (Parshley, 1920c). Eggs and nymphs of Melanorhopala have not been described. Adults range in size from 4.25 to 6.8 mm. long and are 1.4 to 2.0 mm. wide. Therefore it is one of our larger tingids. The earliest New England record at hand is June thirteenth for Branford, Connecticut (Parshley, 1923b) and the latest is August twenty- seventh for a specimen from Swans Island, Maine and for one from Durham, New Hampshire. Adults are probably active from early June until September or even later. Genus Hesperotingis Parshley, 1917a Hesperotingis antennata Parshley, 1917a, p. 21, figure 2. Hesperotingis antennata var. borealis Parshley, 1917a, p. 23. Little information is available concerning the habits of Hes- perotingis antennata Parshley. This medium-sized tingid has been so rarely taken that less than a score of records were located ante- dating Bueno’s paper of 1946. Nothing is known concerning the immature stages nor of the actual food habits of this species. The variety borealis is known only from the holotype and possibly two other specimens (Froeschner, 1944; Hurd, 1946). The former was taken in Hampton, New Hampshire, and one of the others came from the District of Columbia, while the third was taken in central Missouri. This variant, therefore, probably occurs throughout the known range of the species since the records of typical forms in- clude Franconia, New Hampshire (Cornell University collection), Dunedin, Florida (Blatchley, 1928b), and Columbia, Missouri (Froeschner, 1944). Adult females vary in length from 3.7 to 4.5 mm. and are 1.5 mm. wide. Variation in length is primarily a matter of hemielytral development. Individuals with long, short, or with intermediate wing lengths are mentioned. In three sum- mers, Bueno (1946) collected the notable total of 504 specimens in 4 The figures given by the author are not consistent with his recorded specimens. 22 Volume XXXI the same locality at Tannersville, New York. Eleven, or about 20%, were macropterous. New England records include Franconia, Hampton (August 15), and Durham (July 16), New Hampshire; Northboro (June 24), Massachusetts; New Haven (September 4; Olsen, 1923) and Granby (July 4), Connecticut. The earliest seasonal record seems to be June 24, 1937 (Northboro, Massachusetts; C. A. Frost) and the latest September 6, 1932 (Tannersville, New York; Bueno, 1946). Concerning the habitat preferences of H. antennata Parshley, Bueno (1946) states: “They were taken by sweeping low — in fact by scraping the ground itself — on both sides of a footpath running through bushes and on the mowed area, in the longish grasses, close to their roots. They also seemed to be found in the shade, preferably of willow; the foodplant was indeterminate, under the conditions. There was the usual weedy growth — grasses, sedges, trailing blackberry, goldenrods, plantain, meadow sweet, overgrown here and there by stunted willows, young aspens and blackberry bushes, except, of course, in the grassy part of the meadow — a wide choice of food plant, in the absence of direct observation. ’ ’ Again : ■ ‘ The greater part of the specimens was taken either on a footpath about 50 feet long, through bushes; or else along the edges of a tree-grown space not over 100 feet long. The species was sought in other parts of the field, but without success, except in one instance. It was never found in or among tall grasses.” And : ‘ 1 The only fairly certain thing about the habitat is that they are found more or less in the shade of trees and bushes, since they seem preferably swept along the edges of footpaths or of mowed areas where the grasses are again tall.” The University of New Hampshire collection includes the Dur- ham specimen which bears the label ‘ ‘ on Andropogon. ’ ’ Froeschner (1944) records one of the Missouri specimens as “swept from an overgrown field that had formerly been under cultivation. ’ ’ A final bit of ecological detail is reported by Blatchley (1928b). On three occasions in January he obtained several brachypterous individuals by beating bunches of Spanish moss hanging from the lower limbs of Quercus rubra L. in open pine woods in Florida. None were taken from the same epiphyte on pines nor on other trees in the same area. This indicates hibernation of adults, and the exclusive association with the oak may or may not be significant. 23 ENTOMOLOGICA AMERICANA Hesperotingis illinoiensis Drake, 1918, p. 88. Four tingids collected in South Meriden, Connecticut on July 3, 1939, by Mr. Harry L. Johnson, and now in the New Haven Agri- cultural Experiment Station collection, could not be certainly iden- tified with any known species on the basis of published descriptions alone. Since types and paratypes were not available, it was neces- sary to send the four specimens to Professor Drake who kindly de- termined them as H. illinoiensis. He originally described this spe- cies from three individuals taken in Palos Park, Illinois, by Mr. W. J. Gerhard. The type is not in the Cornell University collection as erroneously reported (Drake, 1918). These two collections represent the only records for this spe- cies as far as I can discover. The Connecticut material includes a female and three males. In length they vary from 3.8 to 4.2 mm. and in width from 1.3 to 2 mm. Two rather regular rows of costal areolae readily distinguish H. illinoiensis Drake from the other known hesperotingids. Nothing is recorded concerning the biology of this species. Like the genus Melanorhopala, Hesperotingis is strictly North American in distribution. Genus Physatocheila Fieber, 1844 Physatocheila brevirostris Osborn and Drake, 1916a, p. 243. This is another species known only from scattered records of adults. There is nothing available concerning its habits nor about its biology. In size and coloration P. brevirostris 0. and D. closely approaches P. plexg (Say). The two are separable on the basis of the short rostrum and triseriate costal area of the former. Two males before me, collected by Mr. C. A. Frost in Natick, Massachu- setts, have remarkably short beaks that just fail to reach the meso- coxae and their costal areolation is fairly regular. One specimen in the New Plaven Experiment Station collection, taken on May 9 in New Haven by Mr. H. P. Zappe, appears to be identical. How- ever, four others in the same collection, identified as P. brevirostris 0. and D. by Professor Parshley some years ago, have much longer rostra and the costal margins are more nearly biseriate, though partly and irregularly triseriate in two individuals. These four seem more properly P. plexa (Say). Variation in hemielytral areolation is very likely to occur in tingids. Therefore it is unfor- tunate that larger series are not at hand for comparative analysis. In other tingids there is also enough variation in the length of the rostrum to make reliance on that feature alone unsatisfactory. 24 Volume XXXI Under present circumstances this interpretation must stand for want of more adequate materials. Although, for the above reasons, records in the literature are certainly questionable, P. brevirostris O. and D. has been reported from Massachusetts and Connecticut only for New England. Elsewhere it apparently ranges from New York and New Jersey west to Ohio and south to Virginia (Blatch- ley, 1926). The recorded seasonal activity covers the period from early May into early July. This, however, is undoubtedly subject to extension when more information becomes available. In color the adults are a uniform soft reddish brown. They are about 3.2 mm. long and about 1.6 mm. wide. Physatocheila plexa (Say), 1832. Tingis plexus Say, 1832; in Fitch reprint, 1858, p. 794; Say (in LeConte edition), 1859, p. 349. Physatocheila parshleyi Osborn and Drake, 1917b, p. 156. ( non Physatocheila plexa Osborn and Drake, 1916a, p. 242, 1917b, p. 156.) There must always be a degree of uncertainty in the recog- nition of Say’s species. However, in this instance the interpretation of Professor Parshley (1916b) seems the more plausible. Van Duzee (1917b), Barber and Weiss (1922), and Hurd (1946) also concur with Parshley rather than with Osborn and Drake (1917b). P. plexa (Say) is distinguished from P. brevirostris 0. and D. by having the rostrum extend beyond the mesocoxae and by having the costal area usually rather regularly biseriate. Parshley (1916b) describes the color as a uniform dull yellowish brown and states the length as 3.0 to 3.2 mm. and the width as 1.1 mm. There are many more records for this species than for P. brevi- rostris 0. and D. Its known range extends from Quebec south to Virginia and from Maine to Oregon (Hurd, 1946). There are New England records for Maine, Massachusetts, Rhode Island, and Con- necticut. Mr. C. A. Frost collected specimens as early as May 2 and as late as November 12 by sifting duff in Ashland and Sherborn, Mas- sachusetts. This indicates hibernation of the adults. Blatchley (1926) also notes that Bueno took specimens on oak and McAtee found them on hickory and on laurel. Proctor (1946) reported the species from Great Heath on Mt. Desert Island, Maine. Barber and Weiss (1922) record material from willow in New Jersey. These few ecological items are no more than hints concerning the possible 25 ENTOMOLOGICA AMERICANA host plants of P. plexa (Say). This species is apparently active from early May (May 2, Kingston, Rhode Island, Parshley, 1917c) until late September (September 27, Mt. Desert, Maine, Proctor, 1946). Localities cited for P. plexa (Say) in New England include Westbrook, Maine (Parshley, 1914) ; Pinkham Notch, New Hamp- shire (Museum of Comparative Zoology) ; Wilmington, Massachu- setts (fide Sailer) ; Providence, Rhode Island (Parshley, 1917c) ; and Brookfield (Parshley, 1923b) and Lyme ( fide Sailer), Connec- ticut, in addition to those mentioned above. Physatocheila variegata Parshley, 1916b, p. 166. Physatocheila plexa Osborn and Drake, 1916a, p. 242. P. variegata Parshley is the most frequently collected of our three New England species. It is also most easily distinguished. In color it is predominantly greyish brown, but there is a mottling effect of darker and lighter areas. The areoles of the costal area show considerable variation between a biseriate and a triseriate condition. The arrangement is usually intermediate between these extremes and noticeably irregular. The length of the rostrum is also diagnostic. It always exceeds the length of the rostral sulcus and usually projects beyond the base of the second abdominal ster- nite, often reaching the middle of that segment. P. variegata reaches a length of 3.4 to 4.0 mm. and a width of 1.3 to 1.5 mm. Again there is no definite information concerning its food habits. A few field notes suggest an association with trees when they flower in spring. Barber and Weiss (1922) report the species from hickory and willow (May 21) . White pine is noted by Blatch- ley (1926). Froeschner (1944) took two females from small "branches of a willow near a stream in Missouri (May 11). Proctor (1946) mentions specimens from catkins on Mt. Desert Island, Maine, April 27). There are specimens in the United States Na- tional Museum ( fide Sailer) taken on willow catkins in Bar Harbor, Maine (April 28 and May 11). Proctor (1946) also recorded speci- mens from the Great Heath on Mount Desert Island (September 25). Mr. C. A. Frost has collected this species in Framingham, Massachusetts by sifting on November 17 and 25 and on December 25 ( fide Sailer). This is all that is known concerning the hiberna- tion of adults. He has also collected them by sweeping in Sherborn (May 30 and June 20) (fide Sailer) and in Framingham (June 1), Massachusetts. The extent of seasonal activity is shown by the Mt. Desert Island records for April 27 and September 25 given 26 Volume XXXI above. New England localities include Orono, Maine (University of Massachusetts) ; Zealand Camp in the White Mountains (Uni- versity of Massachusetts) and Durham (University of New Hamp- shire), New Hampshire; Blue Hills, Holliston (Museum of Com- parative Zoology), Natick (Blatchley, 1926), Ware (H. G. Barber) and Wellesley (Parshley), Massachusetts; and Brookfield and Port- land, Connecticut (Parshley, 1917c), in addition to those previ- ously listed. Outside of New England the species ranges from New York (Osborn and Drake, 1917b) to British Columbia (Parshley, 1919c) and south to Missouri (Froeschner, 1944) and Virginia (Os- born and Drake, 1917b). The last reference lists most of the states from which the species has been recorded. In this genus about thirty species are recognized. Only four are North American. The rest are chiefly Palearctic in distribution. Genus Leptoypha Stal, 1873 Leptoypha costata Parshley, 1917a, p. 16. Leptoypha distinguenda Heidemann, 1917, p. 218, PI. 17, Fig. 1. The genus Leptoypha is primarily North American, and three species of these small tingids are recorded from New England. L. costata Parshley is easily distinguished from the other two by its costal area which is biseriate in the humeral region and then uni- seriate to the apex of the hemielytron. L. mutica (Say) has the costal area so reduced that areolae are distinguishable only behind the middle of the hemielytron and even fewer areolae may be seen in L. ilicis Drake. The lateral carinae are barely discernible in L. costata , but quite obsolete in the other two species. A specimen of L. costata before me is noticeably broader and much paler than any local material of L. mutica at hand. This male measures 2.7 mm. long and 1.15 mm. wide. Parshley (1917a) gives the length of the female as 2.8 mm. and the width as 1.3 mm. The only New England record located for L. costata Parshley is on the list kindly furnished by Dr. Sailer. The locality is Stam- ford, Connecticut (July 20, 1939), and carries the notation “reared from red ash.” A Louisiana specimen on the same list was taken on Hicoria alba (now Carya tomentosa Nutt.). McAtee (1923) re- ported a collection ftom Fraxinus caroliniana Mill, and suggests that an earlier indication that witch-hazel is a food plant for this species is probably an error since repeated sweeping of witch-hazel by him failed to yield any specimens. Drake (1925) took large numbers of L. costata Parshley on Fraxinus in Mississippi, which 27 ENTOMOLOGICA AMERICANA further supports McAtee’s view. Outside of New, England the species is known from Maryland, the District of Columbia, Virginia, Colorado, Arkansas, and Illinois (Drake, 1918). Leptoypha ilicis Drake, 1919a, p. 420. Drake (1919a) emphasizes the close affinity of this species and L. mutica (Say). It differs from the latter in being much smaller and in having shorter antennae. L. ilicis is said to be dark reddish brown and, like L. mutica, lacks the lateral carinae and has even fewer costal areolae. These features separate this species from L. costata Parshley. The length is given as 2.21 mm. and the width as .87 mm. The few records (Drake, 1919a, Blatchley, 1926 and 1928b) report this species from Stone Mountain, Georgia, and from Flor- ida. However, Hurd (1946) adds Texas, Oklahoma, and New Hampshire to the list. The distribution of L. ilicis is so predomi- nantly southern that the New Hampshire record certainly needs verification. As the specific name implies, this species was collected on Ilex in Georgia (Drake, 1919a). Blatchley (1928b) found one while beating Vaccinium and Hurd (1946) records palm jungle sweep- ings as a source of specimens. Leptoypha mutica (Say), 1832. Tingis mutica Say, 1832; in Fitch reprint, 1858, p. 794; Say (in LeConte edition), 1859, p. 349. Leptoypha mutica (Say) is better known than the two species just considered. However, it is by no means common and only occa- sionally locally abundant in my experience. In color it is usually dark reddish brown but may be lighter or, according to McAtee (1917a), almost black. Drake (1919a) stressed a close resemblance to L. ilicis Drake, which is distinguished chiefly by its smaller size. L. mutica is 2.9 to 3.0 mm. long and 1.0 to 1.1 mm. wide (Osborn and Drake, 1916a). The species ranges from Quebec to North Dakota and south to Florida and Texas (Blatchley, 1926). Records for New England include the first for Maine (by Little Wilson Stream near Willi- mantic, September 5, 1949) and two previously unreported for New Hampshire (Durham, September 15 and 30 and Lakeside, Septem- ber 9, 1949). Parshley (1922b) noted the species from Lexington, Massachusetts (June 25) and my collection includes specimens from 28 Volume XXXI Sherborn (June 8 and August 24), and Dover (June 16), Massa- chusetts. These scattered records for the northeast indicate that the species is more prevalent than can be demonstrated at present. Egg: Dickerson and Weiss (1916) describe the eggs as follows: “Length 0.36 mm.; greatest width 0.22 mm. The some- what flask-shaped, smooth, whitish eggs were found on the under surface of the leaf, inserted as a rule in the mid-rib, but sometimes in the leaf tissue adjoining the mid-rib. Usually they occurred in small clusters, being stuck sometimes verti- cally in the tissue and at other times at an angle. The necks of the eggs seemed to be bent slightly so as to bring the cap on a level with the leaf surface. Where many eggs were found in a mid-rib, a distortion was present, the rib extending out on one side and being thickened at that point. The tissue sur- rounding the eggs was somewhat hard and corky and each egg- cap was topped by a brownish scab-like crust evidently depos- ited by the parent insect. ’ ’ In this case the host plant was Chionanthus virginicus L. grow- ing in a nursery in Hammonton, New Jersey. Nymphs: In the same paper the authors (Dickerson and Weiss, 1916) describe each of the five nymphal instars. A fine plate figur- ing the egg, nymphal stages, and the adult accompanies the article. The paper is summarized and the plate reproduced in a pamphlet entitled “ The Lace Bugs of New Jersey” (Barber and Weiss, 1922). Both publications are generally available. Dickerson and Weiss (1916) observed many nymphs on the upper side of the leaves where the foliage was dense and shaded. They found that after the second instar the nymphs dispersed somewhat and fed singly or in groups on any shaded portion of a leaf. The plants in the nursery were infested from top to bottom. All nymphal stages were present on July 7 and very abundant on August 15. By September 1 only fifth instar nymphs were found. Adults: Plants in the family Oleaceae are the hosts for this species. My four New England collections of Leptoypha mutica (Say) were all from Fraxinus. In only one place (Sherborn) were they at all numerous. There they were mostly feeding on the upper surface of leaves of suppressed ash seedlings growing under a can- opy of mature hardwoods on a dry, gravelly knoll. The majority were resting parallel with the slightly depressed mid-rib and were very inconspicuous in that position. When the leaves were touched the tingids would commonly let go and drop off, thus avoiding 29 ENTOMOLOGICA AMERICANA capture. Although the species is often reported from Chionanthus where that plant occurs, Dr. Sailer states that specimens from this host show constant differences in repeated collections from Plum- mers Island (near Washington, D. C.), and in material from Clif- ton, Virginia he observed the same tendencies. Hereabouts Chio- nanthus is occasionally grown as an ornamental, but there has been no opportunity for me to make an intensive search for Leptoypha on this host. Drake (1918) reported specimens collected on Adelia acuminata (now Forestiera acuminata (Michx.) Poir.) in Texas and stated that Heidemann got a long series at a light in Maryland. Adults have been found hibernating among old leaves (McAtee, 1917a). Characteristic lace bug feeding damage is described by Dicker- son and Weiss (1916). Practically all leaves of the white fringe bushes in the Hammonton nursery were injured. At first there was a whitish discoloration of the upper surface near the mid-rib. Such light patches enlarge with continued feeding until the entire leaf becomes mottled and in severe infestations the leaves actually turn yellowish brown and eventually wither. On the under side the leaves are spotted with the brownish excrement of the nymphs and adults. If the leaves are exposed to direct sunlight most of the insects remain on the lower surface. Where the leaves are shaded they will be found on the upper side. The New England records reveal a period of activity extending from June 8 to September 30. McAtee (1919b) reports captures from April 11 to October 12 on Plummers Island. Although the various stages have been adequately described, there is as yet no definite information concerning the time necessary for develop- ment. Genus Dictyonota Curtis, 1827 Dictyonota tricornis Schrank var. americana Parshley, 1916b, p. 164. There are about twenty-four species in the genus Dictyonota and the majority of them are Palearctic in distribution. The only American representative is this variety of a European species known to occur in the northeast from Nova Scotia to Mt. Desert Island, Maine. Parshley (1916b) and Hurd (1946) both mention the possibility of its introduction from Europe. However, Parshley (1916b) was able to compare it with European material of D. tri- cornis Schrank and considered it distinct in a varietal sense. This insect is a dull greyish brown to black. The general form 30 Volume XXXI is oval. Antennae are thick, rugose and spinous. The pronotum is tricarinate and the small hood does not project over the head. There are no discal elevations on the hemielytra and the costal area is as wide as the discoidal and has two slightly irregular rows of areoles. The bucculae are open anteriorly and the sternal ridges are but slightly elevated. The rostrum reaches a little be- yond the metasternum. Males have strongly developed claspers. The variety is 3 mm. long and 1.2 mm. wide. Since the North American records for this insect are so few they are given in detail. The holotype is a male from Eastport, Maine (July 15, 1909) in the Parshley collection. There are two paratypes, one from Machias, Maine (July 26, 1906), and the other from Roque Bluffs, Maine (July 15, 1907). Both are in the Mu- seum of Comparative Zoology collection. Parshley (1923a) gives two later records for Nova Scotian localities. One is Kings County (July 4, 1921) and the other is Truro (July 21, 1920). The final record seems to be the one given by Proctor (1946) for Mt. Desert Island, Maine (August 29). It is interesting to note that five of the six specimens known were taken in July. On September 8, 1949, a full forenoon was spent sweeping a wide variety of herbs, shrubs, and other vegetation in Roque Bluffs for this tingid but without success. Since there is no other information about D. tricornis var. americana available, it is worthwhile to quote Butler (1923) con- cerning the habits of the species in Europe. The eggs are said to be unknown, but he describes the last instar nymph and states that adults occur from June to October. Then : £ ‘ Unlike most Tingidina, this species is usually found singly, and occurs, as a rule, by promiscuous sweeping, but I have not found it anywhere common. Curtis found the speci- men upon which he erected the genus under a stone in a meadow near Bognor, Sussex; and Bedwell tells me he fre- quently finds it in the crevices between blocks of chalk, flint, etc., on the Downs in the Box Hill district. As food-plants Horvath gives Mentha, Artemisia, and Thymus, and Carr adds broom ( Cytisus scoparius). Dubois took it in summer by beating pine trees. Douglas and Scott give ‘ among grass and moss in dry sandy places.’ Morley gives sandpits, nests of Myrmica rubra, and at roots of Sedum acre . Donisthorpe has found both imago and larva in nests of A. flavus, and J. Sal- berg gives A. niger 31 ENTOMOLOGICA AMERICANA Genus Acalypta "Westwood, 1840 Orthosteira Fieber, 1844; Orthostira Fieber, 1861; Fen- estrella Osborn and Drake, 1916a; Drakella Bergroth, 1922. Approximately 35 species have been described in the genus Acalypta. Butler (1923) states that 24 Palearctic species are known, and Hurd (1946) notes ten American and one Japanese species. Very little, however, is recorded concerning the biology of any of them. The European representatives have usually been collected in association with mosses on stumps, on the dry borders of woods and hills covered with low grasses and heath, about the roots of certain herbs in sandy places, sometimes apparently living with ants (like Dictyonota) , and even under dry ocean drift (But- ler, 1923). They have been more often found by patient search of such habitats than by sweeping. The description of. one American species is based on a female found in drift on the shore of a reservoir in British Columbia (Downes, 1927). In my collection there are a dozen specimens given to me by K. A. Christiansen. They were obtained from oak leaf duff from Corvallis, Oregon, which he was putting through Berlese funnels for Collembola. Sources of our New England spe- cies will be enumerated below. Although the eggs of all species remain undescribed, Butler (1923) describes the nymphs of three of the six British species and figures one of them in an outline drawing which completely fails to suggest the actual appearance of the immature insect since the numerous simple and stellate dorsal spines are entirely omitted. The few specimens at hand suggest that these processes would fur- nish useful criteria for distinguishing the different nymphal in- stars if only enough material were available for examination and for study of the limits of variation in the form and position of these structures. They might also aid in specific identification of the nymphs. Acalypta lillianis Bueno, 1916, p. 39. Acalypta ovata Osborn and Drake, 1916b, p. 9, Fig. 1. Acalypta grisea Heidemann, 1917, p. 218, PI. 17, Fig. 2. Acalypta modesta Parshley, 1921, p. 4. Acalypta lillianis Bueno is the best known of the three New England species. Members of this genus are small, rather oval, and the hemielytra are only slightly convex. The paranota and 32 Volume XXXI costal margins are explanate and the relatively large areolae of these areas form a rather continuous border. The pronotum is tri- carinate and the hood but slightly produced between the conspicu- ously large and coarsely faceted compound eyes. Areolae are gen- erally rounded and the nervures strongly raised. The color is greyish black. Both brachypterous and macropterous forms are known. Brachypterous individuals are 2.3 mm. long and 1.2 mm. wide, while macropterous specimens are 3.0 mm. long and 1.6 mm. wide. Since this species is widely distributed some variation in size and color is to be expected. Figures of both brachypterous and macropterous individuals are given by Osborn and Drake (1917a), (see Parshley, 1917a for criticism), and Heidemann (1917) figures the brachypterous form. In New England, specimens have been taken at The Forks (Parshley, 1917c), and Orono (Parshley), Maine; Durham (Os- born and Drake, 1916b — as A. ovata 0. and D.), and Franconia ( fide Sailer), New Hampshire; Amherst (University of Massachu- chusetts), Andover ( fide Sailer), Attleboro (Parshley), Canton and Dover (Bailey), and Provincetown ( fide Sailer), Massachu- setts. Acalypta lillianis is widespread and has been reported from Quebec to British Columbia and south to Iowa and Maryland (Hurd, 1946). Nymphs : Bueno (1916) reported nearly mature nymphs found under stones on November 25, December 5, March 5 and 19, and April 2. This makes it evident that nymphs hibernate. Three nymphs in my collection were taken on July 7 and each represents a different instar. Another taken on August 2 in ’the same locality (Canton, Massachusetts) seems to be the final stage. These imma- ture specimens were taken by sweeping the haircap moss, Poly- trichum, where adults had been collected earlier in the season. Drake (1928b) found nymphs and adults in low wet mossy areas on the virgin prairie near Ames, Iowa. Adults: The Andover specimen (fide Sailer) was found under a stone in April, which suggests that adults also hibernate. The period of adult activity is only between June 2 and June 27 ac- cording to the New England records seen. However, Drake (1928a) reports them from May into August for New York, and Bueno (1916) took most of his type series in late May. The little evidence at hand points to a short period of adult availability during late spring. My first success collecting them came on June 10, 1948, when 18 were taken in less than an hour of sweeping Polytrichum. 33 ENTOMOLOGICA AMERICANA On June 17 a few more were collected in the same Canton locality. Since then, however, I have never found more than one or two at a time. During* the spring, summer, and autumn of 1949 and 1950, numerous attempts in the same patches of moss were fruitless. There is something about the habits of this species yet to be ex- plained. Since European species of Acalypta have been found associated with plant roots and in ant nests (Butler, 1923), it may be that A. lillianis spends most of its time beneath the surface of the soil and possibly only for a few days early in the season are adults at all common above ground, when they may emerge for mating or some other special purpose. Or, more simply, the spe- cies may be nocturnal in habit and retire under stones by day. There is, nevertheless, something unusual about the habits of spe- cies of Acalypta which may account for the scattered records and the varied conditions under which they have been found. Other habitat situations noted for this species include damp marshy meadows where the tall tree-like moss ( Climacium ameri- canum L. — according to Blatchley, 1926) grows in clumps (Bueno, 1916), under mosses on rocks in May (Parshley, 1921) ; and H. 0. Barber collected a long series under a board (Drake, 1928b). Drake (1928b) also stated that Henderson took hibernating adults in Iowa in early April. In the type locality Bueno (1916) col- lected them in small numbers from mid-May until the end of the month and records a specimen taken in Michigan on July 14. An abortive attempt was made to keep some specimens alive from a series of about eighteen adults and a few nymphs taken in Canton on June 10, 1948. Several adults and the nymphs were placed with fresh Polytrichum in a plastic vial and taken home. A sod of the moss was also taken and placed on clean sand in an aquarium. This was watered well and a day later the lace bugs were placed on the moss. For the next day or two an occasional in- dividual could be found apparently feeding on the fresh moss cap- sules from which the calyptra had fallen. This was also very defi- nitely observed while the insects were being transported on the moss in the plastic vial. Later a nymph was seen in the process of molting to the adult stage. Due to unsatisfactory humidity or other conditions it was unable to successfully extricate itself, how- ever. In a few days the insects disappeared entirely. If more specimens can sometime be obtained another effort will be made to rear them under conditions more nearly simulating the out-of-doors. The house in summer was probably too hot and dry. 34 Volume XXXI Acalypta nyctalis Drake, 1928b, p. 5. All that is known concerning this species is given in the paper cited above. It is described as ovate and yellowish brown. Al- though it resembles A. lillianis Bueno, this species may be distin- guished by its much narrower paranota and by differences in the hood, carinae, and hemielytra. Two brachypterous females only are recorded. The holotype came from Franconia, New Hamp- shire, and is in the United States National Museum. The paratype was collected on June 1, 1924, in Alberta, Canada. They are 2.6 mm. long and 1.2 mm. wide. Acalypta thomsonii Stal, 1873, p. 122. Acalypta madelinae Bueno, 1926, p. 117. Although this is the first American species described, it is rarely collected. It may be immediately separated from the two species previously mentioned by the low lateral carinae which diverge strongly behind. A. thomsonii is smooth, rather shiny and brownish to dark greyish brown. Only brachypterous individuals are known. Osborn and Drake (1916a) give the length as 2.5 to 3.0 mm. and the width as 1.5 to 1.75 mm. Bueno (1926) cites three specimens sifted from leaves around the bases of alder clumps in a swamp by Mr. C. A. Frost in Sher- born and Framingham, Massachusetts. In the United States Na- tional Museum there is a specimen from Rhode Island ( fide Sailer). Elsewhere the species occurs in Maryland ( fide Sailer), South Caro- lina, Virginia, and the District of Columbia (Drake, 1928b). Mr. Frost made his sifting collections in October, which means the insects were probably hibernating. A Virginia record for Sep- tember 28 and a January 23 record for Maryland are both from Sphagnum moss ( fide Sailer). Blatchley (1926) says they occur in late autumn beneath weed debris along the margins of swales. And Drake (1928b) states that adults hibernate in moss. Genus Leptopharsa Stal, 1873 Leptostyla Stal, 1873; Gelchossa, Kirkaldy, 1904. More than sixty species of this large and variable genus occur in South America and thirty-three in North America. Few are found elsewhere. Only three of the North American species have been reported from New England. They feed mostly on leguminous plants, but there are some interesting exceptions noted in the lit- erature. Their elongate sub-rectangular form makes them readily 35 ENTOMOLOGICA AMERICANA distinguishable from other local genera. There is a small areolate hood at the anterior end of the median carina which covers only the occiput. Lateral carinae are present and, like the median, have a single row of areolae. The membranous paranota are flar- ing erect. The hemielytra are constricted near the end of the ab- domen and extend well beyond the terminalia. Areolae of the costal area are conspicuously large and the discoidal area is some- what depressed. Leptopharsa clitoriae (Heidemann), 1911b. Leptostyla clitoriae Heidemann, 1911b, p. 180, Fig. 4. Leptostyla costofasciata Drake, 1916, p. 326, Fig. 1. This species is readily separated from the other two by its rela- tively broader and shorter form. Of the three in New England, L. clitoriae alone has the costal area completely biseriate. It is also most distinctively colored and the nearly black pattern con- trasts sharply with the discrete whitish areas of the paranota and the costal borders. The figure in Drake’s paper (1916) is a good representation of the species. It is about 2.2 to 2.4 mm. long and about 1.1 to 1.2 mm. wide. McAtee (1919b) reports the species from Massachusetts, but I can find no more definite indication of its occurrence there. Proc- tor (1946) lists it for Mt. Desert Island, Maine. The only New England specimens seen are about a dozen in the New Haven Ex- periment Station collection, all taken by Mr. J. P. Johnson in New Haven on September 24, 1942. This Connecticut material is the best evidence of its presence in New England. Outside of this region the range extends to Indiana, Arkansas, and South Carolina (McAtee, 1919b). McAtee (1917a) reports the insect from Clitoria mariana L. and various species of Lespedeza, and Meibomia (now Desmodium). Heidemann (1911b) said his type specimens were found close to the ground on Clitoria. Blatchley (1926) notes that it may be found on such legumes growing along high wooded slopes. Another host is said to be Lappula (Boraginaceae) (fide Sailer). In Proc- tor (1946) the notation “On alder, which is its food plant” raises an interesting question. The same source gives July- August as the time when specimens were obtained. In the District of Colum- bia dates of collection range from June 26 to October 14 (McAtee, 1923). McAtee (ibid.) notes eggs on July 19 and saw the species in copula on July 26. He also states that they are attracted to light. 36 Volume XXXI Eggs: Heidemann (1911a, PL 10, Fig. 6) described the eggs of L. clitoriae just before his description of the adult insect was pub- lished. The eggs are said to be about .5 mm. long. They are laid singly and with the abopercular end attached to the lower side of the leaf. The egg stands upright, is ovate, and tapers noticeably toward the point of attachment. It is black except for the aboper- cular end, which is whitish. The chorion is thick and hard. It is covered with numerous coarse granules. There is a rim at the opercular end with channeled chorial processes running vertically around the extension of this collar-like structure on its inner side and continuing onto the low, broadly conical operculum to its center. The processes may be noticed on the outside since they budge somewhat. The ripples thus formed are usually covered with a whitish substance. There seems to be no literature on the nymphs nor on the other biological relationships of this species. Leptopkarsa heidemanni (Osborn and Drake), 1916a. Leptostyla heidemanni Osborn and Drake, 1916a, p. 238. Van Duzee (1917) erroneously synonymized this species with L. clitoriae (Heidemann), which is very distinctive. L. heide- manni (0. and D.) is, however, very similar to L. oblonga (Say). It is slightly larger than L. oblonga and appears to be somewhat broader in proportion to its length. The head spines of L. heide- manni and the transverse costal nervures are dark, while the same parts of L. oblonga are distinctly white or colorless. In the costal region of L. heidemanni there may be a few extra areolae which make the margin irregularly uniseriate throughout, but the costal area of L. oblonga is very regularly uniseriate from the base of the wing to just beyond the apex of the discoidal area. Specimens be- fore me show the hood of L. heidemanni to be somewhat shorter and broader, with only three areolae along the median ridge where L. oblonga has four. Seen together they are readily separated. L. heidemanni is 3.0 mm. long and 1.0 mm. wide. This is a wide ranging species and has been commonly collected in Massachusetts, Connecticut, and elsewhere in New England. Records for the two states named are so numerous it seems unneces- sary to list them (see Parshley 1917c and 1923 for some published records) . Parshley (1920b) also reports the species from Woodford, Vermont, and there is a specimen in the Rhode Island State Col- lege collection from Kingston. There are apparently no records from Maine nor from New Hampshire. The species is known from 37 ENTOMOLOGICA AMERICANA New York to Ohio (Hurd, 1946), from Arkansas and Louisiana (Blatchley, 1926) and from Missouri (Froeschner, 1944). With one exception the food plant is always given as Baptisia tinctoria (L.) R. Br. My collections locally have always been from that herbaceous host which commonly shows severe feeding damage. The Woodford, Vermont, material mentioned by Parshley (1920b) was taken on alder. This should be compared with the Mt. Desert record for L. clitoriae (Heidemann) (Proctor, 1946). McAtee (1923) records the seasonal activity as extending from May 2 to October 10 in the District of Columbia. New England records run from May 29 (Natick, Massachusetts — fide Sailer) to September 16 (East Hartford, Connecticut — Parshley, 1923). During June and July I have found nymphs and adults feed- ing on the under side of Baptisia leaves. The injured foliage is conspicuously whitened on the upper surface. The leguminous host grows commonly on dry soils and often along roadsides or woodland openings. Favored situations are often warm and dry in good weather, but there are usually trees and shrubs nearby which offer some shade during the day. Weiss and West (1924) give a good account of the habits and early stages of L. heidemanni (0. and D.). Their paper is readily available, and, therefore, their notes on the biology of the species are quoted in full, but the detailed descriptions of the egg and nymphal instars are omitted. “In the central portion of New Jersey overwintering adults appear about the last week of May and persist in more or less plentiful numbers until about the middle of June. As a rule the adults inhabit the lower surfaces of the leaves and do noticeable feeding, causing white areas to appear on the upper surfaces. Eggs are deposited during the last of May and first part of June and the young nymphs become plentiful about the middle of June. By the last of June and first week of July many last stage nymphs are in evidence and adults issue shortly afterwards, becoming numerous from the begin- ning of the second week of July until the end of the month. There is apparently only one brood and the adults gradually disappear during August. The eggs are deposited singly, or in irregular groups of two or three, in the lower tissue of the leaf either near or away from the midrib. Each egg is em- bedded well in the tissue with the long axis of the egg parallel with the leaf surface and with only the truncated, oval end of 38 Volume XXXI the neck of the egg visible as it projects slightly beyond or re- mains flush with the leaf surface. These oval ends resemble stomata somewhat and are similar in color to the leaf. After hatching the young nymphs appear to feed close to the midrib, assuming a position as a rule, parallel to this part of the leaf. As they become older they feed in colonies of 10 or 12 on the lower leaf surface, although many leaves may contain more and some only one or two nymphs. Usually both adult and nymphal feeding is well scattered over the plant. In severe infestations, which often occur, every leaf is white and hun- dreds of whitish nymphs and adults inhabit each plant. Such a condition existed at Prospertown, N. J., on July 18, when hundreds of adults and a very few last stage nymphs were ob- served on the whitened plants. Five nymphal stages were ob- served. . . Leptopharsa oblonga (Say), 1825. Tingis oblonga Say, 1825, p. 325. Leptopharsa oblonga (Say) closely resembles L. heidemanni (0. and D.) except in the features mentioned under that species. It is included on the basis of specimens in the Museum of Com- parative Zoology collection taken by Blanchard in Tyngsboro, Mas- sachusetts, on March 17, 1895. This is the only record for the spe- cies in New England as far as I can discover. That early in the year they must have been in hibernation. Hurd (1946) gives the range as New Jersey to South Dakota to Arkansas to Virginia and Brazil. The species is about 2.8 mm. long and about .9 mm. wide. Dates of collection for L. oblonga (Say) in the District of Co- lumbia run from June 5 to August 22 (McAtee, 1923). Most food plant records are Falcata comosa (L.) (now Amphicarpa bracteata (L.)). Blatchley (1923) also gives Amorpha fruticosa L. and Kuhnistera (now Petalo sternum) as hosts and states the insects were swept from herbage in dense upland woods. McAtee (1917a) re- ports specimens from a leaf of Tilia in Wisconsin and from Petalo- stemum growing on dry gravelly knolls in Iowa. Genus Galeatus Curtis, 1833 Galeatus peckhami (Ashmead), 1887. Sphaerocysta peckhami Ashmead, 1887, p. 156. Galeatus peckhami (Ashmead) is one of the most unusual members of the New England tingid fauna. The paranota and the 39 ENTOMOLOGICA AMERICANA costal area of the hemielytra have a single row of large subquadran- gnlar areolae. The dorsal membranous parts are all hyaline except for some infnscation of the much inflated lateral carinae which rise above the median carina and the tumid apex of the pronotum. All nervures are yellowish brown. The hood is very small and does not cover the head, which bears five remarkably long, sharp, slender and erect spines. The body is reddish brown, while legs and an- tennae are yellowish. This species is 4 to 4.5 mm. long. Parshley (1917c) gives Princeton, Maine (July 12, 1909), as one New England locality for Galeatus, while Proctor (1946) re- ports the species from Salisbury Cove on Mt. Desert Island in the same state (July 15), and there was a nymph from Bar Harbor, also on Mt. Desert, in the Museum of the Boston Society of Natural History. New Hampshire records include the Glen House (July 20, 1915) (Parshley 1917c) and Fabyans, also in the collection of the Boston Society of Natural History. The type locality is in Wisconsin and the species ranges from Ontario to Manitoba in the north (Blatchley, 1926). Kecords from Michigan, Minnesota, and New York (Drake 1922c) are be- fore me. Uhler (1896) recognized this species in material from Japan. However, his (Uhler, 1904) report of the species from New Mexico is not valid, since Horvath (1923) has described a new species based on the Las Yegas specimen. Nymphs: Osborn and Drake (Drake, 1922c) found the species numerous and breeding on Aster macrophyllus L. and Eupatorium on the summit of Crataegus Hill, Barber Point, Cranberry Lake, New York, during the last week of July, 1920. At the time only adults were present, but the exuviae of four different nymphal in- stars were found clinging to the lower surface of aster leaves. The three later stage exuviae were nearly perfect and these are fig- ured (PI. 4, Figs, f, i, j) and described. The number of nymphal instars is probably five. Eggs: Drake (1922c) says that: “The eggs of Galeatus peckhami (Ashmead) are almost en- tirely inserted in the stem of the host plant, usually somewhat near the surface of the ground, upon which the insects are feed- ing. They are generally placed singly (Plate 4, Fig. e) and in no definite order in the stems of the plants. Only a small portion of the egg or the neck-like structure and cap protrudes from the plant. However, occasionally two or rarely three eggs are inserted in the same slit or egg puncture. Sometimes five 40 Volume XXXI or six or even a dozen may be laid in rather close proximity to each other. They are deposited during the latter part of July, August, and September. In proportion to the size of the abdomen the eggs are quite large and only a few fully formed ova can be contained within the body of the female at the same time. This probably accounts for the long period of egg-de- position. There is but a single generation a year in the Adi- rondacks and field observation indicates conclusively that eggs, which are laid in the latter part of the summer do not hatch until the following spring. Asters, in which eggs had been deposited in the stems in the field, were transferred to small pots and placed in the laboratory at Syracuse. Adult males and females were also carried to Syracuse on the host plants, but all died during the latter part of September and October. The plants were destroyed by mildew during November and De- cember. The eggs, which had been deposited during July or later, failed to incubate in the laboratory, but seemed to have remained in a living state until they were destroyed by desicca- tion of the stems of the asters a few weeks after the plants had been killed by the mildew. “The egg (Plate 4, Pig. e) is slightly curved, from .7 to .8 mm. long and about one-third as wide. The cephalic end is distinctly neck-shaped and closed by a round cap or lid. The color varies from brown to dark brown or black, usually con- siderably darker on the cephalic half and with a much lighter cap.” Adults: The genus Galeatus is predominantly Palearctic in distribution and in some species the hemielytral length is variable. Only macropterous individuals of G. peckhami are known, however. According to Drake (1928a) the species is locally distributed, but may be found in large numbers on knolls in semi-shady, dry situa- tions. Pour predaceous Heteroptera were found associated with Galeatus in the Cranberry Lake region. Whether or not the adults overwinter is unknown. Van Duzee (1889) recorded this insect as swept from low weeds, probably a dwarf vaccinium or a species of aralia, which were growing together among pines on a rocky island in Canada. Bueno (1915) noted thirty specimens found in beach drift on the shore of Lake Superior in Michigan, which suggest a migratory movement over the lake. A good figure (PL 4, Fig. a) of the adult accompanies the paper by Drake (1922c). 41 ENTOMOLOGICA AMERICANA Genus Gargaphia Stal, 1873 This genus may be readily separated from all others by the transverse carina that interrupts the rostral channel between the meso- and metasternum. In addition, it differs from the related genera Corythaica and Corythucha by having a much smaller hood that never extends beyond the anterior margin of the eyes, and from Stephanitis by its low median carina and flattened discoidal area. Fifty-five species, all from the Western Hemisphere, compose this genus. Twenty-six of these are from North America, and of these, only three are known from New England (Hurd, 1946). Gargaphia angulata Heidemann, 1899, p. 301. Gargaphia angulata Heidemann is the smallest of the three species occurring in New England. In this species the free rim of the paranota is about level with the crests of the pronotal carinae. In the other two the paranotal margin is noticeably higher. Hood, carinae, and paranota bear an abundance of long, silky setae. Except for the body, the dark disc of the pronotum, and the fuscous fourth antennites and tarsi, the color is dull stramineous. The insect is 3.4 mm. long and 1.6 mm. wide. New England records are not numerous, and only two of the states are represented. Massachusetts records are from Framing- ham, June 15, August 30, and September 1, Mr. C. A. Frost, col- lector; Natick, June 4, fide H. G. Barber; and Westfield, July 10, (Parshley, 1917c). Connecticut records include Brookfield, July 27, (Parshley, 1917c) ; Hartford, June 15, New Haven, June 23, July 23, and August 19, New Haven Experiment Station collection. Hurd (1946) gives the full range for this species as Massachusetts to Colorado, Alabama to Minnesota. Mr. C. A. Frost made his collections on Ceanothus americanus L. (Rhamnaceae), which, from several other reports ( e.g McAtee, 1917b), seems to be the preferred host. The species first came to scientific notice, however, as a pest of beans (Chittenden, 1900). The paper just referred to has a good figure of G. angulata Heide- mann. The same drawing was used later by Blatchley (1926). The season of activity in New England runs from about June 4 ( fide Barber) to September 1 .(fide Sailer). Although Mr. Frost has collected them in the vicinity of Framingham, Massachusetts, for several years, I have been unsuccessful in searching the same host in the Boston park system and in Dover. The distribution appears to be very local and there is no other biological data on record. 42 Volume XXXI Gargaphia solani Heidemann, 1914, p. 136, figure 1. The preference of G. solani Heidemann for solanaceous plants has given this tingid considerable economic importance. This is especially true in parts of its range where eggplants are grown commercially. The species is readily distinguished from G. angu- lata Heidemann by its larger size and from both G. angulata Heide- mann and G. tiliae (Walsh) by its high-angled hood which is lat- erally compressed, by the lobes at the middle of the paranota which are wide and flaring-erect, and by the infuscated first, second, and fourth antennites, nervures of the paranotal lobes, and many ner- vures of the hemielytra. Hood, paranota, and pronotal carinae are conspicuously covered with long, soft hairs. They are about 4 mm. long and 2 mm. wide. The only New England record discovered is for Milford, Con- necticut (September 21, 1944), on eggplant ( fide Sailer). The species is primarily southern and southwestern in distribution. Its known range includes Virginia (Fink, 1915), Maryland, Ohio, Arizona, Texas, and Missouri (Blatchley, 1926). Hurd (1946) also lists Canada. Fink (1915) contributed importantly to our knowledge of the habits and life cycle of Gargaphia solani Heidemann. With the exception of a few host plant records and one or two other items, the following biological information is derived from his paper. Eggs: The eggs are about .37 mm. long and .18 mm. wide. They are light to dark green where attached, but become brownish towards the opercular end. There is a whitish, lacy border around the cap and a screw-like rim. The operculum is said to be crater- like. The eggs are attached to the under side of the food plant leaves near their abopercular ends and usually lean in all direc- tions and at many angles. They are laid in roughly circular masses of about 116 to 188 in all, over a period of five to eight days. A single female will oviposit on four or five days, laying from 15 to 57 eggs on each occasion, to complete the egg mass. After oviposition a sticky secretion is spread over the eggs. The female attends the eggs during the entire incubation period and leaves them only at intervals to feed. The incubation period for the eggs in late May, June, and July at Norfolk, Virginia, ranged from five to eight days. Nymphs: The five nymphal instars are fully described by Fink (1915) and the final stage shown in his PI. 1, Fig. 1. Maternal solicitude is an interesting phenomenon in the genus Gargaphia. The attendance of the female on the eggs is not peculiar to this 43 ENTOMOLOGICA AMERICANA species nor is her subsequent care for the young unique. Bueno (1942) made similar observations on G. iridescens Champion in Arizona and noted that G. tiliae (Walsh) has been frequently seen watching over its nymphs. This I can confirm from local observa- tions of the species last named. Fink (1915) points out that the nymphs always feed in groups. After the first molt they become yellowish and also move to new feeding positions. When they move from one leaf to another the female usually directs them and keeps the brood together with her antennae. With a colony of over one hundred to care for, the female moves hurriedly from one end of the migrating brood to the other in an effort to keep them together and to urge them on in the proper direction. Once the movement is satisfactorily completed, the nymphs settle down in the new area and begin to feed. Not only does the female guide the activities of her nymphs, she also attempts to protect them from predators. On one occasion Fink (1915) saw a ladybird beetle ( Hippodamia convergens Guer.) approach a feeding colony of nymphs. The adult in attendance suddenly darted toward the intruder with wings outstretched and somewhat raised. She actually succeeded in driving the beetle from the leaf. During their growth the nymphs molt five times. Fink (1915) followed the development of nine individuals and found that two days was the usual interval between molts. Since the eggs require an incubation period of 5 to 8 days, and the nymphs take another 10 days to mature, he estimates that the life-cycle is usually com- pleted in about 20 days. This makes some allowance for the time necessary for mating and oviposition. Adults: Fink observed that females laid their first eggs as soon as two days after copulation. In the Norfolk, Virginia, area they have a breeding season of nearly six months, extending from late May into early November. There is a possibility of seven to eight generations each year in that area. In the field these generations overlap so that all stages may be found practically throughout the season. The insects feed naturally on such solanaceous plants as Solanum carolinense L. and S. elaeagnifolium Cav. (Heidemann, 1914). S. Melongena L. (Fink, 1915), S. tuberosum L., and Ly co- per sicum esculentum Mill. (Somes, 1916) are crop plants which attract this lace bug. Damage to eggplant is particularly severe and typical of tingid injury to foliage. This insect has been col- lected from such diverse hosts as Amphiachrus (possibly Amphi- 44 Volume XXXI achyris Nutt, which is now Gutierrezia Lag., Compositae ) , Cassia species, Legnminosae, Gossypium herbaceum L., Malvaceae, and Salvia Pitcheri Torr. (now Salvia azurea Lam., Labiatae) (Gibson, 1919b). The adults hibernate and emerge early to feed on S. carolinense L. Whereas six broods develop on the eggplant, probably seven or eight a season are produced on the native Solanum (Fink, 1915). Froeschner (1944) reports adults all year in Missouri, where they usually are found hibernating in grass clumps during the winter months, but may also be collected under bark or the basal leaf rosettes of mullein ( Verbascum ). Fink’s (1915) discussion of the. predators of G. solani is of particular importance since the natural enemies of tingids are but rarely mentioned. He observed both larval and adult ladybird beetles of two species (Hippodamia convergens Guer. and Megilla maculata De Geer) feeding on nymphs and adults. Usually the beetles turned the lace bugs on their backs before feeding on them. The soldier-bug, Podisus maculiventris (Say), feeds on the nymphs, as does another common hemipteron, Triphleps insidiosus (Say) (now Orius insidiosus (Say)). In addition to these insects, three species of spiders feed on all stages. They are Epeira domiciliorum Hentz, Plectana stellata Hentz, and Chiracanthium inclusum Hentz. Many lace bugs were found decapitated and their bodies mutilated. A few specimens of a parasitic hymenopteron ( Microdus sp.) were reared with the tingids, but it was not proved that they parasitized them. Artificial control on egg-plants can be effected with nicotine sulfate and fish oil soap sprays thoroughly applied to the under side of the leaves. Adults are not killed by the nicotine sulfate alone, but seven pounds of the fish oil soap in fifty gallons of water will kill all nymphs and 90 to 95% of the adults. Gargaphia tiliae (Walsh), 1864. Tingis tiliae Walsh, 1864, p. 408. In addition to the differences already mentioned, G. tiliae may be distinguished from the two other species by its generally pale whitish color. Usually only a few nervures just beyond the middle of the discoidal area are infuscated enough to produce a spot on each hemielytron which extends onto the adjacent costal area. The pubescence of G. tiliae is less pronounced, which results in a neater appearance. There is, however, a fine, dense, and somewhat matted pubescence on the disc of the pronotum. 45 ENTOMOLOGICA AMERICANA Rhode Island is the only New England state for which there seem to be no records. There are no published records for Maine and Vermont nor for New Brunswick and Quebec in eastern Can- ada. Specimens collected in those places, or to be found in other collections, will be noted. Four Maine localities can be reported. They are Paris ( fide Sailer), Willimantic, Lambert Lake, and Al- bion (Bailey). Previously unpublished (see Parshley 1917c) New Hampshire records include Antrim ( fide Sailer), Durham (Uni- versity of New Hampshire) and Harrisville (Bailey). The only specimens seen from Vermont are some in my collection from Sun- derland. Massachusetts stations not listed before are Montgomery ( fide Sailer), Amherst (University of Massachusetts), Boston and Swansea (Boston Society of Natural History), Mt. Holyoke (Cor- nell University), Arnold Arboretum, Boxford, and Georgetown (Bailey). There is material in the New Haven Experiment Sta- tion collection from Canaan and from Salisbury. Although Pro- vancher (1886) suggested that the species occurred in Quebec, no definite records were found. There are, however, specimens from that province in Professor Parshley ’s collection and the species was taken in Applegrove on September 1, 1949. There are also specimens in my collection from Richmond Center, New Bruns- wick. Beyond the limits of New England, G. tiliae ranges from New York to Ontario, Nebraska, Arizona, Missouri, Alabama, and Virginia (Blatchley, 1926 and Hurd, 1946). Some notes on the life history and immature stages are given by Weiss (1919). The interest of the females in their eggs and young has already been discussed (see G. solani). Since there is nothing further to add, it need not be repeated. The habits of this species evidently follow the same pattern. Eggs: The eggs are .48 mm. long and .18 mm. in diameter. They are sub-oval, since one side is more convex than the other. The basal portion is somewhat acute, but rounded at the tip and slightly constricted where it is inserted into the leaf. The opercular end is truncate and bears a rim-like collar. The operculum is conical and has a slight nipple-like projection in the center. Ex- cept that the upper third of the egg may be covered with the usual varnish-like secretion, the eggs are translucent. In Pennsylvania eggs are laid in late May and early June. They are deposited in groups of less than 60 to as many as 300 on the lower leaf surface. Most project at right angles to the leaf, but they may lean in any direction. The surface of the leaf above the egg mass usually be- 46 Volume XXXI comes brown as a result of injury to the tissues. Incubation of the eggs takes about a week. Weiss (1919) observed two annual broods, with the second egg-laying period coming before mid- July. Nymphs: Detailed descriptions of the five nymphal instars are given. Development of the nymphs is said to require about three weeks. The nymphs feed in compact groups with the female par- ent usually at hand. They are brownish white and, on the leaves somewhat discolored by their feeding injury, are rather inconspicu- ous. The upper surface of the damaged leaves becomes first mot- tled with whitened areas. These later turn brown as the tissue is killed and dries out. Adults: The adults emerge from hibernation in late May or early June, feed on the linden foliage, and soon start to lay eggs. Early in the season the adults are scattered and the injury to the foliage is not very noticeable. About a month after the eggs are laid the first summer brood of adults appear. They continue the cycle by laying eggs before mid- July. These eggs produce the second brood adults by mid- August or early September. It is the adults of this second brood that hibernate to emerge the following spring. All of the food plant records for this species are Tilia, except that Gibson (1919b) mentions wild cherry. G. tiliae has been so consistently associated with linden in most cases that the cherry record may be simply accidental. All of my collections and obser- vations in the northeast have confirmed this host association. The approximate period of activity in eastern Massachusetts (Boxford) is indicated by collections at hand made on June 4, 1948, and Sep- tember 3, 1948. The species is widespread and more prevalent than records suggest. It is very likely to be found wherever lindens will grow. Genus Corythaica Stal, 1873 This genus is represented only in the Western Hemisphere. There are eight South American species, one from the Galapagos Islands, two in Central America, two in the West Indies, and four in North America (Hurd, 1945). Only one of these occurs in New England. Members of this genus have a prominent pronotal hood extending over the head beyond the proximal end of the third an- tennite. This one feature distinguishes Corythaica from all north- eastern tingid genera except Stephanitis and Corythucha. From both of these genera Corythaica is separated by the narrow para- nota and costal area, by the distal tapering of the hemielytra, by 47 ENTOMOLOGICA AMERICANA the uniformly biseriate hypocostal ridge, and by the small size of our native species. Corythaica bellula Bueno, 1917, p. 19. Corythaica floridana Blatchley, 1926, p. 471. The occurrence of C. bellula Bueno in the northeast is of con- siderable interest since the other members of the genus are largely restricted to milder climatic zones. Up until about 1947 the spe- cies was known only from the type locality in White Plains, New York, and from Florida (C. floridana Blatchley — see Drake, 1930). In White Plains, Bueno and Mr. C. E. Olsen (Bueno, 1917) swept them in some numbers from fine low grasses and moss on a meadow sloping up from a rich marshy swale. They were taken from April to September. Our next information concerning this lace bug came from Miss Hilda Vilkomerson who was working on the cyto-genetics of Pani- cum Lindheimeri at Brown University. She found some insects as- sociated with the Panicum collected in Pawtucket, Rhode Island, and causing enough damage to interfere with seed production. Some of these eventually were sent to Professor C. J. Drake and he identified Corythaica bellula Bueno. In response to a letter con- cerning New England Tingidae, Professor Drake informed me of this and suggested that the species should be found in Massachu- setts. Acting on this suggestion, an effort was made to find Cory- thaica locally and the species was soon discovered to be rather gen- eral in distribution and, in some places, fairly abundant. My first success came in Canton on July 26, 1948. A few days later a pair was taken in West Newbury. About a week later a nymph and an adult were swept from Panicum growing on a wood road in Dover. During the remainder of the season seven more collections were made at the Canton locality, yielding a total of over 30 specimens. In all cases Panicum was the chief element in the low herbs swept. The last specimen of the 1948 season was actually taken by shaking a tuft of Panicum over the palm of my hand. The next season the first two adults were taken in Canton on June 1. Nymphs were found there on June 10 and other nymphs and adults were subsequently collected. Another locality was added on July 7 when teneral adults were swept from Panicum on the shore of a pond in Milton. However, of greatest interest during the 1949 season was a successful attempt to rear Corythaica. Three clumps of Panicum 48 Volume XXXI were brought into the greenhouse from the Canton station and potted up after weeding out other grasses. This was done on June 13. Plants A and B were fairly large, showed signs of previous feeding injury (a mottled whitening of blades), and had nymphs present when potted. When examined on June 20, there were still nymphs on Plants A and B and four adults were also seen. On June 22 six adults were counted on Plant A and five on B. A pair were observed in copulation on this date and another pair on June 27. While mat- ing the male is nearly at right angles to the female. His wing tips are above hers, and the tip of his abdomen is beneath her wing tips. As the male retracted his aedaegus after separating from the female, he appeared to push it first with the tip of one clasper and then with the other. The female of this pair was still somewhat teneral. Plant C was small and showed no signs of injury. It appeared to be quite free from infestation. On Plant C two pairs of Cory- thaica bellula were placed on June 21, and the pair seen copulating on June 27 was added on that date. A large lamp chimney with fine copper screening at the upper end was put over the plant. On July 29 the plant was thinned out considerably and in the process nymphs in various stages were seen and eggs located in the grass blades. Only two adults could be found. On August 4 Plant C was again examined and eight adult males were collected. Five of the eight were very teneral and the other three probably somewhat so. Many nymphs were still present. These results prove conclu- clusively that Corythaica bellula Bueno can complete its develop- ment on Panicum and, with the other evidence noted, indicate that the grass is the natural host of this species. Since the period from June 21, when pairs of adults were first placed on Plant C, to August 4, when teneral adults were first observed, is 44 days, the life cycle of the species is completed in six weeks at the most and probably in a somewhat shorter time. My collections to date tend to show that Corythaica bellula Bueno may be found almost anywhere that the Panicum will grow. Bueno ’s note on the type locality was given above and the wood road station in Dover, Massachusetts, mentioned. The only things the situations I have collected them in seem to have in common are the presence of the Panicum and the possibility of some shade. Since there is considerable variation in the amount of shade avail- able, the Panicum is actually the only common factor. However, it may be of some value to describe the range of habitats. In the 49 ENTOMOLOGICA AMERICANA Canton station the Panicum grows in small openings between such trees as Prunus serotina Ehrht., Betula populifolia Marsh., Acer rubrum L., young Pinus Strobus L., Pinus rigida Mill., Juniperus virginiana L., Comptonia peregrina (L.) Coult., Vaccinium and Quercus species, and other vegetation. Often the Panicum grows from a lichen mat around the edges of these areas and with other sparse grasses (e.g., Danthonia spicata (L.) Beauv.) along the bridle trails. In mid-summer the station, though low, is often rather dry and hot. Some of the most heavily infested plants were in the full shade of young pines whose lower branches are only about 3 to 4 feet from the ground. West Newbury specimens came from grasses along the edges of a ploughed field with a dense shrub border and trees just beyond. Other new records include Errol, New Hampshire (September 9, 1949), where they were picked by hand from small clumps of Panicum on the gravelly shore of a small pond surrounded by shrubs on the bank; Dummer Hill, New Hampshire (September 10, 1949), where eight adults (some ten- eral) were shaken from Panicum growing in low dense herbs at the edge of a large field; Eaton Center, New Hampshire (September 11, 1949) , where about fourteen adults were taken from Panicum grow- ing in the gravel between the road pavement and a pond. Other New Hampshire records are for Harrisville (August 14 and 15, 1950) , Nymphs and adults were found on small Panicum plants growing in the crevices of granite ledges with lichens on a wooded hillside. On August 5, 1950, a Connecticut record was added when nymphs and adults were collected in East Hartland on Panicum, again growing from granite crevices and along paths where the soil nearby was moist enough to support a lush growth of Vaccinium corymbosum L., Acer rubrum L., some Kalmia latifolia L., and other vegetation. Strangely enough, there was a ploughed field just beside the area covered with shrubs and trees which was car- peted with a rank mat of what appeared to be the same Panicum . In the freshly turned soil the growth was much more luxuriant. Sweeping those plants in the open field gave no specimens, however. My earliest record is May 14, 1950, for West Newbury and the latest is a Canton record for September 26, 1948. Although col- lecting was started on April 25, 1950, in Canton, no specimens were taken there until June 16, 1950. Nymphs have been taken from June 10 until August 15 in central New England. It is probable that the adults hibernate, but they have yet to be collected in winter. Eggs : The eggs are .54 mm. long and .13 mm. wide. They are colorless, smooth, and sub-cylindrical. There is a slight curvature 50 Volume XXXI on the long axis and the abopercnlar end is bluntly rounded. The female inserts them between and parallel to the longitudinal veins of the Panicum blade from either the lower or upper side. Only the operculum protrudes beyond the epidermis of the leaf. A con- striction and ring-like thickening mark the area of opercular at- tachment at the leaf surface. The opercular end is bent somewhat to bring it above the leaf surface. The cap itself bulges slightly at the base above the point of constriction and then tapers from its base to a low, somewhat striated, and rounded peak. The striated appearance is due to the alveolar nature of the opercular tissue. The eggs are apparently laid singly and in no particular relation to one another. One or several may be found in a grass blade. Nymphs: The characteristic form of a late nymphal instar is well shown in Fig. 2 (drawn by Mr. F. Y. Cheng). The nymphs are broad in proportion to their length and rather flat dorso-ven- trally. The dorsal surface of living nymphs has a greyish lavender tone and a shagreened texture. This apparent roughness results from the numerous short white stellate hairs scattered generously over the surface beneath the larger, more conspicuous, branching spines which are arranged in a rather definite pattern. The earliest instars are paler. Cotton (1917) gives a good account of Cory- thaica monacha Stal, a pest of egg-plant in Porto Rico. For this species he describes five nymphal instars, and his data show that the cycle from egg to adult may require only 15 to 17 days. He also notes two species of coccinellid beetles and two reduviid bugs as predators. Adults: Corythaica loellula Bueno differs so much from all other New England tingids that it is easily recognized. The hood is low, flat, and broadly rounded behind, and tapers to a sharp point for- ward where the apex curves down over the front of the head. The pronotal carinae are all membranous and uniseriate. Paranota are narrow and erect. The costal area has a single row of rather large areolae and the hypocostal ridge has two rows of small areolae. The median nervure of the hood is carinate and nearly black, as are most of the hemielytral nervures. All areolae are greyish white and opaque. Hood, carinae, paranota, and costal area are color- less, giving this minute insect a striking appearance. Both brachyp- terous and macropterous forms are known, but the latter are un- common and I have not yet seen a macropterous specimen from New England. Hurd (1945) gives a figure (PI. 1, Fig. 2) of the short-winged form. Macropterous individuals are 2.2 mm. long 51 ENTOMOLOGICA AMERICANA Fig. 2. Late nymph of Corythaica lellula Bueno. Drawn by Mr. F. Y. Cheng. 52 Volume XXXI and brachypterous forms 1.98 mm. long. Both forms are .8 mm. wide. Genus Stephanitis Stal, 1873 The species in this genus are predominantly Palearctic in distri- bution. Of the five now known to occur in North America, two were evidently introduced from Japan and a third is native to eastern North America (Weiss, 1918b). Only these three species will be found in New England. One of the three, Stephanitis glo- bulifera (Matsumura), is apparently a recent introduction and this is the first account of its occurrence in North America. Members of the genus are easily distinguished from repre- sentatives of the related genera in this area. In our species of Stephanitis the hood covers the head, the rostral sulcus is not in- terrupted, and the discoidal area of the hemielytron is raised. These features distinguish them from the New England species of Gar- gaphia, in which the hood does not cover the head, the rostral chan- nel is interrupted, and the discoidal area is not elevated. From Corythaica, Stephanitis species are distinguished by their greater size, their wider and nearly erect paranota, their high arched median carinae, and by their more rounded hoods. Finally they differ from Corythuchdi species principally in their more erect, less broadly rounded paranota, in the narrow, evenly curved humeral angle of their hemielytra, and in the much greater length of their antennae. Stephanitis globulifera (Matsumura), 1905. Tingis globulifera Matsumura, 1905, p. 36, PI. 19, Fig. 16. In studying a collection of Tingidae from the New Haven, Con- necticut Agricultural Experiment Station, two somewhat damaged specimens of an unfamiliar Stephanitis were noted. Since no de- scription of the species could be found in the literature dealing with American species, they were sent to Dr. Reece I. Sailer for deter- mination. They proved to be Stephanitis globulifera (Matsumura) when compared with specimens in the National Museum. The spe- cies was first described by Matsumura as Tingis globulifera in 1905. Later Horvath (1912) properly transferred it to the genus Steph- anitis and redescribed it in some detail. In a 1930 publication Mat- sumura supplied an English translation of the description which is not very satisfactory and a very small, unsatisfactory figure is also given. Since these three references are not generally available, it seems desirable to include a brief comparative description of this recent addition to our insect fauna. 53 ENTOMOLOGICA AMERICANA The two specimens mentioned above were sent to the Experi- ment Station by Mrs. L. B. Winton of Greenwich in late October, 1946. Therefore, correspondence was initiated to gather more de- tails of their occurrence. Mrs. Winton kindly kept me well in- formed concerning the appearance and development of the popula- tion in her garden during the summer of 1950. However, it was after mid-August before many adults were observed. On August 23 I visited her garden and found a heavy infestation of nymphs and adults (mostly somewhat teneral) on a splendid specimen of Pieris japonica (Thunb.) Don, planted in a sheltered corner be- tween the house and an open porch. More than 150 adults were col- lected in a few minutes and a score or so more were kept alive for further study. Mrs. Winton reported that the lace bugs were first troublesome on the Pieris in 1945. By the following year they were destruc- tively abundant. For a time she considered removal of the host plant because it was so seriously injured by them. However, by frequent spraying, continued intermittently even throughout the mild winter of 1949-50, the population was somewhat controlled and the plant was still vigorous at the time of my visit. The late appearance of the adults suggests that this species overwinters in the egg stage, as do the other two species of Steph- anitis that occur in New England and that also infest members of the plant family Ericaceae. Of added interest is the fact that Stephanitis pyrioides (Scott) was collected on a deciduous azalea on the opposite side of the house. This species was not found on Pieris. However, a few specimens of S. globulifera were associated with S. pyrioides on the azalea. Such other ericaceous plants as Kalmia and Rhododendron in her garden supported no lace bugs at that time. It is evident that this recently introduced species may become a serious pest of Pieris and possibly of other ornamental Ericaceae. Through Dr. C. L. Remington I learn that for two or three years the nurserymen of Fairfield County, Connecticut, have complained of serious damage to Pieris by lace bugs. Since other species are not known to feed on that host, it is apparent that 8. globulifera is already well established. At this time it is only possible to sug- gest that eggs of the species were probably introduced before 1945 in the foliage of evergreens shipped from Japan or elsewhere. Mrs. Winton knew of infested plants in four or five gardens within three to eight miles of her home. She thought these infestations were probably of earlier origin than the one on her Pieris. 54 Volume XXXI The following notes provide criteria for the identification of the three species of Stephanitis now established in the northeast. Both 8. pyrioides (Scott) and 8. globulifera (Matsumnra) may be readily distinguished from 8. rhododendri Horvath by their some- what greater length, by their darker hood and hemielytral mark- ings (which become intensely black in 8. globulifera) , by their much more inflated hoods (again extreme in 8. globulifera) , and by their much abbreviated lateral carinae. The paranota of 8. rhododendri flare conspicuously. This species also differs from both the other species in the greater width of the hemielytra and in the abundance of silky setae on all the nervures of the mem- branous parts. The hypocostal ridge of the hemielytron of 8. rhododendri is mostly biseriate and partly triseriate. It is uni- seriate in the other two species. The differences between 8. pyrioides and 8. globulifera are less obvious, but, nevertheless, pronounced. The most noticeable dis- tinguishing features include the conspicuously dark color pattern of 8. globulifera. In this species the entire hood of mature specimens is black. All the pronotal (including paranotal) nervures, except the apex of the median carina, are black, as are most of the hemi- elytral nervures. Areolae of the hood, the discoidal, and the su- tural areas are fumeous, as are the cells of the basal and apical bands. Although the color pattern of 8. pyrioides is similar, the paranota and the discoidal elevations are nearly colorless and in all areas the coloration is brownish and much less intense. Only the areolae of the hemielytral bands are fully infuscated. Interesting differences are seen in the relative proportions of the hoods and pronotal carinae of these two species. In 8. globu- lifera the much inflated, globose hood is twice as high at its peak as the crest of the median carina, while in S. pyrioides the hood and carina are sub-equal in height. Both species have the lateral cari- nae much reduced in length as compared with 8. rhododendri. However, in 8. globulifera they are half again as long as the dis- tance between their anterior ends and the back of the hood, while in 8. pyrioides they are about as long as the distance between their anterior ends and the back of the hood. In 8. globulifera the hood is much wider than the distance between the lateral carinae, while in 8. pyrioides the hood is only slightly wider. Both species have the paranota almost vertical rather than flaring as in 8. rhodo- dendri. Although differences in the relative lengths of the anten- nites and differences in other features can be shown, they are slight and those indicated are adequate for the ready separation of the 55 ENTOMOLOGICA AMERICANA three species now occurring in New England. 8. globulif era (Mat- sumura) is 3.7 to 4.0 mm. long and 2.0 mm. wide. Stephanitis pyrioides (Scott), 1874. Tingis pyrioides Scott, 1874, p. 440. Stephanitis azaleae Horvath, 1905, p. 568. As previously indicated, 8. pyrioides is readily distinguished from 8. rhododendri by its much larger globose hood and by its uniseriate hypocostal ridge. From 8. globulifera it is easily sep- arated by its paler color, by the sub-equal height of the hood and median carina, and by the smaller hood that is only slightly wider than the distance between the lateral carinae. This species is 3.6 to 4.0 mm. long and 2.0 mm. wide. Records for 8. pyrioides are relatively few and scattered. This is a very typical situation, which reflects more accurately the dis- tribution of collectors than of the insects. There was a specimen in the collection of the Museum of the Boston Society of Natural History from Pocasset, Massachusetts, and in my collection are specimens collected in Newton Centre on August 15 and October 29, 1949. In the Rhode Island State College collection there is one taken in Kingston. On August 23, 1950, I made a collection in Greenwich, Connecticut. Other records for that state are for New Canaan on September 10 and 20 and for Hamden on August 15. 8. pyrioides (Scott) is a native of Japan (Scott, 1874; Uhler, 1896; Drake, 1923) and was introduced in the egg stage on azaleas imported from that country (Weiss, 1916a and 1918a). It is now locally abundant in New Jersey (Barber and Weiss, 1922), and is known from Pennsylvania and the District of Columbia (Blatchley, 1926). Froeschner (1944) found it also in Missouri. The species has also become established in Holland (Dickerson and Weiss, 1917). All of the host records are for azaleas, and both the evergreen and deciduous kinds, as well as hardy and greenhouse varieties, are susceptible to infestation. My Newton Centre specimens were found on Rhododendron calendidaceum (Michx.) Torr., an Amer- ican deciduous azalea. They were plants set out several years ago, and the lace bug infestation is of fairly recent origin. Weiss 1918b) records Azalea indica (Rhododendron indicum Sweet) and Azalea amoena (R. oltusum Planch, var. amoenum Rehd.), both semi-evergreen forms, as the hosts. He states (see also Dickerson and Weiss, 1917) that infestations on the deciduous varieties are 56 Volume XXXI not as severe as on the evergreen kinds. This is undoubtedly re- lated to the fact that Stephanitis pyrioides (Scott) overwinters in the egg stage. Since the eggs are deposited in the leaves, they probably have a better chance of surviving the winter in the living leaves on the plants than in fallen leaves that, under garden condi- tions, are likely to be raked up and destroyed. This, in part, may also account for the relative rarity of the species in New England, since the evergreen azaleas are near the limit of their hardiness and are not too commonly grown. Dickerson and Weiss (1917) record the following species of azalea and their hybrids and varie- ties as susceptible to infestation: Rhododendron molle Don and R. schlippenbachii Maxim., both deciduous; R. mucronatum Don and R. yedoense Maxim., both semi-evergreen, and R. ponticum L., an evergreen species. As usual, the tingids live on the under side of the leaves. Nymphs and adults pierce the leaves and destroy the mesophyll which results in a blanching and browning of the upper surface. In heavy infestations the leaves are dried out and many drop dur- ing the summer. The lower leaf surface is also mottled with the excrement of the lace bugs. The early stages and life cycle were studied in some detail by Dickerson and Weiss, and a good account with an excellent plate, showing egg, nymphal instars, adult, male terminalia and parts of the female external and internal genitalia, is given in their paper (1917). Some of their findings are quoted below. Eggs : The eggs are 0.4 mm. long and 0.18 mm. wide. They are smooth, white and flask-shaped, with the opercular end bent to one side. Since they are inserted in the leaf tissue along the mid-rib and larger veins, the curved neck brings the cap just above the lower leaf surface. Each operculum resembles an oval or irregu- larly circular whitish disc. They are sometimes covered with a brownish, crust-like deposit. One to ninety eggs may be found in a single leaf. Usually the younger leaves are chosen by ovipositing females. Most are placed irregularly along the mid-rib. The eggs overwinter and in southern New Jersey hatch towards the end of May. By late June adults appear and lay eggs that hatch in about two weeks. In late July and early August, second brood adults appear and another two week period of oviposition follows. These produce a third generation of adults by late September, and during early October they lay the eggs that will overwinter. Nymphs: Five nymphal stages occur. The length of each in- 57 ENTOMOLOGICA AMERICANA star varies from three to six days (Weiss, 1918b). The average time for development from egg to adult in southern New Jersey is only a month. As the figures show, the nymphs are elongate oval and bear conspicuous spines dorsally and at the posterior dorso- lateral angle of each abdominal tergite. Adults: As noted above, three generations appear each season in southern New Jersey. At the peak of each cycle of adult abun- dance the proportion of males and females is about equal. Females, however, seem to live longer than the males and the former are always more numerous during the egg laying periods. The last brood adults may be found until late in November and these linger- ing individuals are usually females. The males apparently die soon after copulation. The deposition of a single egg takes the female two to three minutes. She pushes the ovipositor from its sheath and touches the surface of the leaf until she locates a satisfactory position. Then the ovipositor is thrust into the leaf tissue until the abdomen comes to rest against the leaf and is then withdrawn. On account of the several extended periods of oviposition, all stages may be found together from late June until October. In central New Jersey two full broods and a partial third one de- velop. Further north the season is shorter and the cycle probably takes longer. This also accounts in part for the smaller populations that occur in central New England. Stephanitis rkododendri Horvath, 1905, p. 567. Leptobyrsa explanata Heidemann, 1908, p. 105, PI. 4. Figs, d, e, £. This species is at once distinguished from both the others oc- curring in New England by the irregularly biseriate hypocostal ridge. This structure is uniseriate in the others. Horvath (1905) described the insect from cultivated rhododendrons growing in Holland and observed that it had been introduced into Europe. This view is supported by others (Dickerson, 1917), although the fact that tingid eggs have been found on rhododendrons imported from Holland (Weiss, 1915) may have led some to the erroneous belief that this lace bug was introduced into the Atlantic states from Europe (Blatchley, 1926). On August 5, 1950, I collected them on native Kalmia latifolia L. in East Hartland, Connecticut, where infestation from cultivated plants was most unlikely. Such evidence, coupled with the marked structural differences between 58 Volume XXXI S. rhododendri and the two established Palearctic species, points to eastern North America as the place of origin of this species. There are records from all of the New England states except Vermont. It is the species of Stephanitis most commonly collected. Parshley (1917c and 1923b) has published numerous Massachusetts and Connecticut records which suggest a general distribution. The few Maine records include Bar Harbor ( fide Sailer), Southwest Harbor (Proctor, 1946), both on Mt. Desert Island, and Orono (Parshley). There are specimens in the University of New Hamp- shire collection from Durham. The Rhode Island State College collection contains specimens from Kingston and Newport. The range extends from New England west to Ohio and south to Florida. In actuality, it is likely to be found wherever rhododendrons are grown since it is commonly distributed in the egg stage with nurs- ery stock. Like other species in the genus, the host plants are all Ericaceae. The native Kalmia latifolia L. and Rhododendron maximum L. are the hosts usually recorded and the plants from which my collec- tions have been made. Barber and Weiss (1922) report Kalmia angustifolia L. as another host. White (1933) observed that plants fully exposed to the sun were most heavily infested and most seri- ously injured. He adds Pieris japonica (Thunb.) Don and Pieris floribunda (Pursh) B. and H. to the list of hosts. Dickerson (1917) quotes from Dutch sources that the species has been found on 120 varieties of rhododendrons and azaleas. Our present knowledge of the biology of Stephanitis rhododendri Horvath results primarily from the studies of Heidemann (1908), Crosby and Hadley (1915), and Dickerson (1917). The following account is based on their papers. Each of these publications con- tains plates of significance. Some of their figures are reproduced in the articles by Weiss (1918b) and by Barber and Weiss (1922) which summarize their contributions. Eggs : The egg is ovoid-cylindrical in shape with the opercular end slightly constricted and bent to one side. They are smooth and yellowish white. The length is about 0.4 mm. and the diameter about half the length. The operculum is oval and makes the end obliquely truncate, whereas the opposite end is rather evenly rounded. Most of the eggs are laid in irregular rows along the mid-rib. Some may be found inserted in the other large leaf veins or even in the blade a few millimeters from any vein. They are always laid on the lower side of the leaf and the younger leaves are 59 ENTOMOLOGICA AMERICANA selected (see Crosby and Hadley, 1915, PL 22, Pigs. 2 and 3, PL 23, Fig. 2; Dickerson, 1917, Pl. 8, Pig. 6). The leaf cells surrounding the eggs become corky and form a small gall-like mass that may be easily separated from the normal tissue. The egg cap is level with the leaf surface and, as is usual among the Tingidae, the females deposit a brownish substance that forms a varnish-like scab over the eggs. This crust often peels away before the eggs hatch. Pew to as many as 176 eggs may be counted in a single leaf, but the number laid by a single female has not been ascertained. In laying the egg the abdomen of the female is parallel with the leaf and the extended ovipositor is driven backward and ventrally into the leaf. The eggs of this species overwinter in the evergreen foliage of the host plants. In the District of Columbia, Heidemann (1908) found un- hatched eggs and recently hatched nymphs on April 20. In New Jersey (Dickerson, 1917) they are said to emerge in early May. In Ithaca, New York, they hatch in late May and early June (Crosby and Hadley, 1915). Egg laying probably starts in late June and continues through July. There is evidence that the eggs hatch over a prolonged period since nymphs are sometimes seen in early August. In New Jersey two broods may develop. There is need for clarification of such details. Nymphs: Most notable is the fact that this species apparently has only four nymphal instars. This is an exceptional condition among the Tingidae, where five usually occur. Heidemann (1908) has described the last stage and figured a nymph (Pl. 4). Crosby and Hadley (1915) figure the four instars (Pl. 23) and describe them in detail. They describe hatching in the following manner : 1 ‘ In hatching, the end of the egg enlarges slightly, becom- ing almost transparent. Then the nymph gradually emerges, the red eye-spots showing very conspicuously. The body sways back and forth slowly, during emergence, with slight spas- modic movements. When all the body has emerged except the last two or three segments, the spines along the sides of the body and the legs stiffen out. Then the leaf surface is grasped by the claws of the legs, and the insect is able with this help to withdraw the body entirely. Often the egg shell is drawn partly out of its pocket by this last effort. The newly emerged insect rests for several minutes after its exertions, then it slowly walks around, seeking a suitable feeding place. When 60 Volume XXXI first emerged the nymph is colorless, almost transparent, ex- cept for the bright red eyes. Soon after feeding commences, it begins to darken up. 1 1 The nymphs feed in groups, remaining in a place a short time, then moving to a new location. The characteristic feed- ing attitude is with the body inclined upward towards the head, the antennae straight outward slightly diverging. At intervals the body sways slightly from side to side. After feeding a short time, the proboscis is withdrawn, and the slen- der inner suckang tube is carefully cleaned with the forefeet, in much the same fashion as the ordinary house fly. ’ ’ Under insectary conditions in Ithaca they found that the length of the various nymphal stages was as follows : ‘ ‘ First stage, 6 to 7 days ; Second stage, 4 to 6 days ; Third stage, 3 to 6 days ; Fourth stage, 12 to 15 days. ” They also suggest that development may require somewhat longer out-of-doors. Their observations give a minimum time of 25 days and an average of about 30 days for that region. In New Jersey, Dickerson (1917) found that some nymphs completed their growth in only twenty days. This supports his view that two broods may mature there and further south. Dickerson (1917) observed that the first three nymphal instars stay together and move very little, but the final stage nymphs move about more freely. He also noted that the rostral sheath doubles on itself when the insects feed and the tip of the beak guides the extended lancets. Adults: The feeding of adults and nymphs causes the usual mottled whitening of the upper leaf surface characteristic of lace bug injury. In heavy infestations the leaves become dry and shriv- eled. This makes the plants unsightly in the garden and may hinder the sale of nursery stock. The under side of the leaves be- comes thickly dotted with the rusty brown excrement of the feed- ing bugs. Adults have been collected from June 6 to September 2 in Connecticut. In favorable seasons their period of activity probably somewhat exceeds those limits. Specimens have been taken in Princeton, Massachusetts, as late as October ( fide Sailer). Stephanitis rhododendri Horvath is usually a little shorter than the other two species and varies from 3.3 to 3.7 mm. long. It 61 ENTOMOLOGICA AMERICANA is, however, somewhat wider because the hemielytra are unusually broad beyond the discoidal area. They are about 2.4 mm. in width. Heidemann’s figure of the adult (1908, PI. 4, Pig. d) is an excellent representation. Dickerson (1917) gives one of the first and one of the few de- scriptions of tingid external genitalia. He figures (Pl. 8) the male terminalia from the dorsal (Pig. 1) and ventral (Fig. 2) aspects, the ventral aspect of the .three sternites of a female showing the ovipositor in position (Fig. 3), the ovipositor in transverse section showing the inter-relationship of the valves (Pig. 4), and a greatly magnified view of the serrated element of the ovipositor that makes the incision in the leaf tissue (Pig. 5). Genus Corythucha Stal, 1873 All members of this large genus are restricted to the Western Hemisphere. About sixty-three species and a few varieties have been described (Hurd, 1946). Of this number, fifty-one are North American, and of the fifty-one, fifteen species and one variety are found in New England. Many of the species are strikingly simi- lar and, as a group, show evidence of close relationship. Professor Parshley (1923b) pointed out that our knowdelge of the limits of variation within any one species is far from adequate. This is still true and, because of their close affinities the species are, in some cases, difficult to identify. Although several species seem to have very specific food habits which aid in their identification, if host records are available, such evidence can be misleading since other species are less selective and since recent observations will show that the same species may have different host preferences in dif- ferent parts of its range. Species of Corythucha are among our commonest tingids. Since they are gregarious, their feeding damage is often conspicuous and announces their presence on a variety of common native and cultivated plants. Accordingly, some of them are frequently col- lected. However, the biology of some of the most common species has been neglected, and there is much we could profitably learn about their habits and the ecological factors controlling their dis- tribution. The New England species usually live on the under side of the leaves of trees and shrubs but a few live on herbaceous plants. As far as we know they all overwinter as adults under loose bark on trees, amongst fallen leaves, and under stones or other shelter. 62 Volume XXXI The genus Corythucha is readily distinguished from other local forms. Only three genera represented in this region have large pronotal hoods covering their heads. In addition to Corythucha , these are Corythaica and Stephanitis. From Corythaica bellula Bueno, our only native species, Corythucha is readily distinguished by the hood which is distinctly globose posteriorly, by the broadly rounded paranota, by the strongly constricted and reflexed humeral area of the hemielytra, and by the larger size of all local species. Corythucha differs from Stephanitis also in the humeral hemielytral constriction and reflection, in the marked constriction and forward projection of the hood which reaches beyond the base of the third antennite, and in the more broadly rounded, medially bullate para- nota. Corythucha species are sub-quadrangular in outline, which increases their distinctiveness. Uhler (1886), Provancher (1886), Banks (1910), Blatchley (1926), and others commonly misspelled the generic name Corythuca , leaving out the final h. Corythucha arcuata (Say), 1832. Tingis arcuata Say, 1832; in Fitch reprint, 1858, p. 794; Say (in LeConte edition), 1859, p. 349. Corythucha mali Gibson, 1918, p. 98. Corythucha arcuata var. mali Drake, 1921, p. 54. Froeschner (1944), Dr. Sailer (letter), and I are agreed that, since all gradations from the typical C. arcuata (Say) color pat- tern, with a full basal and apical bar, to the extreme mali pattern, with the apical bar entirely lacking and other markings reduced, may be collected at one time from the same oak tree, there is no valid reason for using a varietal name. Blatchley (1926) gives the range of this species as New England and Quebec west to North Da- kota and Colorado and south to Texas and Alabama. Froeschner (1944) lists Missouri stations, and specimens have been seen from Virginia. Parshley (1915) lists three localities for Maine, and to the three mentioned Albion (September 8, 1949) should be added. His (1917c) New England records give two for New Hampshire (Durham and Hampton), numerous Massachusetts and Connecticut localities, and Kingston, Khode Island. Another addition seems to be the first for Vermont. Specimens were found on Quercus Prinus L. on August 31, 1949, in Wells. This lace bug probably occurs, usually on members of the white oak group, throughout this region. On the campus of the University of Massachusetts, Morrill (1903) found Quercus alba L ., Q. Prinus L., Q. rubra L., Q. Muehlenbergii 63 ENTOMOLOGICA AMERICANA Engelm. heavily infested and a light infestation on Q. macrocarpa Michx. However, other oaks growing nearby, in some cases with their branches touching infested trees, seemed to be quite immune from attack. These species were : Quercus coccinea Muenchh., Q. ilicifolia Wang., and Q. laurifolia Michx. Drake (1928a) states that they occasionally breed on chestnut and very rarely on maple and apple. Quercus prinoides Willd. is also mentioned as a host (fide Sailer). The season of activity is suggested by Massachusetts records which extend from May 20 (Bailey) to October 9 (Bueno, 1924a). For the District of Columbia the period lasts from May 2 to No- vember 21, and McAtee (1923) observed pairs mating as late as September 27. Bueno (1924a) found an oak in Amherst, Massa- chusetts, heavily populated with eggs, nymphs, and adults on Oc- tober 9. They hibernate beneath leaves and in other sheltered places on the ground (Drake, 1928a). In his paper describing the immature stages of Corythucha ar- cuata (Say), Morrill (1903) set a fine example for subsequent studies of the Tingidae. A splendid plate of the egg and five nymphal stages is included (PL 3). There are, however, many gaps yet to be filled concerning details of the life cycle of this species. Eggs: The eggs of this species are noticeably tapered at the basal end, but are otherwise of the usual sub-cylindrical shape. They are about .56 mm. long and the greatest diameter is .24 mm. There is a narrow collar at the apical end on which the operculum rests. The cap itself is a low cone with fifteen to twenty ridges con- verging from the base to meet at the apex. There may be a slender filament of variable length rising from the apex where the ridges meet. It is commonly one-fifth as long as the egg. The eggs are smooth and shiny black. They generally are covered with a dark rough coating that may be easily rubbed away. Eggs are laid on the lower side of the leaves and may be found from late May (Morrill, 1903) until early October (Bueno, 1924a) in Massachusetts. They are usually laid in roughly circular patches in which the individual eggs are well-spaced. The eggs are often near the larger leaf veins but may be anywhere on the leaf blade. Single eggs are found, but as a rule the clusters contain twenty-five to fifty, and over one hundred may be counted on a leaf. Each female probably lays many eggs, but no definite figures are available. The late copu- lation mentioned by McAtee (1923) suggests that impregnated females may hibernate and lay their eggs the following spring. The incubation period of the eggs is not established. 64 Volume XXXI Nymphs: Morrill’s full descriptions and figures (1903) of the five nymphal instars make any detailed account unnecessary. Like other tingid nymphs, those of C. arcuata feed in groups. As they move around on the lower leaf surface they deposit shiny black excrement in dots. Their feeding results in a whitening of the upper leaf surface. As is usual in heavy lace bug infestations, the foliage injury may be very conspicuous and seriously detracts from the appearance of the plant. The early stadia last two or three days, but from six to seven days pass between the fourth and fifth moults. It therefore takes at least eighteen days for the nymphs to mature and may take twenty-four. In central New England the early summer brood probably matures in thirty or forty days. Whether or not another generation develops before late fall has not been determined. Morrill (1903) found nymphs of a predaceous heteropteron very active in destroying the young nymphs of C. arcuata. Adults: Since adults of our New England species of Corythucha may be identified by use of the key given previously, only a few of the more striking features will be mentioned in the discussion of the species. The few details given here should not be considered ade- quate for determination of the species. In many cases host records are useful indicators, but any species may at times be collected from plants on which it does not breed nor even feed. Therefore, even the host records of careful collectors should not be relied on com- pletely. Pour species in this region have the hood and median carina sub-equal in height. C. arcuata (Say) is somewhat smaller than the other three. One of the greatest differences is seen in the ex- ternal male genitalia (see Pigs. 5A, D, and E, and Pig. 6G). The figures are all on the same scale and it is immediately obvious that the genital capsule of C. arcuata is only about half as large as any of the other three. There are also differences in the actual form of the hood and in other features. In both C. caryae sp. nov. and C. pruni 0. and D. the hood is appreciably larger and more broadly globose and flattened behind. Spines on the hood nervures are few in C. caryae and practically absent in C. pruni. The hood of C. ciliata (Say) is noticeably broader and slightly flatter than that of C. arcuata. The whitish opacity of the hood nervures and areolae of C. ciliata and the numerous black-tipped spines on the nervures also differentiate it from all other species. C. arcuata (Say) is 3.0 to 3.3 mm. long and about 1.6 mm. wide. 65 ENTOMOLOGICA AMERICANA Corythucha as so data Osborn and Drake, 1916b, p. 14. Corythucha spinulosa Gibson, 1918, p. 79. The only New England state from which this species is now known is Connecticut. Localities are sufficiently numerous (New Haven Experiment Station and Parshley collections) to suggest the general occurrence of C. assodata 0. and D. in that state. Records available for Connecticut range from May 27 to September 23. McAtee (1923) gives records for the District of Columbia ranging from May 18 to October 2. The species is found from New York to Ohio and Indiana and south to Mississippi and Georgia (Monte, 1940). Prunus serotina Ehrh. is the host for this species (Osborn and Drake, 1916b and Drake, 1928a). Under the name of C. spinulosa Gibson the early stages of this species are described and some biological information given by Dickerson and Weiss (1918). A plate (7) showing the egg, five nymphal stages, the adult, and feeding injury to the cherry leaf accompanies their paper. Their study is the source of the following details. Eggs: The eggs are about .55 mm. long and are .2 mm. wide. They are sub-elliptical. The basal half is translucent and the apical half is dark brown. The end attached to the leaf is acute, but the tip is rounded. There is a slight tapering toward the truncate apical end, which has a collar-like rim and a low cone-shaped opercu- lum. In New Jersey females emerge in early June, feed, and soon begin to oviposit. By June 20 most of the eggs have been laid and the females have mostly disappeared. The eggs are usually inserted irregularly in and along the mid-rib on the lower side of the cherry leaf. Most project almost parallel to the leaf surface, but some are perpendicular to the blade and a few may be placed at an angle in the mid-rib tissue. Only the rounded abopercular end is inserted into the leaf. The exposed part of the egg is covered with the black varnish-like excrement of the female. From 4 to 35 eggs may be counted on a single leaf, but the average number is about fifteen. Eggs start to hatch about June 25 and by July 15 adults are seen again. Incubation of the eggs takes two or three weeks. Nymphs: Full descriptions of the five instars and a figure of each are given by Dickerson and Weiss. Each of the first three stadia lasts two or three days, the fourth takes three or four days, and the fifth requires seven to nine days. The minimum time for development from egg to adult would be about thirty days and the 66 Volume XXXI maximum probably about forty days. Depending largely on temp- erature, the life cycle would require four to six weeks. The nymphs feed in the usual groups and preferably close to the mid-rib. If scattered they will soon reassemble. The last two instars tend to be a little more independent. By late July most of the first brood nymphs are mature. Adults: There are nine species of Corythucha in New England with hoods twice as high as the median carina. Three of the six have very small hoods and extremely low carinae and would, therefore, never be confused with the others. The three with small hoods and low carinae are C. mollicula 0. and D., C. pergandei Heidemann, and C. ulmi 0. and D. The small size of C. bellula Gibson, C. coryli 0. and D., and C. marmorata (Uhler) (from 2.8 to 3.4 mm. long) immediately distinguishes them from C. associates. C. pallipes Parshley is most like C. associata in size and in hood form. However, the areolae of the hood of C. pallipes are less than half as large as the cells of the hood of C. associata and the nervures of the latter are much embrowned, while the nervures are pale in C. pallipes. A comparison of Pig. 5B and Fig.6E will reveal differ- ences in the male genitalia. Adults of C. associata measure about 4.0 to 4.2 mm. in length and are 2.4 mm. wide. They feed on the under surface of the leaves with the nymphs and cause a whitening of the central part of the blade along both sides of the mid-rib on the upper surface. In New Jersey adults of the first brood begin to appear in abundance by late July. By the end of August second brood adults are found and they continue to develop well into September. It is this generation that later hibernates. Corythucha bellula Gibson, 1918, p. 93. This small species somewhat resembles C. coryli 0. and D. and is also closely related to C. hewitti Drake. It may be distinguished from the former by its slightly larger size, by the much smaller hood areolae, and by the more conspicuous spots on the paranota. From C. hewitti it differs in having shorter marginal spines, a more angu- late and narrowed hood and in usually having some large hyaline areolae in the apical band (Drake, 1918) . In all these features there is considerable variation and the specimens from northern New England suggests that these two species should be studied further for clarification of their specific limits. Males of the specimens from Maine have a larger genital capsule and their claspers are stouter and blunter than in males of C. coryli from Massachusetts 67 ENTOMOLOGICA AMERICANA (Pigs. 5B and P). They are about 3 mm. long by about 1.7 mm. wide. My collection contains material from Harvey, New Brunswick ; East Brighton, Vermont; Daigle, Houlton, Madrid, Moose River, and Pleasant Landing, Maine. They were collected from September 2 to 9 in 1949. The host plant in all cases was Corylus. Outside of New England published records include New York and Ohio (Hurd, 1946). The type specimens were found on Crataegus (Gibson, 1918). Drake (1928a) reports the following plants as hosts: Crataegus succulenta var. neofluvialis (Ashe) Palmer, C. punctata Jacq., C. pruinosa (Wendl.) K. Koch (Rosaceae), Alnus incana (L.) Moench (Corylaceae), and Bibes oxycanthoides L. (Saxifragaceae) . In the same paper he states that adults hibernate in bark crevices and among fallen leaves. In New York they are active from May into September. No other biological information is available. Corythucha caryae sp. nov. This lace bug from hickory resembles Corythucha pruni 0. and D. in the shape of its hood and superficially resembles both 0. arcu- ata (Say) and C. ulmi 0. and D. in the lack of an apical hemielytral band. C. caryae is a little larger than the last two species named and it differs from both in having a larger, broader hood that is noticeably flattened. The hood of C. caryae is very slightly higher than the crest of the median carina. Although the hood of C. ulmi O. and D. is small, it is about twice as high as the median carina. Males of C. caryae (Pig. 5D) are easily distinguished from C. arcu- ata (Say) (Fig. 5A) by the much larger genital segment and both sexes are distinguished by the large areolae of the hood and the whitish opacity of the paranota of C. caryae. C. pruni 0. and D. is appreciably larger than C. caryae and the hood of the former is smoother, almost spineless, and much less sharply constricted. In C. pruni 0. and D. the hood areolae are somewhat clouded with yellowish brown and the nervures are all amber. The hood areolae of C. caryae are sub-hyaline and a few nervures of the crest are dark brown, the rest pale amber. Males of C. caryae (Fig. 5D) are at once separated by their claspers, which taper to a rather acute point while the claspers of C. pruni 0. and D. (Fig. 6G) have almost spatulate tips. The antennae of C. caryae are more or less densely clothed with bristly hairs of various lengths. On the terminal antennite the 68 Volume XXXI bristles are especially short and numerous, giving the tip a tufted appearance. Antennite one is barely twice as long as antennite two and, except where it tapers proximally, of the same diameter as the second. Antennite three is more slender and much longer than the others. It is twice as long as antennite four, which is clavate and broadest distally. The rostrum extends just beyond the meso- metasternal groove and reaches the hind coxae. The sternal ridges have a single row of conspicuous hyaline areolae set off by stout brownish black nervures. The greatest width of the hood is 0.6 mm. and it is 0.9 mm. long. It is constricted rather abruptly to less than half its greatest width anterior to the middle, and rather abruptly tapers at the tip. A few short spines occur on the nervures of the hood, especially on the sides of the constricted portion. The pos- terior part is sub-globose and noticeably flattened dorsally, which gives this species a resemblance to C. pruni 0. and D. The areolae of the hood are more than twice as large as those of the paranota. The paranota are evenly areolate and bordered by sharp, black- tipped spines, some of which also occur on the nervures of the upper surface. The anterior and posterior margins are strongly raised and the mid-area of the paranotal membrane is bullate. The pronotum and the base of its triangular process are punctate. The posterior portion of the process gradually approaches an areolate condition. The lateral carinae are highest anteriorly and the 3-4 areolae become increasingly smaller posteriorly. The median carina is foliaceous, slightly raised anteriorly, but slightly lower than the crest of the hood. It consists of a single row of areolae with a partial second row beneath and just behind the large anterior cell. A few spines occur on the nervures of the carina and sides and apex of the hood. The apex of the hemielytron is evenly rounded and the lateral margins are sub-parallel. Areolae of the costal area are in four rows and are small from the posterior margin of the basal cross-bar forward. Behind the bar there are three rows and the cells rapidly become much larger. The subcostal area is tri- seriate and the areolae are very small except posteriorly, where a single row of appreciably larger areolae occurs. The humeral ele- vations are a little lower than the crest of the median carina. Spines along the costal margin of the hemielytra become increasingly shorter from the proximal end towards the distal and are entirely absent on the posterior third of the wing. Males are 3.4 to 3.7 mm. long and females are 3.4 to 3.8 mm. long. Males vary in width from 1.87 to 2.0 mm. while females are from 1.9 to 2.2 mm. in width. The 69 ENTOMOLOGICA AMERICANA Fig. 3. Corythucha caryae sp. nov. Drawn by Mr. F. Y. Cheng. average male is about 3.4 mm. long and the average female is about 3.55 mm. long. These measurements are based on thirty males and thirty females selected at random from the type series. Color: A few nervures of the crown of the hood are somewhat embrowned. The hood areolae are sub-hyaline. The paranota have a small brown spot anterior to the tumid center and the areolae are mostly whitish and sub-opaque. The pronotum is dark brown, with a varnished look over the disc, but becoming almost white at 70 Volume XXXI the apex of the triangular process. A few central nervures of the median carina are embrowned. The hemielytra have a single bar just behind the anterior margin. Areolae anterior to the bar are sub-opaque. Areolae and nervures of the bar are fusco-testaceous. The discoidal and sutural areas are also embrowned nearly to the apex, except that the anterior portion of the discoidal elevations has hyaline areolae and pale nervures. The sub-costal area is colorless except in the region of the basal band. The areolae of the costal area and the apical margin are distinctly hyaline. There is no apical band, but a few nervures near the posterior union of the costal and subcostal areas may be darkened. Without magnifica- tion the cross-shaped pattern of the dorsal markings is rather strik- ing. Antennae and legs are amber, while the rostrum and tarsi are slightly darker. The ventral side of the body is black, except for some reddish amber areas about the caudal end. I am indebted to Mr. F. Y. Cheng for Fig. 3 which is an excellent representation of this species. The first specimens seen were given to me by Dr. William L. Nutting, who had collected them on young hickories in a Salem, New Hampshire, field in July of 1946. Since that time I have col- lected the same species on various occasions from July 13 to Sep- tember 6 in the towns of Georgetown, Newbury, and West New- bury, Massachusetts (Essex County). They have always been found on hickories (probably Carya ovata (Mill.) K. Koch). There are specimens in my collection taken during each of the last four seasons. On March 25, 1948, a single adult was found in hiberna- tion under the exfoliating bark on a hickory trunk. Early stage nymphs were numerous on July 13, 1950, and in 1948 a few late in- stars were found on September 6. A lot of about 250 adults col- lected on Carya in Newbury on August 7, 1950, is herein desig- nated the type series. The holotype (a male) and an allotype from this series will be deposited in the Harvard Museum of Comparative Zoology. Paratypes will be placed in the United States National Museum collection. Numerous paratypes and topotypes remain in my collection and some of these will gladly be distributed on request. On August 22, 1947, specimens were taken in Newbury (near Byfield Center) and several of them had mites attached. Some of these were sent to Dr. Edward W. Baker at the United States Na- tional Museum. He kindly identified them as larval Erythraeidae of the genus Leptus, which are known to be insect parasites in the larval stage and free living predators in the nymphal and adult 71 ENTOMOLOGICA AMERICANA stages. Other instances of mites on tingids will be occasionally noted. Corythucha ciliata (Say), 1832. Tingis ciliata Say, 1832; in Fitch reprint, 1858, p. 793; Say (in LeConte edition), 1859, p. 348. Gibson (1918) states that C. ciliata (Say) may be found on the sycamore tree {Plat anus) wherever it grows east of the Rocky Mountains. This species differs from all others in the area in the whitish opacity of all the areolate areas. The concolorous pattern is complete, except for a brown patch on the medial posterior as- pect of the discoidal elevations. Short black-tipped spines are nu- merous on the dorsal nervures. In C. ciliata the hood is small, somewhat flattened and sub-equal with the median carina in height. Its areolae are only a little larger than those of the paranota. The external male genitalia (Fig. 5E) are large and the bluntly rounded tips of the claspers separate this species from all except C. pruni 0. and D. In 0. pruni the clasper tips are decidedly spatulate (Fig. 6G). C. ciliata is about 3.75 mm. long and 1.6 mm. wide. Parshley (1914) notes that the species has been collected in Maine. There seem to be no New Hampshire records, but a heavily infested sycamore was seen while driving through South Lee on September 11, 1949. On August 31, 1949, specimens were collected on Plat anus in Wells, Vermont. Massachusetts and Connecticut records are fairly numerous (for some of them see Parshley 1917c and 1923b). There are a few specimens from Kingston in the Rhode Island State College collection. Drake (1919a) reports that both nymphs and adults of this species have been collected on Fraxinus, Cary a ovata (Mill.) K. Koch and Broussonetia papyrifera (L.) Vent, in Missouri. In the same state Froeschner (1944) records adults from cypress. A note on Dr. Sailer’s list gives the interesting information that on April 5, 1949, Mr. Roy Latham found adults abundant on Chamaedaphne growing in a Long Island, New York, cedar swamp. C. ciliata (Say) is almost invariably found on mature sycamores no matter how isolated they may be from other trees of the same genus. This constant association is probably favored by the nature of the sycamore bark, which exfoliates in large irregular pieces. As the outer bark gradually loosens, it offers apparently ideal shelter for hibernation, and overwintering adults may be found in abun- dance under the loose bark of such trees from October until as late 72 Volume XXXI in the spring as early June in central New England. Froeschner (1944) says they also hibernate under logs, under the bark of other trees and in grass clumps. To such situations, McAtee (1923) adds among old leaves and on the foliage of scrub pine in the Dis- trict of Columbia. Massachusetts records at hand run from mid-March until early October. The extremes mentioned are records for hibernating adults. The duration of their seasonal activity will correspond closely with the period in which the trees are in leaf. By mid- summer most sycamores show obvious feeding damage. The leaves become much whitened except around the margins. Some may wilt, turn brown, and even fall prematurely. Despite the abun- dance of this rather common species, the life cycle has not been studied in any detail. Eggs: Heidemann (1911a) says the eggs are hidden in the pubescence on the leaf. Barber and Weiss (1922) state that the egg bases are inserted into the tissue along the larger veins and their forks on the lower side of the leaf. They are said to occur singly or in groups of no more than ten. Nymphs: Morrill (1903) showed that each of the five nymphal stages could be distinguished from the corresponding instars of C. arcuata (Say). Barber and Weiss (1922) suggest that nymphal development takes about three weeks in New Jersey. They esti- mate that maturation from egg to adult requires about five weeks and that probably two broods develop annually. Osborn and Drake (1917a) state that the adults are para- sitized by a red mite. Parasitic mites were noted for C. caryae and for Piesma cinerea (Say) and other instances will be cited below. Weiss (1913) found that hibernating adults placed in a warm room demonstrate a strong negative geotropism. They will climb up on twigs or anything else available. Corythucha coryli Osborn and Drake, 1917a, p. 299. Five of our New England species have high-crested globose hoods about twice as high as the median carina. In addition to C. coryli 0. and D., they are C. pallipes Parshley and C. associata 0. and D., G. bellula Gibson and 0. cydoniae (Fitch). C. coryli is at once distinguished from the first two species by its small size. This is reflected in the relative size of the male genital capsule of C. coryli (Fig. 5F) which should be compared with that of C. asso- ciata (Fig. 5B) and that of C. pallipes (Fig. 6E). The male geni- talia of C. cydoniae are appreciably larger (Fig. 5G) and the 73 ENTOMOLOGICA AMERICANA claspers of C. bellula males are much blunter (Fig. 5C). This species is usually smaller than any other in this region, being only 2.8 mm. long and 1.5 mm. wide. No one has yet described the early stages of C. coryli 0. and D. We know only that the usual host is reported as Corylus americana Walt. Mr. Nathan Banks has collected the species in Holliston, Massachusetts, on Corylus (Museum of Comparative Zoology). These, and the records below, are the only ones available for New England and are all from this state. McAtee (1923) records it from Maryland and Virginia, while Blatchley (1926) gives the range as from New Jersey and Maryland west to southern Indiana. There are specimens at hand from Groveland and Georgetown from Corylus. However, it is of particular interest to record Ostrya virginiana (Mill.) K. Koch as a host. The hophornbeam is an- other native plant in the family Corylaceae. Dr. John T. Wood- land brought me specimens from Stoneham which he had taken on this plant. Since then we have collected them on the same host in the Arnold Arboretum, Forest Hills, and in Stony Brook Reserva- tion, Hyde Park. There are also specimens at hand from a road- side tree in the town of Newbury (Bailey). That Ostrya is a true host is indicated by the fact that eggs and the various nymphal stages have been taken along with numerous adults. The collec- tions known cover the months from late May until mid-September. Early stage nymphs were collected on July 22 and late stages oc- curred until September 10. In the Stony Brook Reservation some plants of Corylus occur under the mature trees, but no lace bugs have been found on them. My only specimens from Corylus came from plants growing along the roadside, where they were fully ex- posed to sunlight. On a few specimens from the Arnold Arboretum large ovoid mites were found. The mites were attached to the abdomen on the dorsal side beneath the wings. They have yet to be identified, but are larger and unlike any other that I have found on lace bugs. Corythucha cydoniae (Fitch), 1861. Tingis cydoniae Fitch, 1861, p. 114, Fig. Corythucha arcuata Comstock, 1879, p. 221. Corythucha arcuata crataegi Morrill, 1903, p. 132. Corythucha crataegi Osborn and Drake, 1916a, p. 229. In Massachusetts C. cydoniae (Fitch) is one of the commonest and most abundant tingids. Like C. coryli 0. and D., the hood of this species is more than twice as high as the median carina. While 74 Volume XXXI small, C. cydoniae averages somewhat larger than C. coryli and the cells of its hood, though large, are also somewhat smaller than the hood areolae of C. coryli. Another marked difference is seen in the general infuscation of the hood and the central area of the paranota. C. cydoniae is more completely and intensely pigmented than other New England species of Corythucha. The external male genitalia of C. cydoniae are appreciably larger than those of C. coryli (Figs. 5F and G). C. cydoniae (Fitch) is about 3.2 to 3.5 mm. long and about 1.7 to 2.0 mm. wide. Although Monte (1940) and Hurd (1946) say this species is distributed generally throughout the United States, some qualifica- tion of such statements is probably desirable. Records for Massa- chusetts are so numerous and scattered that there is no doubt of its general occurrence in this state. Professor Parshley (1923b) indi- cates the same for Connecticut, and specimens in the New Haven Experiment Station collection support him. Plowever, the only specimens seen from northern New England are a few from Dur- ham in the University of New Hampshire collection. On my 1949 collecting trip through Maine, New Hampshire, and Vermont, not a single specimen was found although the usual host plants are fairly common in those states and even locally abundant. While driving in New Brunswick, for example, a hillside for some dis- tance along the highway was covered with clumps of Crataegus. But there were no tingids on them. In the northeast this species is apparently not common north of Massachusetts. Eggs: In his description of the eggs of this species, Professor Comstock (1879), on advice from Professor Uhler, referred them to Corythucha arcuata (Say). Morrill (1903) in his noteworthy paper pointed out that differences in the eggs alone were enough to distinguish the two and proposed a sub-specific name. For some time the fact that Fitch (1861) had already made a distinction be- tween these species was overlooked or disregarded. Comstock (1879) described the eggs as smooth, whitish, glis- tening and semi-transparent. They are ovoid and their broad ends are somewhat inserted into the leaf tissue. The female covers them with a brown, sticky substance that soon hardens and adheres so firmly to the upper part of the egg that it cannot be removed with- out crushing the egg. The opercular end of the egg is sharply truncate, which gives each egg the appearance of a small cone (ibid., PI. 4, Fig. 3). The round opercular openings make it easy to dis- tinguish the empty egg shells from unhatched eggs. They are 75 ENTOMOLOGICA AMERICANA usually laid in groups of from ten to thirty along the sides of the more prominent veins on the lower side of the leaf. In my experi- ence, as many as 100 eggs may be counted on a single leaf and 40 or more frequently occur, as shown by the following observations. I. Eggs of Corythucha cydoniae (Fitch) on Amelanchier leaves at Stony Brook Reservation, May 16, 1949. Leaf 1 carried a total of 12 eggs 4 eggs 65 eggs 104 eggs 100 eggs 17 eggs In all 302 eggs were found, an average of about 50 to a leaf. II. Amelanchier leaves with eggs taken June 1, 1949. Leaf 1 with 1 mass of 22 eggs. 1 1 2 with 1 mass of 61 eggs. i i 3 with 2 masses of 32 and 15 eggs. i i 4 with 1 mass of 3 eggs. 1 1 5 with 1 mass of 60 eggs. 1 1 6 with 2 masses of 40 and 6 eggs. 1 1 7 with 3 masses of 24, 45 and 35 eggs. c ( 8 with 2 masses of 22 and 43 eggs. 1 1 9 with 1 mass of 14 eggs. i c 10 with 1 mass of 57 eggs. t i 11 with 3 masses of 29, 15 and 14 eggs. l ( 12 with 1 mass of 16 eggs. ( t 13 with 1 mass of 12 eggs. c t 14 with 1 mass of 13 eggs. 1 1 15 with 1 mass of 48 eggs. 1 1 16 with 1 mass of 3 eggs. i i 17 with 1 mass of 48 eggs. 1 1 18 with 1 mass of 10 eggs. l < 19 with 1 mass of 50 eggs. i c 20 with 1 mass of 36 eggs. 1 1 21 with 3 masses of 14, 82 and 14 eggs. In all 883 eggs were found, an average of about 42 eggs to a leaf. My notes show that eggs are most abundant from mid-May until mid- June, but some may be found until nearly the end of August. Comstock (1879) states that living eggs may be found in 76 Volume XXXI dead leaves under the bushes in winter. Adults, however, are known to hibernate and it is doubtful if any eggs survive. The incubation period and other details have not been accu- rately determined. However, in 1950 the first eggs were observed on May 27 and the first nymphs were seen on June 17. Early in the season incubation takes three or four weeks, but the period is probably shortened by warmer weather then or later in the summer. Nymphs: Comstock’s (1879) few remarks concerning the nymphs are also of interest. He compares their dirty brownish color to the color of the material applied over the eggs and the slightly lighter color of the leaves injured by their feeding. Their bodies are broadly oval and flat and spines project from all parts. My field notes record first instar nymphs by early June and various stages may be found more or less abundant until late September. Details of nymphal anatomy and development have yet to be studied for this common species. Adults: The plants on which C. cydoniae (Fitch) is most com- monly found in abundance are all in the family Rosaceae. In this family species of at least six genera serve as the usual hosts. Fitch (1861) based his description of this tingid on specimens he re- ceived from Leominster, Massachusetts. His correspondent re- ported that quince trees ( Cydonia ) were being seriously damaged by heavy infestations of the lace bugs. Locally they have been found abundant and breeding on garden varieties of Chaenomeles, native and cultivated Amelanchier, Crataegus, and Sorbus ( Pyrus ameri- cana (Marsh.) DC. and P. Aucuparia (L.) Gaertn.), and on wild Pyrus melanocarpa (Michx.) Willd. In the Parshley collection there are some with the host label ‘ ‘ elderberry. ” Specimens in the United States National Museum were collected from loquat, Malus, pear, and Pyracantha coccinea Roem. ( fide Sailer). McAtee (1917b and 1923) reports Cephalanthus occidentalis L. (Rubi- aceae) as a host. Bueno (1933) swept them from Quercus. The food plants of C. cydoniae are unusually varied even if we con- sider only the rosaceous kinds as true hosts. My records for 1949 and 1950 show, however, that this species is reasonably selective even amongst rosaceous plants. During those two seasons regular collections were made from a bushy Amelanchier growing at the branch tips of a fairly large Prunus serotina Ehrh. On the same weekly visits C. pruni 0. and D. was collected on the Prunus. Dur- ing two entire seasons (see graphs) only two specimens of C. cy- doniae were taken from the wild cherry — one female in 1949 and 77 ENTOMOLOGICA AMERICANA a single male in 1950. Such evidence strongly suggests that rec- ords are accidental unless breeding populations are present on the plants in question. Adults may be found hibernating in leaves on the ground (Drake, 1928a), under loose bark, and under sticks and stones (Comstock, 1879). They become active by mid-May, somewhat later than C. pruni 0. and D., when the Amelanchier foliage is fairly well expanded in eastern Massachusetts. As suggested above, this species and C. pruni were studied for two full seasons in the field. A station was chosen in the Stony Brook Beservation, Hyde Park, Massachusetts. This reservation is part of the Boston park system and supports a mixed hardwood stand. There are numerous small hills consisting basically of con- glomerate outcrops. A fairly large pond, small streams, and swampy woods provide varied habitat conditions. The site chosen for convenience is near the edge of the reservation on the Hyde Park side. A wood road enters the reservation there. Along the road Prunus serotina Ehrh., Vaccinium, Amelanchier , Bubus and other shrubs and small trees grow. On each side is a low swale supporting a rank growth of herbs and various woody plants. A clump of Prunus serotina , a short distance in, served during both seasons as the collection station for C. pruni 0. and D., since it had a heavy infestation of this tingid. At its branch tips a shrubby Amelanchier infested with C. cydoniae (Fitch) was a most con- venient station for that species. Collections were made regularly at weekly intervals for the two seasons. It was necessary to develop a technique that would give comparable results and not critically deplete the populations of the two tingids. After brief experimentation in the early spring of 1949, it was found that a small vial of alcohol could be easily held under the leaves and the insects flicked into them. Vials with plastic caps proved very satisfactory. They are If inches deep and f of an inch in diameter at the top. Separate vials were used for each species and for each collection. The specimens could then be studied when convenient. The time limit set for each weekly col- lection at each plant was five minutes. At the height of their abundance as many as sixty were taken in the time allowed. This time limit seemed to give a reasonable indication of population density without depleting the populations enough to influence the trends significantly. There is a possibility, however, — that on the smaller Amelanchier supporting, of necessity, a lighter infestation 78 Volume XXXI than the P minus, 1949 collections may, in a small way, partly ac- count for the low numbers of overwintering adults taken in the spring of 1950 (see below). During the 1949 season, temperature, humidity, and light readings were taken on the wood road on each collecting visit. Since it was impossible to go at precisely the same hour each week, and because of rapid and local fluctuations in all three, it was con- cluded such data had no useful significance. The insect popula- tions responded to the broader climatic trends that were reflected by the leaf development of the host plants in spring and the gradual cessation of plant growth in the fall. Reference to Graphs 1 and 2 will reveal interesting differences in the population trends for C. cydoniae (Pitch) during the 1949 and 1950 seasons. In the spring of 1949 there was a large popula- tion of overwintering adults. Many of them were feeding by mid- May and by June 1 the peak was reached. The population then declined rather rapidly and reached its low point between June 15 and 22. By that time teneral adults were appearing and the new generation reached its peak of abundance about July 6. Maturation of this generation took about five weeks, since eggs were first found on May 16 and teneral adults appeared by June 22. The sudden drop in the population by late July suggests some dispersal to other plants and also, perhaps, early hibernation of adults after a short feeding period. By late August very few could be found. The curve for 1950 is very different. Apparently only a relatively small number of adults survived hibernation. This is probably chiefly the result of a ground fire that spread through the underbrush of the area during the period of dormancy. Much of the surface litter was burned, and many tingids were undoubtedly destroyed. The irregularities of the late summer curve reflect primarily the element of chance in the collection method and in small part the vagaries of New England weather and its influence on insect activity. In both 1949 and 1950 it will be clearly seen that the hibernat- ing female population is greater than that of males and that the females emerge earlier. The 1949 brood produced somewhat more males than females and they matured about a week sooner. Ir- regularities in the 1950 curve obscure these relationships. In 1949 a total of 181 males and 178 females were collected. In 1950 the totals were 126 males and 147 females. The males for both seasons 79 ENTOMOLOGICA AMERICANA 1949 POPULATION TRENDS CORYTHUCHA CYDONIAE (FITCH) (&a,y) , 1832 (P. parshleyi 0. and D.) Physatocheila variegata Parshley, 1916b (P. plexa 0. and D., 1916a and 1917b) Leptoypha costata Parshley, 1917a (L. distinguenda Heidemann) Leptoypha ilicis Drake, 1919a Leptoypha mutica (Say), 1832 Dictyonota tricornis Schrank var. americana Parshley, 1916b Acalypta lillianis Bueno, 1916 (A. ovata 0. and D., 1916b; A. grisea Heidemann, 1917, A. modesta Parshley, 1921) Acalypta nyctalis Drake, 1928a Acalypta thomsonii Stal, 1873 ( A . madelinae Bneno) Leptopharsa clitoriae (Heidemann), 1911b (. Leptostyla costofasciata Drake) Leptopharsa heidemanni (Osborn and Drake), 1916a Leptopharsa oblonga (Say), 1825 Galeatus peckhami (Ashmead), 1887 Gargaphia angulata Heidemann, 1899 Gargaphia solani Heidemann, 1914 Gargaphia tiliae (Walsh), 1864 6 The synonyms are enclosed in parentheses. 113 ENTOMOLOGICA AMERICANA Corythaica bellula Bueno, 1917 (0. floridana Blatchley) Stephanitis globulifera (Matsumura), 1905 Stephanitis pyrioides (Scott), 1874 (S. azaleae Horvath) Stephanitis rhododendri Horvath, 1905 (. Leptobyrsa explanata Heidemann) Corythucha arcuata (Say), 1832 ( C . mali Gibson, C. arcuata var. mali Drake) Corythucha associata Osborn and Drake, 1916b ( C . spinulosa Gibson) Corythucha bellula Gibson, 1918 Corythucha caryae sp. nov. Corythucha ciliata (Say), 1832 Corythucha coryli Osborn and Drake, 1917a Corythucha cydoniae (Fitch), 1861 (C. arcuata Comstock, C. arcuata crataegi Morrill, C. era - taegi Osborn and Drake) Corythucha heidemanni Drake, 1918 ( C . borealis Parshley) Corythucha juglandis (Fitch), 1857 (C. contracta 0. and D., C. parshleyi Gibson) Corythucha marmorata (Uhler), 1878 Corythucha marmorata var. inf or mis Parshley, 1919c Corythucha mollicula Osborn and Drake, 1916b (C. salicis 0. and D., C. canadensis Parshley) Corythucha pallipes Parshley, 1918 (C. cyrta Parshley, C. betulae Drake) Corythucha pergandei Heidemann, 1906 Corythucha pruni Osborn and Drake, 1916a Corythucha ulmi Osborn and Drake, 1916a HOST PLANTS OF NEW ENGLAND TINGOIDEA7 Phylum Bryophyta Class Musci Climacium americanum L. and Poly trichum Acalypta lillianis Bueno 7 This list gives only the hosts definitely recorded as food plants. The scientific names of the lace bugs follow their hosts. Starred species are known to breed on the plant indicated. Common names of plants in the literature are disregarded here (though included 114 Volume XXXI Phylum Tracheophyta Class Angiospermae Family Gramineae Panicum *Corythaica bellula Bueno Family Cyperaceae Scirpus atrovirens Willd Piesma cinerea (Say) Family Salicaceae Populus species ^Corythucha mollicula 0. and D. Salix elegantissima Koch Corythucha pallipes Parshley Salix species Corythucha juglandis (Fitch) *C. mollicula 0. and D. C. pergandei Heidemann Physatocheila variegata Parshley Family Juglandaceae Carya illinoensis (Wang.) K. Koch * Corythucha juglandis (Fitch) C. ovata (Mill.) K. Koch Corythucha ciliata (Say) C. tomentosa Nutt Leptoypha costata Parshley Carya species * Corythucha caryae sp. nov. Juglans cinerea L * Corythucha juglandis (Fitch) J. nigra L * Corythucha juglandis (Fitch) J. sieholdiana Maxim * Corythucha juglandis (Fitch) in the earlier discussion) unless the generic reference is assumed to be unmistakable. The plant families follow the order in Gray’s Manual and the binomials of native plants are also those of the new edition (Fernald, 1950). Names of cultivated plants agree with those used by L. H. Bailey (1949). Genera and species are arranged alphabetically for convenience. 115 ENTOMOLOGICA AMERICANA Family Corylaceae Alnus glutinosa (L.) Gaertn .^Corythucha pergandei Heidemann A. incana (L.) Moench Corythucha bellula Gibson # Corythucha pergandei Heidemann A. glutinosa (Du Hoi) Spreng * Corythucha pergandei Heidemann Alnus species * Corythucha heidemanni Drake *0. pergandei Heidemann Leptopharsa clitoriae (Heidemann) L. heidemanni (0. and D.) Betula lutea Michx * Corythucha pallipes Parshley C. pergandei Heidemann B. nigra L * Corythucha pergandei Heidemann B. papyrifera Marsh * Corythucha pallipes Parshley B. populifolia Marsh Corythucha pergandei Heidemann Corylus americana Walt * Corythucha coryli 0. and D. Corylus species * Corythucha bellula Gibson Ostrya virginiana (Mill.) K. Koch Family Fagaceae Fagus grandifolia Ehrh. . Quercus alba L Q. macrocarpa Michx * Corythucha coryli 0. and D. *C. pallipes Parshley * Corythucha pallipes Parshley * Corythucha arcuata (Say) * Corythucha arcuata (Say) 116 Volume XXXI Q. Muehlenbergii Engelm. * Corythucha arcuata (Say) Q. prinoides L Corythucha arcuata (Say) Q. Prinus L ...* Corythucha arcuata (Say) Q. rubra L * Corythucha arcuata (Say) Quercus species ... Corythucha cydoniae (Fitch) Piesma cinerea (Say) Family Ulmaceae Celtis Occident alis L ... Corythucha pergandei Heidemann TJlmus americana L * Corythucha. ulmi 0. and D. JJ. rubra Miilil * Corythucha ulmi 0. and D. V. Thomasi Sarg . ^Corythucha ulmi 0. and D. TJlmus species ... Corythucha juglandis (Fitch) C. pergandei Heidemann Family Moraceae Broussonetia papyrifera L. Corythucha ciliata (Say) Family Chenopodiaceae Beta vulgaris L Chenopodium album L Family Amaranthaceae Amaranthus caudatus L. ... A. hybridus L A. retroflexus L Family Saxifragaceae Bibes oxycanthoides L Piesma cinerea (Say) ...*Piesma cinerea (Say) ...*Piesma cinerea (Say) *Piesma cinerea (Say) *Piesma cinerea (Say) ... Corythucha bellula Gibson Ribes species ... Corythucha mollicula 0. and D. Family Platanaceae Platanus occidentalis L . * Corythucha ciliata (Say) Piesma cinerea (Say) 117 ENTOMOLOGICA AMERICANA Family Rosaceae Amelanchier species * Corythucka cydoniae (Fitch) Chaenomeles species * Corythucka cydoniae (Fitch) Crataegus pruinosa (Wendl.) K. Koch Corythucka bellula Gibson C. punctata Jacq Corythucka bellula Gibson C. succulenta var. neoflu- vialis (Ashe) Palmer ... Corythucka bellula Gibson Crataegus species * Corythucka cydoniae (Fitch) Cydonia oblonga Mill * Corythucka cydoniae (Fitch) Malus species Corythucka cydoniae (Fitch) Prunus americana Marsh. ... Corythucka pergandei Heidemann P. serotina Ehrh # Corythucka associata 0. and D. *C. pruni 0. and D. Pyracantha coccinea Roem. ... Corythucka cydoniae (Fitch) Pyrus ( Sorbus ) americana (Marsh.) D. C * Corythucka cydoniae (Fitch) *C. heidemanni Drake *C. pallipes Parshley P. ( Sorbus ) Aucuparia *C. juglandis (Fitch) (L.) Gaertn * Corythucka cydoniae (Fitch) P. communis L Corythucka cydoniae (Fitch) P. melanocarpa (Michx.) Willd * Corythucka cydoniae Bubus species (Fitch) Corythucka juglandis (Fitch) 118 Volume XXXI Family Leguminosae Amorpha fruticosa L ^Leptopharsa oblonga (Say) Amphicarpa br aetata (L.) Fern ^Leptopharsa oblonga (Say) Baptisia tinctoria (L.) R. Br ^Leptopharsa heidemanni (0. and D.) Clitoria mariana L ^Leptopharsa clitoriae (Heidemann) Desmodium species Leptopharsa clitoriae (Heidemann) Lespedeza species Leptopharsa clitoriae (Heidemann) Petalostemum species Leptopharsa oblonga (Say) Family Aqnifoliaceae Ilex species Leptoypha ilicis Drake Family Aceraeeae Acer pensylvanicum L Corythucha pallipes Parshley A. saccharinum L Corythucha pallipes Parshley A. saccharum Marsh Corythucha pallipes Parshley A. spicatum Lam Corythucha pallipes Parshley Family Hippocastanaceae Aesculus species Piesma cinerea (Say) Family Rhamnaceae Ceanothus americanus L *Gargaphia angidata Heidemann Family Yitaceae Vitis species Piesma cinerea (Say) Family Tiliaceae Tilia species Corythucha juglandis (Fitch) *Gargaphia tiliae (Walsh) Family Ericaceae Chamaedaphne calyculata 119 ENTOMOLOGICA AMERICANA (L.) Moench Kalmia angustifolia L K. lati folia L Pier is floribunda (Pursh) B. and H Pieris japonica (Thunb.) Don Rhododendron calendula- ceum (Michx.) Torr. R. indicum Sweet R. maximum L R. molle Don R. mucronatum Don R. obtusum Planch, var. amoenum Rehd R. ponticum L R. yedoense Maxim Rhododendron species Corythucha ciliata (Say) *Stephanitis rhododendri Horvath *Stephanitis rhododendri Horvath *Stephanitis rhododendri Horvath * Stephanitis rhododendri Horvath *8. globulifera (Matsumura) *8tephanitis pyrioides (Scott) *8tephanitis pyrioides (Scott) *Stephanitis rhododendri Horvath *8tephanitis pyrioides (Scott) *Stephanitis pyrioides (Scott) *Stephanitis pyrioides (Scott) *Stephanitis pyrioides (Scott) *Stephanitis pyrioides (Scott) . Stephanitis globulifera (Matsumura) Family Oleaceae Chionanthus virginicus L. . *Leptoyphamutica (Say) Forestiera acuminata (Michx.) Poir Leptoypha mutica ( Say) Fraxinus caroliniana Mill. ... Leptoypha costata Parshley Fraxinus species Corythucha ciliata (Say) * Leptoypha costata 120 Volume XXXI Family Boraginaceae Lappula species Family Solanaceae Lycopersicum esculentum Mill Solanum carolinense L 8. elaeagnifolium Cay 8. Melongena L 8. tuberosum L Family Rubiaceae Cephalanthus Occident alis L Family Compositae Ambrosia trifida L Aster macrophyllus L Aster species Chrysanthemum mori- folium Ramat. • Eupatorium species Helianthus species Solidago semper virens L. Solidago species Parshley *L. mutica (Say) Leptopharsa clitoriae (Heidemann) . Gargaphia solani Heidemann .* Gargaphia solani Heidemann * Gargaphia solani Heidemann .* Gargaphia solani Heidemann * Gargaphia solani Heidemann Corythucha cydoniae (Fitch) * Corythucha marmorata (Uhler) *Galeatus peckhami (Ashmead) * Corythucha marmorata (Uhler) * Corythucha marmorata (Uhler) *Galeatus peckhami (Ashmead) . Corythucha marmorata (Uhler) * Corythucha marmorata (Uhler) * Corythucha marmorata (Uhler) Melanorhopala clavata Stal 121 ENTOMOLOGICA AMERICANA Tanacetum vulgar e L * Corythucha marmorata (Uhler) CONCLUSIONS AND SUGGESTIONS Although more of the tingids of the New England area have been the subjects of biological studies than in all the other American territories combined, the number of species in this area about which we still know very little remains unduly large. The species in six of the fourteen genera recorded from this part of the country are so poorly known in most cases that we cannot even assign them to definite host plants. Their eggs and nymphal stages have never been described. Even several relatively common and accessible species of Corythucha have been neglected. The most comprehen- sive studies available have by no means provided full details con- cerning the life history and habits of even one species. We have much to learn about the longevity of individuals, the local stability of populations, the mobility of species, and the ecological determi- nants of their distribution. This paper fully reviews the literature and supplies some additional biological data. However, its most significant feature may be the revelation that the study of tingid biology has only been initiated. Therefore, it seems pertinent to suggest some lines of investigation that should prove both fruitful and interesting. We shall confine our remarks primarily to the genus Cory- thucha, with the understanding that they apply, with only minor qualifications, to the less known genera. In a few species, general figures for the time of development are available. However, the time necessary for incubation and for the completion of the various nymphal instars should be more accurately determined. There is reason to think that some females may be impregnated in late sum- mer and go into hibernation without ovipositing. This should be investigated since it could be an important factor in the survival of the species. The ecological factors that influence the local distribution of a species should be carefully studied. The ecology and range of the host plants should also be surveyed and the relationships between the lace bugs and their hosts could then be better evaluated. The vigor of the host plants in relation to the tingid population deserves consideration. By marking large numbers of adults, some knowl- edge of their tendency to move to other plants or to remain on the same host could be acquired. It is rather apparent that species in the genus Corythucha are 122 Volume XXXI closely related, and it is likely that speciation is still taking place with relative rapidity. Attempts to hybridize some of the species now recognized would be extremely interesting. For this purpose a leaf cage has been designed that, with only minor improvements, should prove satisfactory for such experiments. It has been used enough to demonstrate that it is practical to employ this method of keeping the insects under observation under otherwise natural con- ditions. Such breeding experiments might also shed some light on the problems of host specificity and phytophagic speciation. Isola- tion of genetically plastic populations on certain plants has probably contributed significantly to speciation in this genus. Special attention should also be given to predators and para- sites. They may be more important than the sparse data now at hand suggest. Only a few spiders, coccinellid beetles, and predace- ous bugs have been directly reported as predators. Mites alone have been occasionally observed parasitizing lace bugs. It is evi- dent that no great effort has been expended on this aspect of tingid ecology. It is my conviction that further study of the male and female genitalia in Corythucha and other genera will confirm the critical usefulness of these structures in taxonomy. The eggs and nymphal instars should also be studied comparatively for specific characters. From the morphological point of view, the Tingidae are especially interesting. These are but a few ideas holding much promise for future investigators. Selected References Abbott, Cyril E. 1935. Notes on the Oviposition and Hatching of Corythucha marmorata (Uhler). Brooklyn Entomological Society , Bulletin, 30: 13. Amyot, C. J. B. and J. C. Audinet Serville. 1843. Histoire Nat- urelle des Insectes, Hemipteres. Librairie Encyclopedique de Roret, p. 300, Paris. Ashmead, W. H. 1886. On Two New Hemiptera Heteroptera. Canadian Entomologist, 18 (1) : 18-20. 1887. Hemipterological Contributions. Tingitidae. Sphaero- cysta Stal. Entomologica Americana, 3 : 156. Bailey, L. H. 1949. Manual of Cultivated Plants. Macmillan, New York. Banks, Nathan. 1910. Catalogue of the Nearctic Hemiptera- Heteroptera. American Entomological Society, Phila- delphia. 123 ENTOMOLOGICA AMERICANA Barber, George W. 1924. Notes on Piesma cinerea Say. Psyche, 31 (5) : 229-232, Figs. 1 and 2. Barber, H. G. and H. B. Weiss. 1922. The Lace Bugs of New Jersey. New Jersey Department of Agriculture, Circular 54 : 1-24. Blatchley, W. S. 1926. Heteroptera of Eastern North America. Nature Publishing Company, Indianapolis. 1928a. “Quit-Claim” Specialists vs. the Making of Manuals. Brooklyn Entomological Society, Bulletin, 23 (1) : 10-18. 1928b. Notes on the Heteroptera of Eastern North America with Descriptions of New Species. New York Entomolo- gical Society, Journal, 36 (1) : 1-23. Bruner, L. 1891. A List of Insects Affecting Sugar Beets. Nebraska State Board of Agriculture, Experiment Sta- tion Bulletin, 4 (16) : 66-67. 1895. Insect Enemies of the Grape. Nebraska State Horti- cultural Report, pp. 68-162. Bueno, J. R. de la Torre. 1915. Heteroptera in Beach Drift. Entomological News, 26 (6) : 278. 1916. A New Tingitid from New York State. Brooklyn En- tomological Society, Bulletin, 11 (2) : 39-40. 1917. A New Species of Tingid from New York. Brooklyn Entomological Society, Bulletin, 12 (1) : 19-20. 1924a. On a Few Heteroptera from Massachusetts. Brooklyn Entomological Society, Bidletin, 19 (2) : 48-51. 1924b. A Correction in Acalypta. Brooklyn Entomological Society, Bulletin , 19 (3) : 93. 1924c. Gaditanus, Being Additional Words on Tingitidae. Entomological News, 35 (9) : 333-334. 1925. Food plant of Corythucha marmorata (Uhler). Brook- lyn Entomological Society, Bulletin, 20 (4) : 179. 1926. Some Remarks, A1 Yuelo, on Tingitid Names. Brooklyn Entomological Society, Bulletin, 21 (3) : 116-117. 1931. Alveotingis grossocerata. Brooklyn Entomological So- ciety, Bulletin, 26 (3) : 149. 1933. New Records of Heteroptera from Arkansas. Brooklyn Entomological Society, Bulletin, 28 (5) : 228. 1942. Maternal Solicitude in Gargaphia iridescens Champion. Brooklyn Entomological Society, Bulletin, 37 (4) : 131. 1945. Random Notes on Heteroptera. Brooklyn Entomologi- cal Society, Bulletin, 40 (3) : 68. 124 Volume XXXI 1946. On Hesperotingis antennata Parshley. Brooklyn En- tomological Society , Bulletin, 41 (3) : 94-95. Butler, E. A. 1923. A Biology of the British Hemiptera-Heterop- tera. H. F. and G. Witherby, London. Chittenden, F. H. 1900. A new Tingitid on Bean. U.S.D.A. Division of Entomology, Bulletin, 23 (n.s.) : 32-33. Comstock, J. H. 1879. The Hawthorn Tingis. Report of the Entomologist of the United States Department of Agri- culture, pp. 221-222.' Cotton, R. T. 1917. The Eggplant Lace-bng in Porto Rico. Porto Rico Department of Agriculture, Journal, 1 (3) : 170-173. Crosby, C. R. and C. H. Hadley, Jr. 1915. The Rhododendron Lace-bng, Leptohyrsa explanata Heidemann. Journal of Economic Entomology, 8 (4) : 409-4*14. Curtis, John. 1827. British Entomology. Vol. 7, item number 154, London. 1833. Characters of Some Undescribed genera and species. Entomological Magazine 1 : 196. Dickerson, E. L. 1917. Notes on Leptobyrsa rhododendri Hor- vath. New York Entomological Society, Journal, 25 (2) : 105-112, PL 8. Dickerson, E. L. and H. B. Weiss. 1916. Notes on Leptoypha mutica (Say). Entomological News, 27: 308-310, PI. 16. 1917. The Azalea Lace-bug, Stephanitis pyrioides Scott. En- tomological News, 28 : 101-105, PI. 9. 1918. Corythucha spinulosa Gibson, a New Lace-bug on Wild Cherry. Entomological News, 29: 121-125, PL 7. Downes, W. 1927. A New Species of Drakella. Canadian En- tomologist, 59 : 60. Drake, C. J. 1916. A New Tingid from Tennessee. Ohio Journal of Science, 16 (7) : 326-328. 1917. Key to Nearctic Species of Gargaphia with the Descrip- tion of a New Species. Entomological News, 28: 227-228. 1918. Notes on North American Tingidae. Brooklyn Ento- mological Society, Bulletin, 13 (4) : 86-88. 1919a. On Some North American Tingidae. Ohio Journal of Science, 19 (7) : 417-421. 1919b. On Some Tingidae New to the Fauna of Canada. ' Canadian Entomologist, 51 (6-7) : 159-160. 1921. Notes on Some American Tingidae with Descriptions of New Species. Florida Entomologist, 4 (4) : 49-54. 125 ENTOMOLOGICA AMERICANA 1922a. On Some North and South American Tingidae. Flor- ida Entomologist, 5 (3) : 37-43, 48-50. 1922b. Heteroptera in the Vicinity of Cranberry Lake, Tin- gidae. New York State College of Forestry, Syracuse Uni- versity, Technical Publication 16, vol. 22 (5) : 64-66. 1922c. Contribution toward the Life History of Galeatus peckhami (Ashmead). New York State College of For- estry, Syracuse University, Technical Publication 16, 22 (5) : 105-110, PI. 4. 1922d. The Life History of the Birch Tingitid, Corythucha pallipes Parshley. New York State College of Forestry, Syracuse University, Technical Publication 16, 22 (5) : 111-116. 1923. Some Tingitidae from Japan. Ohio Journal of Science, 23 (2) : 102-106. 1925. Concerning Some Tingitidae from the Gulf States. Florida Entomologist, 9 (3) : 36-39. 1926. The North American Tingitidae Described by Stal. Carnegie Museum, Annals, 16 (3-4) : 375-380, PI. 34. 1928a. A List of the Insects of New York. Families Piesmidae and Tingitidae. Cornell University Agricultural Experi- ment Station, Memoir 101 : 99-103. 1928b. A Synopsis of the American Species of Acalypta. Brooklyn Entomological Society, Bulletin, 23 (1) : 1-9. 1928c. Synonymical Notes on Tingitid Genera with the Des- criptions of Two New Species from Haiti. Biological So- ciety of Washington, Proceedings, 41 : 21-23. 1928d. Four Undescribed Tingitids from United States. Florida Entomologist, 12 (1) : 3-5. 1930. Notes on American Tingitidae. Brooklyn Entomologi- cal Society, Bulletin, 25 (5) : 268-272. 1948. New Species of Stephanitis Stal Including a List of Species of the World. Musee Heude, Notes d’Entomologie Chinoise, 12 (6) : 45-56. Drake, C. J. and M. E. Poor. 1936. The Genera and Genotypes of the Tingitoidea of the Western Hemisphere. Iowa State College Journal of Science, 10 (4) : 381-390. Felt, E. P. 1903. Injurious Insects. Corythucha marmorata (Uhler). New York State Museum, Bidletin, 76: 125-129. 1933. Observations on Shade Tree Insects. Journal of Eco- nomic Entomology, 26 (1) : 49. 126 Volume XXXI Fernald, M. L. 1950. Gray’s Manual of Botany (Eighth Edi- tion). American Book Company, Boston. Fieber, F. X. 1844. Entomologische Monographien. Mono- graphic der Tingideae, 20-111. Johann Ambrosius Barth, Leipzig. 1861. Die Europaischen Hemiptera. Tingididae, 116-132, Carl Gerolds Sohn, Vienna. Fink, David E. 1915. The Eggplant Lace-bug. TJ. 8. Department of Agriculture, Bulletin, 237 : 1-7. Fitch, Asa. 1857. Third Report on the Noxious Insects of New York. Item 193. Butternut Tingis. pp. 148-149. Al- bany. 1858. Fourth Report on the Noxious and Other Insects of the State of New York. N. Y. State Agricultural Society , Transactions, 17 : 687-814. 1861. The Quince Tingis. Country Gentleman, 17 (25) : 114. Froeschner, Richard C. 1944. Contributions to a Synopsis of the Hemiptera of Missouri, Part III. American Midland Nat- uralist, 31 (3) : 638-683. Gibson, A. 1904. Basswood, or Linden Insects. Entomological Society of Ontario. 34th Annual Report: 52. Gibson, E. H. 1918. The Genus Corythucha Stal. American En- tomological Society, Transactions, 44 (775) : 69-104. 1919a. The Genera Corythaica Stal and Dolichocysta Champion. Biological Society of Washington, Proceed- ings, 32 : 97-104. 1919b. The Genus Gargaphia Stal. American Entomological Society, Transactions, 45 (789) : 187-201. Goldi, Emil A. 1886. Beitrage zur Kenntniss der kleinen and kleinsten Gliederthierwelt Brasiliens. Mittheilungen der Schweizer. Entomologischen Gesellschaft, 7 (6) : 233-241. Heidemann, Otto. 1899. A New Species of Tingitidae. Canadian Entomologist, 31: 301-302. 1906. Account of a New Tingitid. Entomological Society of Washington, Proceedings, 8 (1-2) : 10-13, figs. 2 and 3. 1908. Two New Species of North American Tingitidae. En- tomological Society of Washington, Proceedings, 10 (1-2) : 103-108, PL 4. 1911a. Some Remarks on the Eggs of North American Species of Hemiptera — Heteroptera. Entomological Society of Washington, Proceedings, 13 (2) : 136-137. 127 ENTOMOLOGICA AMERICANA 1911b. A New Species of North American Tingitidae. En- tomological Society of Washington, Proceedings, 13 (3) : 180-181, Fig. 4. 1914. A New Species of North American Tingitidae. En- tomological Society of Washington, Proceedings, 16 (3) : 136-137. 1917. Two New Species of Lace-Bugs. Entomological Society of Washington, Proceedings , 18 (4) : 217-220, pi. 17. Hickman, Dorothy J. 1921. Illustrations of the Male Hooks in Nobis. Brooklyn Entomological Society, Bulletin, 16: 58-59. Horvath, Geza. 1905. Tingitidae Novae vel Minus Cognitae e Regione Palaearctica. Musei Nationalis Hung arid, An- nates, 3: 567-568. 1906. Synopsis Tingitidarum Regionis Palaearcticae Musei Nationalis Hungarici. Separata ex Annalibus historico- naturalibus, 4: 1-118. 1912. Species Generis Tingitidarum Stephanitis. Annales Musei Nationalis Hungarici, 10: 319-329. 1923. A New Species of Galeatus from New Mexico. Carnegie Museum, Annals, 15 (1) : 108-109. Hurd, Margaret P. 1945. A Monograph of the Genus Corythaica Stal. Iowa State College Journal of Science, 20 (1) : 79- 99, PI. 1, Figs. 1-5. 1946. Generic Classification of North American Tingoidea. Iowa State College Journal of Science, 20 (4) : 429-489. Hussey, Roland F. 1922a. On Some Hemiptera from North Dakota. University of Michigan Museum of Zoology, Oc- casional Papers, 115: 1-23. 1922b. Hemiptera from Berrien County, Michigan. Univer- sity of Michigan Museum of Zoology, Occasional Papers, 118: 1-39. Laporte de Castelnau, F. L. 1832. Essai d’une Classification Systematique de l’Ordre des Hemipteres. Magasin Zoo- logique, 1, Nos. 52-55, Supplement. ( fide Blatchley). Leonard, M. D. and A. S. Mills. 1931. Observations on the Bean Lace-bug in Porto Rico. Porto Rico Department of Agri- culture, Journal, 15: 309-323. Le Peletier de Saint Fargeau, A. L. M. and J. G. Audinet-Serville. 1825-1828. Encyclopedic Methodique, 10, Entomologie, Paris. ( fide Blatchley). 128 Volume XXXI Lethierry, L. and G. Severin. 1896. Catalogue General des Hemipteres. Tome III, Heteropteres-Tingidae, 1-26. R. Friedlander et Fils, Berlin. Matsumura, S. 1930. The Illustrated Thousand Insects of Japan. Vol. 1. Rhynchota. Toko-Shoin, Tokyo. McAtee, W. L. 1912. Note on the Hibernation of Corythucha and Milyas. Entomological Society of Washington, Proceedings, 14: 102. 1917a. Key to the Nearctic Species of Leptoypha and Lepto- styla. Brooklyn Entomological Society, Bulletin, 12 (3) : 55-64. 1917b. A Few Notes Chiefly on the Names of Nearctic Tingi- dae. Brooklyn Entomological Society, Bulletin, 12 (4) : 78-79. 1919a. Key to Nearctic Species of Piesmidae. Brooklyn En- tomological Society, Bulletin, 14: 80-93, fig. 1919b. Corrections and Additions to an Article on Leptoypha and Leptostyla. Brooklyn Entomological Society, Bulletin, 14 (4-5) : 142-144. 1923. Tingitoidea of the Vicinity of Washington, D. C. En- tomological Society of Washington, Proceedings, 25 (7-8) : 143-151. Monte, Oscar. 1940. Catalogo dos Tingitideos do Brasil. Arqui- vos de Zoologia do Estado de Sdo Paulo. Revista do Museu Paulista, 2 (3) : 65-174. Morrill, Austin W. 1903. Notes on the Immature Stages of Some Tingitids in the Genus Corythucha. Psyche, 10 (324) : 127-134, plate 3. Myers, J. G. 1926. Heteroptera in Ocean Drift. Psyche, 33 (4-5) : 110-115. Olsen, C. E. 1923. Distributional Notes on Hemiptera. Brooklyn Entomological Society, Bulletin, 18: 164. Osborn, H. and C. J. Drake. 1916a. The Tingitoidea of Ohio. Ohio State University Bidletin, 20 (35) : 217-251. 1916b. Some New Species of Nearctic Tingidae. Ohio Journal of Science, 17 (1) : 9-15. 1917a. Notes on American Tingidae with Descriptions of New Species. Ohio Journal of Science, 17 (8) : 295-307. fig. 2a. 1917b. Notes on Tingidae. Psyche, 24 (5) : 155-161. Osborn, H. and H. A. Gossard. 1891. Reports on Injurious In- sects. Iowa Agricultural Experiment Station, Bulletin, 15: 255-273. 129 ENTOMOLOGICA AMERICANA Parshley, H. M. 1914. List of the Hemiptera-Heteroptera of Maine. Psyche, 21 (5) : 139-149. 1915. Hemiptera-Heteroptera of Maine. Corrections and Ad- ditions. Psyche, 22: 22-23. 1916a. New and Noteworthy Hemiptera from New England. Entomological News, 27 (3) : 103-106. 1916b. On Some Tingidae from New England. Psyche, 23 (6) : 163-168. 1917a. Notes on North American Tingidae. Psyche, 24 (1) : 13-25. 1917b. Insects in Ocean Drift. Canadian Entomologist , 49 (2) : 45-48. 1917c. Fauna of New England, 14. List of the Hemiptera- Heteroptera. Boston Society of Natural History, Oc- casional Papers, 7 : 1-125. 1919a. Note on the Sexes of the Tingid Melanorhopala clavata Stal. Brooklyn Entomological Society, Bulletin, 14 (3) : 102-103. 1919b. A Morphological Note on the Tingoidea. Brooklyn En- tomological Society, Bulletin, 14 (3) : 109-110. 1919c. On Some Hemiptera from Western Canada. Univer- sity of Michigan Museum of Zoology, Occasional Papers, 71:1-35. 1919d. Note of Rectification. Brooklyn Entomological So- ciety, Bulletin, 14 (4-5) : 148. 1920a. Hemiptera from Peaks Island, Maine. Canadian En- tomologist, 52 : 80-87. 1920b. Hemiptera Collected in Western New England. Psy- che, 27 (6) : 139-143. 1920c. Hemipterological Notices I. (Tingidae). Entomologi- cal News, 31 (10) : 271-273. 1921. A Report on some Hemiptera from British Columbia. British Columbia Entomological Society, Proceedings, 18: 13-24. 1922a. Report on a Collection of Henliptera-Heteroptera from South Dakota. South Dakota State College, Technical Bulletin, Number 2: 1-22. 1922b. New England Hemiptera-Heteroptera. New Records II. Canadian Entomologist, 53 : 233-239. 1922c. Tingitidae or Tingidae. Science, 56 (1451) : 449. 1922d. On the Formation of Family Names like Tingidae. Science , 56 (1461): 754-755. 130 Volume XXXI 1923a. Records of Nova Scotian Hemiptera-Heteroptera. Acadian Entomological Society , Proceedings, 8 : 102-108. 1923b. Families Piesmidae and Tingidae in the Hemiptera of Connecticut. State of Connecticut Geological and Natural History Survey, Bulletin, 34: 694-707. 1925. A Bibliography of the North American Hemiptera- Heteroptera. Smith College, Northampton, Massachusetts. Pemberton, C. 1911. The California Christmas-berry Tingis. Journal of Economic Entomology , 4 : 339-343, Pis. 12-14. Proctor, William. 1946. Insect Fauna of Mt. Desert (Part 7), pp. 74-75. Wistar Institute of Anatomy and Biology, Philadelphia. Provancher, 1’Abbe L. 1886. Petite Faune Entomologique du Canada. Vol. 3. Les Hemip teres. C. Darveau, Quebec. Reuter, O. M. 1912. Bemerkungen liber mein neues Heterop- terensystem. Ofversigt Finska Vetenskaps-Societetens F dr- handling ar, 54 (A, 6) : 49. Robinson, B. L. and M. L. Fernald. 1908. Gray’s New Manual of Botany (Seventh Edition). American Book Company, New York. Sailer, R. I. 1945. The Bite of a Lacebug. Kansas Entomologi- cal Society, Journal, 18 (2) : 81. Say, Thomas. 1825. Descriptions of new Hemipterous Insects collected in the Expedition to the Rocky Mountains. Phila- delphia Academy of Natural Sciences, Journal, 4 (2) : 307-344. 1832. Descriptions of New Species of Heteropterous Hemip- tera of North America. New Harmony, Indiana. (See: — New York State Agricultural Society, Transactions, 17 : 755-812, 1858). 1859. The Complete Writings of Thomas Say on the Ento- mology of North America. Edited by John L. LeConte. Bailliere Brothers, New York. Scott, John. 1874. On a Collection of Hemiptera Heteroptera from Japan. Annals and Magazine of Natural History, 14 (4) : 440-441. Singh-Pruthi, Hem. 1925. The Morphology of the Male Genitalia in Rhynchota. Entomological Society of London, Trans- actions, (1-2) : 127-267. Snodgrass, R. E. 1935. Principles of Insect Morphology. Mc- Graw-Hill, New York. Somes, M. P. 1916. Some Insects of Solanum carolinense L. and 131 ENTOMOLOGICA AMERICANA their Economic Relations. Journal of Economic Entomol- ogy, 9 (1) : 39-41. Stal, Carlos. 1862. Hemiptera Mexicana. Stettin. Entomolo- gische Zeitung, 23: 324-325. 1873. Ennmeratio Hemipterorum. 11, No. 2, (3) : 115-134. Stockholm. Summers, H. E. 1891. The True Bugs, or Heteroptera, of Ten- nessee. Tennessee Agricultural Experiment Station Bulletin, 4 (3) : 90. Tilden, J. W. 1950. Biological Notes on Corythucha morrilli 0. and D. Entomological News, 61 : 135-137. Twinn, C. R. 1935. A Summary of Insect Conditions in Canada. Entomological Society of Ontario, 65th Annual Report, p. 126. Uhler, P. R. 1878. Notices of Hemiptera Heteroptera in the Col- lection of the late T. W. Harris, M.D. Boston Society of Natural History, Proceedings, 19: 415-416. 1886. Check-list of the Hemiptera Heteroptera of North Amer- ica. Brooklyn Entomological Society. 1896. Summary of the Hemiptera of Japan presented to the United States National Museum by Professor Mitzukuri. United States National Museum, Proceedings, 19: 265. 1904. List of Hemiptera-Heteroptera of Las Vegas Hot Springs, New Mexico. United States National Museum, Proceedings, 27 : 362. Van Duzee, E. P. 1889. Hemiptera of Muskoka Lake District. Canadian Entomologist, 21 (1) : 5. 1917a. Catalogue of the Hemiptera of America North of Mex- ico. University of California Press, Berkeley. 1917b. Report upon a collection of Hemiptera made by W. M. Gifford. California Academy of Sciences, Proceedings* Series 4, 7 (11) : 257-261. Walley, G. S. 1928. Preliminary Study of Male Genital Arma- ture of the North and South American Genera of Tingi- tidae. (unpublished M.S. thesis in Iowa State College library) . Walsh, B. D. 1864. On Phytophagic Varieties and Phytophagic Species. Entomological Society of Philadelphia, Proceed- ings, 3 (408) : 408-409. Walsh, B. D. and C. V. Riley. 1868. Bugs on grapevine mistaken for chinch bugs. Leaf -Bugs, etc. American Entomologist, 1: 19. 132 Volume XXXI Weiss, H. B. 1913. Notes on Negative Geotropism of Corythucha ciliata (Say). Journal of Economic Entomology, 6: 407- 409. 1915. Insect Importations into New Jersey during the Spring of 1915. Canadian Entomologist, 47 (9) : 326-328. 1916a. Foreign Pests Recently Established in New Jersey. Journal of Economic Entomology, 9: 212-216. 1916b. Newark Entomological Society (Minutes). Ento- mological News, 27 (4) : 189-190. 1918a. The Control of Imported Pests Recently Found in New Jersey. Journal of Economic Entomology, 11 (1) : 122-125. 1918b. Some New Insect Enemies of Greenhouse and Orna- mental plants in New Jersey. New Jersey Agricultural Experiment Stations, Circular 100: 1-19. 1919. Notes on Gargaphia tiliae Walsh, the Linden Lace Bug. Biological Society of Washington, Proceedings, 32: 165— 168. 1921. Notes on the Life History and Early Stages of Cory- thucha celtidis O. and D. Ohio Journal of Science, 21 (3) : 104-106. 1924. Corythucha marmorata (Uhler) on Seaside Goldenrod. Entomological News, 35: 367. Weiss, H. B. and E. L. Dickerson. 1918a. The Early Stages of Corythucha pergandei Heidemann. Entomological News, 29 : 205-209. 1918b. The Life-History and Early Stages of Corythucha parshleyi Gibson. Canadian Entomologist, 50 (12) : 401- 406. Weiss, H. B. and R. B. Lott. 1924a. Notes on Corythucha mar- morata (Uhler) in New Jersey. Entomological News, 35 (2): 68. 1924b. Notes on Piesma cinerea (Say) in New Jersey. Psy- che, 31 (5) : 233-235. Weiss, H. B. and Erdman West. 1924. Notes on the False In- digo Lace Bug. Gelchossa heidemanni Osborn and Drake, in New Jersey. Entomological News, 35: 56-60. Westwood, J. O. 1840. An Introduction to the Modern Classifica- tion of Insects. 2 vols. London. White, R. P. 1933. The Insects and Diseases of Rhododendron and Azalea. Journal of Economic Entomology, 26: 631— 632. 133 ENTOMOLOGXCA AMERICANA Wille, Joh. 1929. Die Riibenblattwanze Piesma quadrata Fieber. Monograpkien zum P flanz ens chut z, 2 : 1-116. J. Springer, Berlin. Index Valid genera and species in Roman; synonyms in Italics ; new name of species and main page references in bold face; # names of plants. Acalypta, 10, 12, 32, 34, 113, 114 *Acer, 50, 93, 119 *Aceraceae, 119 #acre, Sedum, 31 ^acuminata, Adelia, 30 #Forestiera, 30, 120 *Adelia, 30 *Aesculus, 119 *alba, Betula, 93 *Hicoria, 27 *Quercus, 63, 116 * album, Chenop odium, 18, 117 *Alnus, 68, 82, 83, 93, 94, 95, 116 Alveotingis, 9, 20, 113 ^Amaranthaceae, 19, 117 ^Amaranthus, 17, 18, 19, 117 # Ambrosia, 88, 89, 121 #Amelanchier, 76, 77, 78, 86, 101 #americana, Corylus, 74, 84, 116 Dictyonota tricornis var., 10, 30, 31, 113 5X:Prunus, 95, 118 *Pyrus, 77, 93, 118 •Tilia, 91 #XJlmus, 82, 106, 117 ^americanum, Climacium, 34, 114 ^americanus, Ceanothus, 42, 119 *amoena, Azalea, 56 *amoenum, Rhododendron ob- tusum var., 56, 120 *Amorpha, 39, 119 ^Amphiachrus, 44 *Amphiachyris, 44 *Amphicarpa, 39, 119 #Andropogon, 23 ^Angiospermae, 115 angulata, Gargaphia, 13, 42, 43, 113, 119 #angustifolia, Kalmia, 59, 120 antennata, Hesperotingis, 12, 22, 23, 113 Anthocoris, 93 ^Aquifoliaceae, 119 *Aralia, 41 arcuata, Corythucha, 14, 63, 64, 65, 68, 73, 88, 94, 105, 111 (Fig. 5A), 114, 116, 117 Corythucha, 74, 75, 114 Ting is, 63 * Artemisia, 31 associata, Corythucha, 15, 66, 67, 73, 96, 108, 111 (Fig. 5B), 114, 118 * Aster, 40, 89, 121 *atrovirens, Scirpus, 18, 115 #Aucuparia, Pyrus, 77, 118 * Azalea, 56 azaleae, Stephanitis, 56, 114 #azurea, Salvia, 45 *Baptisia, 38, 119 bellula, Corythaica, 10, 48, 49, 51 52 (Fig. 2), 63, 114, 115, 117 134 Volume XXXI Corythucha, 16, 67, 73, 74, 84, 111 (Fig. 5C), 114, 116, 118 #Beta, 117 *Betula, 50, 93, 95, 116 betulae, Corythucha, 91, 114 #Boraginaceae, 36, 121 borealis, Anthocoris, 93 Corythucha, 82, 114 Hesperotingis antennata var., 22, 113 *bracteata, Amphicarpa, 39, 119 breyirostris, Physatocheila, 12, 24, 25, 113 *Broussonetia, 72, 117 *Bryophyta, 114 ^calendulaceum, Rhododendron, 56, 120 #calyculata, Chamaedaphne, 119 canadensis, Corythucha, 90, 114 Cantacader, 21 #carolinense, Solannm, 44, 45, 121 *caroliniana, Fraxinus, 27, 120 *Carya, 27, 71, 72, 86, 115 caryae, Corythucha, 14, 65, 68, 70 (Fig. 3), 73, 105 111 (Fig. 5D), 114, 115 * Cassia, 45 ^caudatns, Amaranthus, 18, 117 *Ceanothus, 42, 119 #Celtis, 95, 117 *Cephalanthus, 77, 121 #Chaenomeles, 77, 118 * Chamaedaphne, 72, 119 #Chenopodiaceae, 18, 19, 117 #Chenopodium, 18, 117 *Chionanthus, 29, 30, 120 Chiracanthinm, 45 * Chrysanthemum, 88, 89, 121 Chrysopa, 93 ciliata, Corythucha, 14, 65, 72, 105,111 (Fig. 5E), 114, 115, 117, 120 Tingis, 72 *cinerea, Juglans, 86, 115 Piesma, 9, 16, 17, 18, 73, 113, 115, 117, 119 Tingis, 16 clavata, Melanorhopala, 9, 21, 113, 121 *Climacium, 34, 114 #Clitoria, 36, 119 clitoriae, Leptopharsa, 13, 36, 37, 38, 113, 116, 119, 121 Leptostyla, 36 #coccinea, Pyracantha, 77, 118 *Quercus, 64 ^communis, Pyrns, 118 *comosa, Falcata, 39 #Compositae, 45, 89, 121 #Comptonia, 50 #Conocarpus, 18 contracta, Corythucha, 84, 114 convergens, Ilippodamia, 44, 45 *Cornus, 93 *Corylaceae, 68, 74, 82, 116 coryli, Corythucha, 16, 67, 73, 74, 75, 84, 111 (Fig. 5F), 114, 116 #Corylus, 68, 74, 84, 116 #corymbosum, Vaccinium, 50 Corythaica, 3, 10, 42, 47, 48, 53, 63, 115 Corythuca, 63 Corythucha, 3, 5, 10, 11 (Fig. 1), 14, 42, 47, 53, 62, 65, 67, 75, 83, 85, 89, 91, 92, 96, 105, 106, 107, 108, 110, 111 (Fig. 5), 112 (Fig. 6) 115, 116, 117, 118, 119, 120, 121, 122, 123 135 ENTOMOLOGICA AMERICANA costata, Leptoypha, 12, 27, 28, 113, 115, 120 costofasciata, Leptostyla , 36, 113 ^crassifolia, Celtis, 95 crataegi, Corythucha, 74, 114 Corythucha arcuata, 74, 114 ^Crataegus, 68, 75, 77, 118 ^Cydonia, 77, 118 cydoniae, Corythucha, 16, 73, 74, 75, 76, 77, 78, 79, 80 (Graph 1), 81 (Graph 2), 97, 101, 111 (Fig. 5G), 114, 117, 118, 121 Tingis, 74 #Cyperaceae, 115 cyrta, Corythucha , 91, 92, 114 ^Cytisus, 31 ^Danthonia, 50 decens, Corythucha, 86 *Desmodium, 36, 119 Dictyonota, 10, 30, 32, 113 distinguenda, Leptoypha, 27, 113 domiciliorum, Epeira, 45 Drakella, 32 #elaeagnifolium, Solanum, 44, 121 ^elegantissima, Salix, 93, 115 Epeira, 45 #erecta, Conocarpus, 18 ^Ericaceae, 54, 59, 119 Erythraeidae, 71 ^esculentum, Lycopersicum, 44, 121 *Eupatorium, 40, 121 explanata, Leptobyrsa, 58, 114 *Fagaceae, 116 *Fagus, 93, 116 *Falcata, 39 Fenestrella, 32 flavus, A., 31 #floribunda, Pieris, 59, 120 floridana, Corythaica, 48, 114 #Forestiera, 30, 120 *Fraxinus, 27, 29, 72, 120 #fruticosa, Amorpha, 39, 119 #fulva, Ulmus, 83, 106 Galeatus, 10, 39, 40, 41, 113, 121 Gargaphia, 10, 13, 42, 43, 53, 113, 119, 121 Gelchossa, 35 globulifera, Stephanitis, 14, 53, 54, 55, 56, 114, 120 Tingis, 53 *glutinosa, Alnus, 95, 116 #Gossypium, 45 ^Gramineae, 115 #grandifolia, Fagus, 93, 116 grisea, Acalypta, 32, 113 grossocerata, Alveotingis, 9, 20, 113 *Gutierrezia, 45 heidemanni, Corythucha, 15, 82, 83, 84, 94, 110, 111 (Fig. 5H), 114, 116, 118 Leptopharsa, 13, 37, 38, 39, 113, 116, 119 Leptostyla, 37 *Helianthus, 89, 121 henshawi, Cantacader, 21, 113 *herbaceum, Gossypium, 45 Hesperotingis, 9, 11 (Fig. 1), 12, 22, 24, 113 hewitti, Corythucha, 67, 84 *FIicoria, 27 ^Hippocastanaceae, 119 136 Volume XXXI Hippodamia, 44, 45 #hybridus, Amaranthus, 18, 117 •Ilex, 28, 119 Mlicifolia, Quercus, 64 ilicis, Leptoypha, 12, 27, 28, 113, 119 *illinoensis, Carya, 86, 115 illinoiensis, Plesperotingis, 12, 24, 113 #incana, Alnus, 68, 95, 116 inclusnm, Chiracanthium, 45 •indica, Azalea, 56 ^indicum, Rhododendron, 56, 120 informis, Corythucha marmo- rata var., 15, 86, 87, 112 (Fig. 6C), 114 infuscata, Melanorhopala, 22 inornata, Piesma cinerea var., 16, 17, 113 insidiosus, Orins, 45 Triphleps, 45 iridescens, Gargaphia, 44 #japonica, Pieris, 54, 59, 120 #Juglandaceae, 115 juglapdis, Corythucha, 15, 82, 83, 84, 85, 86, 107, 110, 112 (Fig. 6A), 114, 115, 118 Tingis, 84 *Juglans, 82, 86, 115 *Juniperus, 50 *Kalmia, 50, 54, 58, 120 *Kuhnistera, 39 *Labiatae, 45 *Lappula, 36, 121 Matifolia, Kalmia, 50, 58, 120 Maurifolia, Quercus, 64 *Leguminosae, 45, 119 *lenta, Betula, 93 Leptobyrsa, 58 Leptopharsa, 10, 11 (Fig. 1), 13, 35, 113, 116, 119, 121 Leptostyla, 35, 36, 37 Leptoypha, 9, 12, 27, 113, 115, 119, 120, 121 Leptus, 71 #Lespedeza, 36, 119 lillianis, Acalypta, 13, 32, 34, 113, 114 #Lindheimeri, Panicum, 48 #Liriodendron, 22 lurida , Tingis , 21, 113 *lutea, Betula, 93, 95, 116 #Lycopersicum, 44, 121 #macrocarpa, Quercus, 64, 116 #macrophyllus, Aster, 40, 121 maculata, Megilla, 45 maculiventris, Podisus, 45 madelinae , Acalypta, 35, 113 mali, Corythucha, 63, 114 Corythucha arcuata var., 63, 114 *Malus, 77, 118 ^Malvaceae, 45 #mariana, Clitoria, 36, 119 marmorata, Corythucha, 15, 67, 86, 87, 88, 89, 111 (Fig. 6B) , 114, 121, 122 Tingis, 86 ^maximum, Rhododendron, 59, 120 Megilla, 45 #Meibomia, 36 ^melanocarpa, Pyrus, 77, 118 Melanorhopala, 9, 20, 21, 22, 24, 113, 121 *Melongena, Solanum, 44, 121 137 ENTOMOLOGICA AMERICANA #Mentha, 31 Microdus, 45 modesta, Acalypta, 32, 113 *molle, Rhododendron, 57, 120 mollicnla, Corythucha, 15, 67, 90, 101, 112 (Fig. 6D), 114, 115 monacha, Corythaica, 51 *Moraceae, 117 #morifolram, Chrysanthemum, 121 *Morus, 91 ^mucronatum, Rhododendron, 57, 120 ^Muehlenbergii, Quercus, 63, 117 *Musci, 114 mutica, Leptoypha, 12, 27, 28, 29, 113, 120, 121 Myrmica, 31 neofluvialis, Crataegus succu- lenta var., 68, 118 niger, A., 31 *nigra, Betula, 95, 116 *Juglans, 86, 115 nyctalis, Acalypta, 13, 34, 113 oblonga, Cydonia, 118 Leptopharsa, 13, 37, 39, 113, 119 Tingis, 39 ohscura, Melanorhopala, 21, 113 ^obtusum, Rhododendron, 56, 120 ^occidentals, Celtis, 95, 117 *Cephalanthus, 77, 121 #Platanus, 117 ^Oleaceae, 120 Orins, 45 Ortkosteira, 32 Orthostira, 32 *Ostrya, 74, 86, 93, 116 ovata, Acalypta, 32, 33, 113 #oxycant,hoides, Ribes, 68, 117 pallida, Corythncha, 91, 105 pallipes, Corythncha, 15, 67, 73, 91, 92, 110, 112 (Fig. 6E), 114, 115, 116, 118, 119 *Panicum, 48, 49, 50, 51, 115 ^papyrifera, Betula, 93, 116 #Broussonetia, 72, 117 parshleyi, Corythncha, 84, 85, 107, 114 Physatocheila, 25, 113 peckhami, Galeatus, 10, 39, 40, 41, 113, 121 Sphaerocysta, 39 #pensylvanicum, Acer, 93, 119 #peregrina, Comptonia, 50 pergandei, Corythucha, 15, 67, 83, 84, 94, 105, 112 (Fig. 6F), 114, 115, 116, 117, 118 #Petalostemum, 39, 119 Physatocheila, 9, 12, 24, 113, 115 *Pieris, 54, 59, 120 Piesma, 9, 11 (Fig. 1), 16, 72, 107, 113, 115, 119 1 Piesmidae, 9, 16, 113 #Pinus, 50 *Pitcheri, Salvia, 45 #Platanaceae, 117 #Platanus, 72, 117 Plectana, 45 plexa, Physatocheila, 12, 24, 25, 26, 113 Physatocheila, 26, 113 Podisus, 45 *Polytrichum, 33, 34, 114 ^ponticum, Rhododendron, 57, 120 #populifolia, Betula, 50, 95, 116 138 Volume XXXI "Populus, 90, 115 "prinoides, Quercus, 64, 117 "Prinus, Quercus, 63, 117 "pruinosa, Crataegus, 68, 118 pruni, Corythucha, 14, 65, 68, 69, 72, 77, 78, 96, 97, 100, 101, 102 (Graph 3), 103 (Graph 4), 104 (Graph 5), 108, 110, 112 (Fig. 6G), 114, 118 "Prunus, 50, 66, 77, 78, 95, 96, 97, 98, 100, 101, 118 "punctata, Crataegus, 68, 118 "Pyracantha, 77, 118 pyriformis, Corythucha, 96 pyrioides, Stephanitis, 14, 54, 55, 56, 57, 114, 120 Ting is, 56 "Pyrus, 77, 93, 118 quadrata, Piesma, 18 "Quercus, 23, 50, 63, 64, 77, 116, 117 reflexa, Melanorhopala, 21, 113 "retroflexus, Amaranthus, 17, 18, 19, 117 "Rhamnaceae, 119 rhododendri, Stephanitis, 14, 55, 56, 58, 59, 61, 113, 120 "Rhododendron, 54, 56, 57, 59, 120 "Ribes, 68, 91, 117 "rigida, Pinus, 50 "Rosaceae, 68, 77, 118 "Rubiaceae, 121 "rubra, Moms, 91 Myrmica, 31 Quercus, 23, 63, 117 "Ulmus, 83, 106, 117 "rubrum, Acer, 50 "Rubus, 78, 86, 118 "rugosa, Alnus, 95 "saccharinum, Acer, 93, 119 "saccharum, Acer, 93, 119 "Salicaceae, 115 salicis, Corythucha, 90, 114 "Salix, 86, 90, 115 "Salvia, 45 "Saxifragaceae, 68, 117 "schlippenbachii, Rhododendron, 57 "Scirpus, 18, 115 "scoparius, Cytisus, 31 "Sedum, 31 "sempervirens, Solidago, 89, 121 "serotina, Prunus, 50, 66, 77, 78, 96, 100, 118 "sieboldiana, Juglans, 86, 115 "Salix, 93 "Sieboldii, Salix, 93 "Solanaceae, 121 solani, Gargaphia, 14, 43, 45, 46, 113, 121 "Solanum, 44, 45, 121 "Solidago, 22, 89, 121 "Sorbus, 77, 83, 86, 118 Sphaerocysta, 39 "Sphagnum, 35, 93 "spicata, Danthonia, 50 "spicatum, Acer, 93, 119 spinulosa, Corythucha, 66, 114 stellata, Plectana, 45 Stephanitis, 10, 14, 42, 47, 53, 54, 55, 59, 63, 120 "Strobus, Pinus, 50 "succulenta, Crataegus, 68, 118 "Tanacetum, 89, 122 "Thomasi, Ulmus, 106, 117 thomsonii, Acalypta, 13, 35, 113 "Thymus, 31 139 ENTOMOLOGICA AMERICANA •Tilia, 39, 47, 86, 91 *Tiliaceae, 119 tiliae, Gargaphia, 14, 43, 44, 45, 46, 47, 86, 113, 119 ^tinctoria, Baptisia, 38, 119 Tingidae, 9, 19, 113 Tingis, 16, 21, 25, 28, 39, 45, 53, 56, 63, 72, 74, 84 Tingoidea, 8 *tomentosa, Carya, 27, 115 #Tracheophyta, 115 tricornis, Dictyonota, 10, 30, 31, 113 *trifida, Ambrosia, 88, 121 Triphleps, 45 Trombidium, 93 ^tuberosum, Solannm, 44, 121 #Ulmaceae, 117 nlmi, Corythncha, 15, 67, 68, 94, 105,106, 112 (Fig. 6H), 114, 117 Corythucha pallida var., 105 *Ulmus, 83, 95, 106, 117 uniformis, Tingis , 21, 113 ^Vaccinium, 28, 41, 50, 78 variegata, Physatocheila, 12, 26, 113, 115 *Verbascum, 45 #virginiana, Jnniperns, 50 *Ostrya, 74, 93, 116 ^virginicus, Chionanthus, 29 *Vitaceae, 119 *Vitis, 119 *vulgare, Tanacetnm, 122 *' vulgaris, Beta, 117 yedoense, Rhododendron, 57, 120 140 - see its VOL. XXXII (New Series) Americana A Journal of Entomology. PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY PUBLICATION COMMITTEE JOSEPH C. BEQUAERT, Editor GEORGE S. TULLOCH E. W. TEALE Published for the Society by the Business Press Inc. N. Queen St. and McGovern Ave., Lancaster, Pa. Subscription, $5.00 per year Date of issue, February 20, 1953 A Journal of Entomology. Volume XXXII (New Series) for 1952 PUBLICATION COMMITTEE JOSEPH C. BEQUAERT, EDITOR GEORGE S. TULLOCH EDWIN WAY TEALE PUBLISHED BY THE BROOKLYN ENTOMOLOGICAL SOCIETY 1953 ENTOMOLOGICA AMERICANA Vol. XXXII (N. S.);> for 1952 CONTENTS PAGES The Hippoboscidae or Louse-Flies (Diptera) of Mammals and Birds. Part I. Structure, Physiology and Natural His- tory. Joseph C. Bequaert. February 20, 1953 1-209 {To be continued and indexed in vol. XXXIII) Americana Volume XXXII OF MAMMALS AND BIRDS PART I. STRUCTURE, PHYSIOLOGY AND NATURAL HISTORY By Joseph C. Bequaert Alexander Agassiz Professor of Zoology, Museum of Comparative Zoology at Harvard College, Cambridge, Massachusetts CONTENTS Page Introduction 3 Measurements 5 Collection and Preservation 6 Acknowledgments 12 External Anatomy 34 Coloration 57 Internal Anatomy and Physiology 60 I. Internal Appendages of the Integument 60 II. Muscular System 61 III. Nervous System 61 IV. Digestive and Excretory Systems 67 V. Adipose System 71 VI. Glandular System 71 VII. Respiratory System 74 VIII. Circulatory System 77 1 ENTOMOLOGICA AMERICANA Page IX. Reproductive System 80 A. Internal Male Organs 80 B. Internal Female Organs 81 Development and Early Stages 86 Egg 87 Larval Instars 88 Imaginal Disks 92 Puparium 93 Natural History 96 I. Food 97 The Blood Diet 97 Intracellular Symbionts and Nutrition 101 Differential Feeding and Host Specificity 105 Mechanism of Feeding 107 Size of Blood Meal Ill Frequency of Meals 112 Resistance to Starvation 114 Cannibalism 116 II. Shelter 116 Physical Characteristics of Feathers and Hair 116 Effect of Macroclimate 120 Preferential Distribution on the Host 124 Protective Behavior 126 Predacious Enemies 129 Parasites 131 Phoresy of Mallophaga 163 Phoresy of Psocoptera 173 Phoresy of Thysanoptera 174 Phoresy of Pseudoscorpions 174 III. Reproduction 174 Sexual Dimorphism 175 Sex Ratio 176 Maturation Period 179 Mating 180 Intra-Uterine Development 183 Larvi position 184 Formation of Puparium 195 Duration of Pupal Development 197 Individual Life Cycle 202 Adult Longevity 202 Rate of Reproduction 204 2 Volume XXXII 1 ‘ Blut ist ein ganz besondrer Saft. ’ ’ Goethe’s Faust, Part I. INTRODUCTION The present work is the outcome of some thirty years’ study of the dipterous family Hippoboscidae, the ectoparasitic louse-flies of birds and mammals, variously called also bird-flies, feather-flies, ked-flies or keds, spider-flies, tick-flies, and flat-flies. As originally conceived, it was to be only a taxonomic account restricted to the species occurring north of Mexico, for which area information is most complete. The Nearctic hippoboscid fauna is, however, so obviously an extension of that of the Neotropics, that eventually it seemed ad- visable to cover the whole of the New World. The larger scope will no doubt make the work more generally useful, while placing the North American fauna in its true perspective. On the other hand, I am only too well aware that the available Central and South American material is very scant, particularly in view of the well- known fact that the Neotropical bird fauna is richer than that of any other of the Earth’s zoological areas. It is hoped, nevertheless, that in its present form the paper may be a safe working basis, as well as an incentive for future investigations. The plan was at first that of my Monograph of the Melophaginae (1942a) ; but the introductory account of structure, function and bionomics has gradually grown into what purports to be a general Natural History of the Hippoboscidae, which is now published as Part I of the work. Several aspects of the structure, physiology and behavior of these insects are almost unique, so that they are of the greatest interest, particularly to the student of parasitism and evolution. Their study is also urgently needed, since many of these flies are doomed to extinction, with their hosts, in the near future. 1 have attempted to compile, to appraise critically from personal observations, and to correlate all published information on their outer and inner structure, physiology, ecology and behavior, calling attention meanwhile to the gaps that should be filled in by future research. Where information is either lacking or very scant for the louse-flies, I have sometimes referred to what is known of their nearest living relatives, particularly the tsetse-flies. Needless to say, it is not claimed that such information can actually be extended to the Hippoboscidae in every case. It is not easy to write a properly balanced summary of a variety of subjects. Like every other naturalist, I have a weakness for certain topics, which is bound to be reflected in the disproportionate 3 ENTOMOLOGICA AMERICANA extension given to certain sections. Other matters are more con- cisely treated, but I trust that no worthwhile contribution has been overlooked. I should also warn that certain publications, clearly out-of-date or obviously unreliable, have been deliberately neg- lected, although all are included in the Bibliography. The general account of Part I called for so many references to papers not mentioned in the taxonomic bibliographies of Part II, that it was necessary to incorporate in the work a reasonably com- plete Bibliography of the Hippoboscidae to date. In the taxonomic section forming the second part of the paper, the key to the genera of the Hippoboscidae of the World which I recognize as distinct, is designed for identification purposes only. It gives no clues to the true relationships, which are discussed in the chapter on “Relationship and Classification.” Much effort has gone into preparing detailed generic descriptions, on a compara- tive basis, as none of those published to date are either reliable or adequate. A careful analysis of generic structure is, in my opinion, essential not only for the study of evolution within the family, but also for evaluating the true specific characters within each genus. I have been led to suppress some of the genera accepted thus far, since they were clearly based on characters which, though readily observable, are nevertheless trivial in the family as a whole. Complete bibliographies of previous writings are given under each species, nearly all citations having been compared with the originals. Each citation also mentions briefly the most important information it contains, particularly all new localities and hosts, except for my own papers, since their contents are included in the text. Thus the bibliographies give credit where it is due and reflect the vicissitudes in the gradual growth of our present knowledge. Slow but steady accretions from many sources is an outstanding historical feature of Entomology. Each addition may be trivial by itself, but the sum total is often impressive, as is perhaps shown by the present memoir. No new detailed descriptions of species were drawn up. In- stead, the keys to the species include nearly all the characters believed to be fully reliable for diagnosis. Together with the illus- trations, they should suffice for identification; but some additional features of minor value have been mentioned in discussing the af- finities. Copies of the original descriptions, if needed in English translations, except for the Latin texts, have been appended, as well as remarks on the type specimens whenever these were avail- able. 4 Volume XXXII The paragraphs entitled “ Distribution and Specimens Exam- ined” detail under each species the localities and hosts from which I have seen material. In addition to new records, they include those published previously (as cited in the bibliographies), when I was able to study the specimens. Inasmuch as I have given full credit to earlier work, I feel that I do not infringe on other people ’s rights if I claim the responsibility for the identification of all specimens I have studied myself. In some cases the published records needed correction, while in others they were based in the first place on my own identifications. When the species or subspecies of the host is definitely known, I use as a rule its scientific name in the body of the paper. As the vernacular names of birds and mammals are in more common use, particularly in Canada and the United States, there might have been some advantage in using these instead. English or other vernacular names, however, could only be confusing if used for for- eign or tropical hosts ; even some of the American names might be meaningless or misleading to British readers. Vernaculars will be added to the scientific names in the 1 ‘ Systematic List of the Hosts and their Parasitic Plies” which will conclude the second or taxo- nomic part. Another concluding section will list the American species of flies according to geographical areas, political divisions (including the provinces and states of Canada and the United States) being used for convenience. A more detailed discussion of several aspects of distribution in the Americas will then be presented. The following information is inserted in square brackets in the text. 1. Entries in the bibliographies that do not conform to bino- mial nomenclature, such as nomina nuda, pre-Linnaean citations, vernacular names, etc. 2. Information received from correspond- ents, but not confirmed by personal examination of specimens. 3. Explanatory words or remarks inserted in the copies or translations of original descriptions, more particularly the metrical equivalents of non-metrical measurements. Measurements. In the Hippoboscidae size is in most cases a fairly reliable specific character, varying within rather narrow limits. The total length of the body is, however, of no significance, as the soft integument of the abdomen is expansible ; in the same individual it is strongly contracted and hence very short upon emerging from the puparium, but increases greatly in size after feeding or, in the female, during the intra-uterine growth of a larva. The abdomen also shrivels after death, particularly in dry, 5 ENTOMOLOGICA AMERICANA pinned specimens. Only two mensurations of length are fully trustworthy and comparable : the combined length of head and thorax, taken from the tips of the apical arms of the frans to the hind margin of the scutellum; and the length of the wing, taken from the base of the tegula to the tip of the membrane. The pro- portion of the greatest width of the thorax (before the base of the wing) to its median length (from the anterior margin to the hind margin of the scutellum) is also of some value, as well as the great- est width of the wing. The proportions of length to greatest width of the head and of the interocular face are also frequently used; but some judgment is needed in determining them. In specimens pinned after being kept for some time in a liquid, the face or eyes often shrivel or shrink from the original width in life. The length (or height) of the head should be measured from the tips of the apical arms of the frons to the occipital margin of the postvertex; while its greatest width is the distance between the extreme outer sides in front view, usually the outermost bulges of the eyes. The interocular face is the area bounded by the inner margins of the eyes, its length being that of an inner eye margin; its width, or horizontal distance between the inner eye margins, usually varies at different levels, the margins being rarely parallel; consequently, it is necessary to indicate the level at which the width was meas- ured. The correct length and width of head and thorax and the proportions between them should be determined on specimens not mounted on slides. Slide mounts are more satisfactory for wing measurements. Most measurements vary individually in the same species within extremes of one-half to one millimeter; so that an approximation of half a millimeter is satisfactory for most purposes. Collection and Preservation. Collecting Hippoboscidae pre- sents special difficulties for the average entomologist, so I offer no apology for the following suggestions. Occasionally a stray speci- men may be collected away from a host or newly emerged flies may be seen flying, particularly species of Lipoptena which sometimes swarm in numbers in search of a host. Such forms as the Melopha- gus of sheep, Lipoptena of goats, and Hippobosca of horses, cattle, camels and dogs, may be collected without much trouble from their domestic hosts; it is relatively simple to obtain Pseudolynchia canariensis and its puparia from domestic pigeons or in pigeon lofts. Some Old World species of Ornithomyia, Crataerina, and Stenep - teryx may be bred from puparia found in birds ’ nests ; this is hardly ever possible in the New World. Olfersia is sometimes abundant in or near the rookeries of certain marine bird hosts. The net re- 6 Volume XXXII suits of such random collecting are practically negligible, as shown by the scarcity of louse-flies in the average dipterist’s or museum collection. Incidental captures can never give a complete insight into the local hippoboscid fauna and its relations to the hosts. During the many years I have been interested in these flies, I have collected personally very few of them, possibly not more than 1 or 2 per cent of the total number of specimens I have used for my work. The student of the louse-flies must enlist the assistance of or- nithologists, game-wardens, wild-life experts and hungers, a prob- lem in human relations which he will have to solve as best he can. Personally I have usually found in such quarters the greatest will- ingness to help. It is particularly recommended to establish friendly contacts with organizations or persons engaged in the banding (or ringing) of birds. Juvenile birds usually predomi- nate among birds trapped for banding during the fall migration and they are often more heavily infested than the older birds of the same species. Some further remarks on bird banding in rela- tion to the study of the Hippoboscidae will be found in the intro- ductory paragraphs of the section on the host-parasite relation. As was first pointed out by H. S. Peters (1930), the systematic and extensive trapping of live birds offers unusual opportunities for the study of many little-known aspects of hippoboscid natural history. Latterly Mr. I. B. Tarshis has made extensive use of the method in studying the two flies of quail ( Lophortyx ) in California, Lynchia hirsuta and Stilbometopa impressa. His results were so far-reaching that I urged him to publish a detailed description of the two types of portable cages he devised for the examination of trapped quail in the field. With Mr. Tarshis’ permission, general views of these cages are reproduced (Figs. 1 and 2). Instructions for their construction and use will be found in his recent paper (1952). A simple, but very efficient method of collecting a bird’s flies without harming it was developed recently by K. Williamson (1952) at the Fair Isle Bird Observatory, Scotland. The live bird is immersed in a bath of chloroform fumes whilst keeping its head clear by means of a cape of oiled silk. It is the only method which provides data suitable for a statistical analysis of fly popula- tions (Butterfield, 1952). Young waders and birds of prey are sometimes banded in the nest; but most flies of large hosts must be obtained from killed birds ; unless trapping can be used, this is at present the only avail- able method for those of tropical birds and of wild mammals. Fully-winged flies usually leave the host shortly after its death or 7 ENTOMOLOGICA AMERICANA in the field. Prom I. B. Tarshis (1952, p. 71). 8 Volume XXXII ENTOMOLOGICA AMERICANA even while it is dying. They may also, in addition, move so swiftly among the feathers or hairs that they are likely to dodge the col- lector’s fingers. In order to reduce losses to a minimum and to make sure of the correct host, it is strongly recommended that each bird be tied up as soon as possible after death in an individual strong paper or cloth bag of suitable size. Either the contents should be exposed to ether or chloroform fumes before opening the bag, or, better yet, the untreated bag should be opened indoors and the bird examined near a closed or screened window. A fly that leaves the body will usually dart for the screen or glass pane, where it can be stuck to a moistened finger and washed off directly in a vial of alcohol. In the field, hosts may be examined inside a mosquito bar with a very fine mesh. Occasionally a fly may remain on a dead bird for several hours and be found while preparing the skin. When several birds of different species lie about during the skinning process, flies may easily move from one bird to another, so that rec- ords obtained under such conditions are often questionable. This is even more true when hosts of several species are brought in from the field in the same bag or container, a frequent source of erroneous host records. Sometimes a fly, after leaving a dead or dying bird, is unable to find a new live host and eventually returns to a dead bird, either the one it had just left or some other one lying near it. Flightless hippoboscids tend to remain on dead hosts much longer than fully-winged species. For best results in later studies, the flies should be dropped as soon as possible after capture in 75 to 80 per cent clean, unadulter- ated ethyl alcohol, and be properly labelled with locality, date, name of collector and of host. It is better not to add glycerine to the alcohol, as this may interfere with later study. The name of the host should be given as accurately as possible and preferably ob- tained from a competent ornithologist or mammalogist. If the skin of the host is preserved, its serial number should be given on the label of the fly so that the name of the host may eventually be veri- fied. These precautions are most important in tropical areas, where many birds are as yet imperfectly known and their parasites even less. Since no stoppered vial has yet been devised from which alcohol will not evaporate eventually, the vials should be plugged with cotton and permanently kept immersed in alcohol in larger containers in the museum. Dry, pinned specimens of known species can usually be cor- rectly identified. However, they show rarely all the specific charac- ters clearly, particularly those of the abdomen, which is often 10 Volume XXXII shrivelled to such an extent that even the sex can no longer he determined. Even prolonged soaking, boiling, or maceration usually fail to soften the dried abdomen and to restore it to the original shape. I advise against mounting hippoboscids on slides, either in Canada balsam or in some other medium, after softening and clearing the specimens by maceration and removal of the inner tissues. Such manipulations are apt to remove or distort certain delicate outer structures, while making visible by transparency various inner structures, apodemes and ridges, which may be readily mistaken for outer structures. On the other hand, the true surface limits of the sclerites and the true external structures may become obliterated, or some of the characteristic chaetotaxy may be lost. Furthermore, it is nearly impossible to determine with any degree of accuracy from a fly mounted under a cover glass the original proportions of the several external body areas, notably those of the interocular face. Cleared specimens are useful only for the special investigation, of certain minute structures and even then should be used with extreme care, lest they lead to erroneous conclusions. A case in point is the determination of the number and location of the abdominal spiracles, which seem to be easily re- moved with the tracheae by the clearing manipulations. I was privileged to examine slides of several species of Ornithoica pre- pared some years ago by an experienced entomologist. Although it was eventually possible to locate all seven pairs of abdominal spiracles by combining the findings on all the slides, no single slide showed the full set and some showed none at all. # Personally, I am reluctant to describe a new species of hippo- boscid solely from either a dry and pinned specimen or a slide mount. Much of the difficulty in recognizing the true identity of the types of earlier authors lies in their being poorly preserved, pinned specimens, on which some of the essential specific or even generic characters cannot be seen. An additional word of caution to the beginner may be pertinent, since it is hoped that the present work will be the incentive for a renewed interest in the Hippoboscidae. He should be wary of regarding as a novelty any specimen that seems to depart from the published figures and descriptions of the known species. Quite apart from individual intraspecific variation, which may be wider than suspected or recognized, the condition, age or method of pres- ervation of the specimen may be deceiving. Published information also may be not fully reliable and even drawings by reputed and skilful workers may be misleading. My long experience with the 11 ENTOMOLOGICA AMERICANA Hippoboscidae leads me to believe that the probability of a begin- ner’s finding an nndescribed species in a limited material is very small and I should think nil for the United States and Europe. If anyone wishes to “split” the now recognized Nearctic and Palearc- tic species, or to subdivide them into subspecies, he should choose names from among those now stored in the synonymy, rather than invent new ones. The descriptions and original drawings in the present work were made by the author almost exclusively from alcoholic material. This was left to dry just enough to disclose the limits of the sclerites, the sutures and the insertion of the setae under the dissecting, bino- cular microscope. Care should be taken to keep the specimen suf- ficiently moist, so as to prevent the appendages from becoming brittle and the abdomen from shrivelling. Illustrations copied or adapted from my predecessors, as far as possible with their per- mission, have been fully credited to them. Acknowledgments. I have been more than usually fortunate in enlisting the assistance of many entomological colleagues, or- nithological friends, game- war dens, museum curators and even casual acquaintances, all of whom have provided specimens or information. Their names will be listed at the close of Part II ; at present I wish to thank them collectively for their kind and gener- ous help. To the following persons I wish to express my appreciation for special contributions to various sections of Part I. Mr. Irvin Barry Tarshis, of Berkeley, California, has most generously communicated some of the unpublished results of his researches on the bird-flies of quail, and I am also indebted to him for many other favors. I have likewise been privileged to read in manuscript the important ob- servations on the pigeon-fly by Dr. J. H. Schuurmans Stekhoven and his co-workers at the Fundacion Miguel Lillo of Tucuman, Argen- tina, which are to appear shortly and perhaps even before my own work. Information obtained from both these sources has enabled me to put some earlier published contributions in the proper pers- pective. Relatively little of this unpublished work, however, has been mentioned in my paper, with the authors’ permission, as I did not wish to detract from the originality of their forthcoming pub- lications. I am indebted to Dr. B. Jobling, The Wellcome Research Lab- oratories of Tropical Medicine, London, England, not only for permission to reproduce some of his published figures, but also for advice on moot problems of anatomy and particularly on the true 12 Volume XXXII homologies of frons and clypeus ; Figs. 6E-F are from his original unpublished drawings. Dr. J. E. Webb, University College, Ibadan, Nigeria, and the Zoological Society of London have kindly allowed me to copy the illustrations in Fig. 16. Similar permission was granted by Mr. K. MacArthur and several other persons as well as Institutions, as mentioned in the explanations of some of the figures. Dr. Clay G. Huff, Naval Medical Research Institute, Bethesda, Maryland, and Dr. G. Robert Coatney, National Institutes of Health, also at Bethesda, have offered valuable suggestions in dis- cussions of the development of pigeon malaria and similar blood parasites. Professor H. R. Hagan, College of the City of New York, read critically the section on the embryology and early de- velopment, and Dr. Kenneth W. Cooper that dealing with the chromosomes. Much of the information on the Laboulbeniaceae was obtained from Mr. Richard K. Benjamin, while he was a Graduate Student at the Biological Laboratories of Harvard University. In connection with these fungi, I received valuable help also from Mrs. F. L. Balfour-Browne, at the British Museum (Natural History). Mr. G. B. Thompson, Cambridge, England, has for many years been most generous in communicating interesting material and valu- able information, particularly on publications otherwise inacces- sible. Dr. F. I. van Emden, British Museum (Natural History), and Mr. E. Seguy, of the Paris Museum of Natural History, have contributed much time and advice, either during visits to study the collections in their care or by correspondence. I owe a special debt of gratitude to Mr. James E. Collin, Newmarket, England, for his unfailing assistance in answering my repeated inquiries and more particularly for his kind hospitality while studying the valu- able types of Bigot’s Hippoboscidae at his home in September, 1951. It would have been utterly impossible to attempt an intelligent discussion of the host-parasite relation of the Hippoboscidae, one of the major sections of this work, without the continued help and suggestions of several ornithologists and mammalogists here and abroad. I wish particularly to express my appreciation to my life- long friend, Dr. James P. Chapin, The American Museum of Nat- ural History, New York, who devoted much time to a criticism of parts of my manuscript dealing with ornithological matters. I am also under special obligation to my colleagues at the Museum of Comparative Zoology, the late James Lee Peters, Mr. James C. Greenway, Mrs. Barbara Lawrence Schevill, and Dr. Charles P. Lyman. 13 ENTOMOLOGICA AMERICANA Finally, I wish to express my gratitude to the Treasurer and the Publication Committee of the Brooklyn Entomological Society, whose generous and sympathetic interest made possible the publica- tion of this work in its present form. EXTERNAL ANATOMY The following account of the external anatomy has a threefold purpose. It intends first to define the terms used in the taxonomic discussions for the several parts or areas of the exoskeleton. It attempts also to trace, in so far as possible, the homologies of these parts in the Hippoboscidae with those of other related Diptera. Finally it explores the most important modifications within the group, preliminary to a discussion of the relationships of the sub- families and genera. In tracing the homologies and in selecting and defining the terms, an earnest attempt has been made to adopt the latest findings of comparative insect anatomy. The professional morphologist is prone to decry the non-morphological terminology used in the Dip- tera by the average taxonomist, whom he seems to regard as a mere willing victim of mental sloth. After spending many tedious hours on a variety of morphological writings, I have reached the conclu- sion that a reasonably stable terminology, based on the true homolo- gies and applicable to all parts throughout the Order Diptera, is as yet a hopeful dream. Morphologists disagree among themselves to such an extent that the unbiased outsider is often at a loss to decide which point of view to adopt. Their involved discussions of moot points sometimes raise the suspicion that they are perhaps not quite so sure of their ground. Meanwhile the taxonomist must use a certain term with a definite meaning and cannot well wait until the morphologists have settled thn matter or change his terminology every other year. With regard to the Hippoboscidae in particular, the morphologists usually stop so far from the goal that they are of little assistance in deciding which terms to adopt. Moreover, I am by no means convinced that, in such highly specialized insects as the ectoparasitic Diptera, it will ever be possible to trace the true homologies of all the parts. Sometimes one interpretation seems to carry as much weight as another, in which case it may be better to “accept the facts as they are” (R. E. Snodgrass) and to adopt conventional terms, which at least have the advantage of not imply- ing more knowledge than is actually attainable. Anticipating a later discussion of the affinities, it may be help- ful to state at the outset that the term “Diptera Pupipara” (or 14 Volume XXXII “Pupipara” for short) will be used as a convenient collective name for the Hippoboscidae, Streblidae and Nycteribiidae, the only dip- terous families of permanent ectoparasites with integral viviparity (so-called pupiparity). It does not intend to cover a taxonomic category nor to imply any close relationship or common origin. The Hippoboscidae will be divided in the present paper into six subfamilies, two of them, the Ornithoicinae and Ornithomyiinae, being restricted to birds and four, the Hippoboscinae, Melophagi- nae, Ortholfersiinae and Alloboscinae, to mammals (except for Hippobosca struthionis of the ostrich). In order to trace the homologies, I have used for comparison the head and thorax of one of the Glossinidae or tsetse-flies, Glossina fusca Walker, original drawings of which are included. Among the higher Muscoidea, these flies appear to be structurally most similar to the Plippoboscidae and possibly their closest living relatives. The structure of the head and thorax of the Hippoboscidae is most readily understood if the adult fly is conceived as a dorso- ventrally flattened, blood-sucking, higher muscoid. The hippobos- cid head capsule was first explained on this assumption by B. Jobling (1926, p. 329; with a useful diagram, text-fig. I). In addition, certain areas or parts of the exoskeleton have been modified, mainly by fusion, reduction or atrophy, in response to permanent ecto- parasitism. The dorso-ventral flattening is less apparent in the abdomen, due to the reduction of the sclerotized areas, itself cor- related with the peculiar viviparous mode of reproduction. The head (Fig. 3A-C ; Microlynchia pusilla) is clearly a strongly flattened muscoid head, in which several of the original areas may be recognized without much difficulty. The upper surface either is horizontal or slants slightly downward and forward. The head shows none of the extreme modifications of the Streblidae and Nycteribiidae and its structure is fairly uniform throughout the family. That of the most highly specialized genus, Melophagus , may be tied up through the related Lipoptena with that of the more generalized Ornithomyia and Ornithoica. The relative uniformity of the head and its appendages, particularly the mouth-parts, points strongly to a monophyletic origin for the entire family. Owing to the dorso-ventral flattening, the mouth-parts are directed forward, not downward as in the Muscoidea, the head being of the prognathous type. As in all Myiodaria, the occipital foramen ( OCF ) is relatively small and occupies a median or submedian position on the hind part of the head, which is connected with the 15 ENTOMOLOGICA AMERICANA Fig. 3. A-C. Microlynchia pusilla (Speiser), female; head from above and below (A), from behind (B) and in profile (C) : A, antenna; AAF, apical arm of frons; EY, eye; EEC, frontal carina; frs, frontal suture; G, gena; GTJ , gula; HA, haustellum; 10, inner orbit; L, lunula; MV, mediovertex; OC, occiput; OCF, occipital foramen; OCM, occipital margin; orb, orbital bristles; FA, palpus; FG, postgena; pts, ptilinal suture; FV, postvertex; 16 Volume XXXII prothorax by means of a short and narrow membranous neck. I fail to see, however, that the neck is much more reduced than in other Diptera (including- the Myiodaria) with a freely movable head. In the Hippoboscinae the head is rounded posteriorly and entirely free from the thorax, with a distinct intervening pronotum (Fig. HE, PT ), so that it can move not only up and down, but also sideways. In all other Hippoboscidae the hind margin of the head juts against the more or less concave, or at least depressed, anterior margin of the thorax. The connection between head and thorax is rather loose in the Alloboscinae, Ortholfersiinae, Mel- ophaginae and Ornithoieinae, in which the humeral callosities form short and broad lobes, similar to those of the Muscoidea, allowing the head some free lateral motion. Im the Ornithomyiinae, how- ever, the humeral angles of the callosities are long and point for- ward, flanking the sides of the head in such a way that it can no longer move sideways. In extreme cases ( Crataerina , Stenepteryx, and Myiophthiria) the head fits so snugly in the thorax that the two form as it were a single streamline unit, at least for motion in one plane. Such an arrangement has obvious advantages for scurrying amidst the feathers of avian hosts. It is therefore note- worthy that the most evolved cephalo-thoracic connection is found only among some of the bird parasites and is most perfect in the flies of swifts and swallows ; while the more primitive condition of a freely movable or loosely attached head occurs, with the ex- ception of the Ornithoieinae, among the parasites of mammals. The most important landmark on the dorsal surface of the head, or face , is the conspicuous, transverse, more or less crescent- shaped ptilinal suture ( pts ), which marks the fissure where the in- flated pjilinum (pt ; ptilinal sac) retracts within the head capsule after the fly emerges from the puparium. The ptilinum is no less developed in most Hippoboscidae than in other Schizophora ; but owing to the flattening of the head and the strong sclerosis of the irons and inner orbits, the edges of the ptilinal suture fit together very tightly. The ptilinal suture divides the median portion of the face into an upper area or vertex ( V ; including the frontalia, parafrontalia and ocellar plate of authors) and a lower area or VO, upper orbit ; the combined mediovertex and postvertex is the vertex (F) ; v~b, vertical bristle. D, Glossina fusca Walker, female; head in profile. The line X — X in O and D marks the level of the dorso-ventral flattening of the head. E, Ornithomyia avicularia (Linnaeus), female; frontal area and antennae: A A, antennal appendage; IAS and 2 AS, first and second antennal segments; PA, palpus. 17 ENTOMOLOGICA AMERICANA frons ( FB ; clypeus of many authors; prefrons of Ferris, 1950; formerly called fronto-clypeus by Jobling, 1926; including the lunula, facialia and interfacialia or facial plate). Most charac- teristic for the Hippoboscidae is the flattened, near-horizontal frons, in the same plane as the vertex, the antennae being pushed forward to near the oral margin of the face. Within the family three types of frons may be recognized, here discussed somewhat in detail, be- cause of their bearing on the evolution within the group.1 1. The most primitive condition is found in the Ornithoicinae, a subfamily which on other grounds also appears to be more gen- eralized than any of the other Hippoboscidae. In Ornithoica (Fig. 5C), the frontal area is essentially as in most Muscoidea, except for the horizontal flattening and the forward position of the antennae. The supra-antennal (or post-antennal) region, or lunula (L; frontal lunule, frontal crescent, or antennaria of authors), is moderately wide and alone visible between the ptilinal suture and the base of the antennae. The antennae are large, flat, covering much of the frons and placed in a single deep antennal cavity (facial depression, facial plate, or interfacialia of authors in Muscoidea). Both enlarged first antennal segments, recognized by the presence of a row of bristles, lie side by side, their inner margins touching without intervening frontal carina (facial carina in Muscoidea, of most authors), except for a very short, triangular area set off by a fine transverse suture from the lower edge of the lunula. 2. Variants of the next known evolutional stage occur in the Ornithomyiinae. In Olfersia (Fig. 7F), Lynchia (Fig. 5A), Pseudolynchia, and M\crolynchia (Fig. 3A), where it is most readily 1 In a recent discussion of the structure of the insect cranium, Snodgrass (1947, p. 3) takes the view that the terms “frons,” “vertex,” “clypeus, ” and “genae” correspond to topographical and not to anatomical areas, so that a great deal of latitude may be allowed in their application to different types of insects. He is also of the opinion that most of the impressed lines or grooves on the surface of the head are not “sutures” in the true sense of the term, but merely “sulci” or furrows, incidental to the forma- tion of secondary strengthening ridges on the inner surface of the exoskeleton. In the present discussion I have found it more con- venient to adhere to the customary use of the term “suture” for most external impressed lines, regardless of their possible origin, which is frequently in dispute. 18 Volume XXXII Fig. 4. Heads of Hippoboscidae : A, Ornitheza metallica (Schiner), female, from above. B, Hippobosca longipennis Fabricius, female, from above. C, Lipopiena rusaecola J. Bequaert, male, from above; D, Stenepteryx hirundinis (Linnaeus), female, from above. E, Allobosca crassipes Speiser, female, from above (at right) and below (at left). F, Crataerina pallida (Latreille), female, from above. G, Lipoptena rusaecola J. Bequaert, male, in profile. 19 ENTOMOLOGICA AMERICANA studied owing to the small size of the antennae, the lunula, more extensive than in Ornithoica, occupies the major part of the frons; the bases of both first antennal segments are often fused with it, except at their inner and outer sides. In addition, the antennae are far apart and separated throughout by the widened, flattened, strongly sclerotized frontal carina ( FBC ). Apically this carina is deeply divided by a broadly triangular notch into long, diverging apical arms ( AAF ), while the basal, undivided portion may be depressed or grooved longitudinally; a distinct, complete trans- verse suture divides the lunula from the frontal carina. Each antenna lies in its own antennal pit ( AP ), open anteriorly, how- ever, where the appendage of the second antennal segment extends beyond it. Narrow lateral extensions of the lunula, corresponding to the facialia of the Muscoidea, form the outer margins of the antennal pits. In other Ornithomyiinae, such as Ornithomyia (Fig. 3E), Crataerina (Fig. 4F), Stenepteryx (Fig. 4D), and Myiophthiria, the frontal carina is often nearly completely fused with the lunula, though abruptly narrowed at the base. In Orni- thoctona the apical arms divide the frontal carina nearly to the base; in Ornitheza (Fig. 4A) they diverge at first, then converge again and run side by side before the tips. 3. The third and most derivative type is that of the Hip- poboscinae (Fig. 5B), Melophaginae (Fig. 4C), and Orthol- fersiinae. In these subfamilies the lunula and greatly widened frontal carina form a single sclerite, only slightly wider basally. The small antennae occupy the extreme lower corners of the frons, each in its own antennal pit, which is usually surrounded com- pletely by a continuous rim (except in the Ortholfersiinae) . In most cases the flaring apical arms of the frons are more or less developed, though rather short and widened basally. In the Melophaginae, by far the most specialized members of the family, the anterior (or oral) margin of the frons is broadly rounded off, with the barest trace of a median notch extended basad as a nar- row, superficial furrow; very rarely there is a faint indication of a transverse line setting off the lunula above the antennae. In some respects the Alloboscinae are intermediate between the second and third types : the frontal carina forms a broad, flattened plate, fused basally with the lunula, though abruptly narrower and sharply set off from both adjoining first antennal segments, themselves fused basally with the sides of the lunula ; the antennal pits are incomplete anteriorly, the appendages of the second an- tennal segments extending far beyond them (Fig. 5D). 20 Volume XXXII Fig. 5. A-D, Frontal areas and antennae of Hippoboscidae : A, Lynchia albipennis (Say), female. B, Bippobosca longipennis Fabricius, female. C, Ornithoica vicina (Walker), female. D, Allobosca crassipes Speiser, female. E, Lipoptena rusaecola J. Bequaert, male; head from below. 21 ENTOMOLOGICA AMERICANA Some of the higher Muscoidea show indications of changes in the lunula and frontal carina which may help to understand the extreme modifications of the Hippoboscidae. In Pollenia rudis, the upper part of the frontal carina is distinctly widened between the antennae ; moreover, the first antennal segment, although very small when compared to the second and third, is shaped much as in Ornithoica. The frontal carina is much more developed in several of the so-called bot-flies, unrelated higher Muscoidea parasitic in the larval stages on mammals, and particularly among the Cutere- bridae and the Hypodermatidae. It reaches its extreme develop- ment in Oestromyia, where the antennal region of the frons is not unlike that of the more derivative Ilippobosciade : the frontal carina, completely fused with the lunula, is very broad, short and flat, with diverging lateral edges forming the inner rims of small, anteriorly open antennal pits. Beyond and beneath the apical arms or edge of the frons may be traced the greatly reduced true clypeus (CL; anteclypeus or epistome of authors), on which the cibarial muscles are attached. It is usually in a ventral position and sometimes far back, as in Melophagus, where it lies near the middle of the ventral surface of the frons. The clypeus serves as a hinge for the rostrum of the proboscis and is connected by lateral sclerotized plates or bars with the cibarial pump. (Figs. 6E-F, drawn by Dr. B. Jobling). A comparison of the frontal and clypeal areas of the higher Muscoidea and Hippoboscidae shows that the head of the latter is flattened approximately at the level of the line X — X in Fig. 3D of Glossina fusca and Fig. 3C of Microlynchia pusilla. The flattening results from an outward transverse fold of the in- tegument running from the occipital foramen across the lower portion of the eyes to the ends of the lateral frontal sutures and to the lower margin of the frons proper (vibrissal angles of the Muscoidea). The area above the line X — X became a horizontal plane ; that below the line folded backward and upward beneath the head, much of it providing the material for the soft rostrum membrane within which the proboscis may be retracted. The head is more flattened anteriorly than behind, so that it is triangular in profile and wedge-shaped (Figs. 3C and 4G), the keel-shaped anterior (frontal) margin being ideal for sliding forward amidst the plumage or pelt of a vertebrate host. The upper part of the vertex is a median, strongly sclerotized, usually well-defined plate, the postvertex ( PV ; vertical triangle or ocellar plate of authors). It bears the three ocelli ( OCI ), when 22 Volume XXXII these are present. The blunt or sharp edge between the postvertex and occiput is the occipital margin (OCM). The entire area of the face limited by the inner margins of the eyes is sometimes called the frons; but it is better referred to as the interocular face ( IF • postfacial area of certain authors). It consists mainly of a median, generally soft, wrinkled portion, the mediovertex (MV ; frontal vitta, frontalia, or interfrontalia of writers on the Muscoidea), with a rough surface of microscopic scales giving it a dull appearance. It is separated on each side from the eye by a narrow, hard sclerite, the inner orbit (10 ; geno-vertical plate, parafrontal, or laterovertex of authors). Olfersia is unusual in that the ptilinal suture has moved so far back (or upward) that it divides the face into two nearly equal areas; the postvertex, devoid of ocelli, en- croaches upon the mediovertex and often reaches the ptilinal suture. The dorso-lateral portion of the head, behind each eye, is the outer orbit (00), usually very narrow and sometimes lacking, particularly when the eyes extend over the ventral side. The upper area, above the eye, is the upper orbit (TJO). The narrow lateral area, continuing the inner orbit between the lower border of the eye and the frontal suture (frs; lateral extensions of the ptilinal suture) is the gena (G; cheek or jowl of authors) ; although the extreme forward position of the antennae in the Hippoboscidae makes it impossible to distinguish the parafacial areas clearly from the genae. The hind or occipital surface of the head is divided by the occipital foramen (OCF) into an upper area, the occiput proper (OC), a lower area, the gula (GTJ), and two lateral postgenae ( PG ). The compound eyes (EY) are as a rule large, always broadly separated in both sexes and with many small facets all of the same size. In some genera they are entirely dorsal, occupying the major part of the sides of the face. In the Ornithoicinae and Melo- phaginae they are, in addition, smaller and particularly narrower than usual. The smallest eyes, with the fewest ommatidia, are those of Myiophthiria and Melophagus (about 135 ommatidia in the latter ). The surface of the eye is always bare. The develop- ment of the ocelli (OCI) varies greatly. In appraising evolu- tionary trends within the family their presence may safely be re- regarded as marking the more primitive types. In nearly half of the genera (8 out of 20) three ocelli are present, although they may be very small or vestigial in certain species. The majority of these genera are bird parasites of the subfamilies Ornithoicinae and Ornithomyiinae ; but two of them are mammalian parasites of the 23 ENTOMOLOGICA AMERICANA Fig. 6. A-C, Pseudolynchia canariensis (Macquart), after Jobling (1926): A, tip of haustellum in side view, showing labella (LA), ends of labial gutter (LG) and of theca (TH), and all prestomal teeth (PET) everted. B, end view of labella with two sets of prestomal teeth inverted. C, third antennal segment and arista removed from cavity of second segment. D, ILippobosca equina Linnaeus; end view of labella with two sets of prestomal teeth (PET) everted, after Jobling (1926). E-F, Ornithomyia avicularia (Linnaeus), after 24 Volume XXXII subfamily Melophaginae. Of the remaining 12 genera, 11 are always without even traces of ocelli, 5 of these occurring on birds and 6 on mammals. The lack of ocelli is the invariable rule among the mammal-inhabiting Hippoboscinae, Alloboscinae and Orthol- fersiinae. The Melophaginae alone among the parasites of mam- mals contain both genera with and without ocelli. In the Orni- thomyiinae, the genus Lynchia is aberrant in that most of the species are entirely without ocelli, while a few show more or less distinct vestiges of them and some (subg. Ornithophila) even have them fully developed. In Ornithomyia, on the other hand, most of the species have distinct ocelli, but in a few (subg. Pseudorni- tliomyia) these organs are atrophied. Attempts have been made to correlate the atrophy or loss of ocelli in the Hippoboscidae with the reduction of the compound eyes and also with the loss of the power of flight (Massonat, 1909). There are, however, many ex- ceptions to these supposed correlations, Olfersia, Pseudolynchia, Stilbometopa, Ortholfersia, and Hippobosca being all without even traces of ocelli, yet fully-winged and active fliers, with large com- pound eyes. Nevertheless, it might be of some significance that, of the five genera in which the wings are either atrophied, reduced or lacking, only one (Stenepteryx) has retained ocelli, moreover of small size or vestigial. There is also a possibility that even in some of the Hippoboscidae with ocelli, these organs may be func- tionless, since they are frequently very small and flat. In insects in general the function of the ocelli is controversial. From their structure they cannot receive an image ; most likely they merely register changes in intensity of illumination. This might be of some assistance before and during flight by accelerating the insect ’s response to shadows and by keeping the body back uppermost (Kalmus, 1945; Parry, 1947). The antennae (A) are, with the male terminalia, the most modi- fied external organs of the Hippoboscidae ; yet it is possible to homologize their structure without much difficulty with that of the higher Muscoidea. They are apparently immovable, placed far forward, near the apical margin of the frontal area, and in most original sketches by Dr. Jobling: E, interocular face and frontal area from a cleared specimen mounted on a slide without any pressure on the cover slip ; F, ideal cross-section of anterior portion of head: CIP, cibarial pump with lateral plate or bars; CL, clypeus; FB, frons; MV, mediovertex; OCI, ocelli; pt, ptilinum; pts, ptilinal suture; PV, postvertex; BM, rostrum membrane; V , vertex. G-, Lipoptena rusaecola J. Bequaert, male; head from behind. H, Hippobosca equina Linnaeus; 6 successive stages of the biting process, in side view, after Hase (1928). 25 ENTOMOLOGICA AMERICANA cases (except in Ornithoica) well separated by the flattened frontal carina. In Ornithoica they lie in a single frontal cavity; in the other genera each of them has its own deep antennal pit ( AP ), sur- rounded by a partial or complete rim. The first antennal segment (IAS; scape) is readily recognized in the more primitive Orni- thoica (Fig. 5C), where it is large and flattened, completely sepa- rated by a deep suture from the lunula, and with strong setae as usual in the higher Muscoidea. In most other genera it is more or less fused with the sides of the lunula, although its position may often be traced by the characteristic setae. In the Melo- phaginae, there is no external indication of this segment; accord- ing to Jobling, in Melophagus ovinus it is deeply merged in the head capsule and its dorsal part, overlapped by the anterior border of the antennal pit, is devoid of setae. The second antennal seg- ment (2AS ; pedicle) comprises the major part of the antenna, be- ing much flattened. Ventrally it is hollowed out into a pouch con- taining the greatly reduced third antennal segment (3 AS ; flagel- lum), hidden from view except for the protruding arista (AR). The shape of the second segment varies greatly and will be de- scribed under the several genera. In the Ornithoicinae (Fig. 5C) and Ornithomyiinae (Fig. 11F), the dorsal face of the second seg- ment is extended into a flattened prolongation, the antennal ap- pendage or process •( AA ) , always bearing many setae and bristles, particularly at the apex. The extreme development of these ap- pendages is reached in Ornithoctona and its subgenus Ornithop- ertha. The Ortholfersiinae have small antennal appendages, while in the Alloboscinae (Fig. 5D) they are of a peculiar form. In the Hippoboscinae (Fig. 5B) and Melophaginae (Fig. 4C), the second segment lacks the appendage, being small, subglobular, only partly filling the antennal pit. The arista (AR) of the third segment varies in shape, spatulate at the tip in Ornithoica, Ornithomyia, and Pseudolynchia (Fig. 11F), dichotomously branched in Hippo- bosca, bifid with comb-like branches in Melophagus, and variously divided in other genera. In nearly all true Muscoidea Calyptrata the dorsal surface of the second antennal segment shows at least a trace of a longitudinal slit or furrow extending some distance from the apical margin. The slit is lacking in all true Acalyptrata. This character is regarded by many dipterists as of paramount importance in classification and in tracing the relationships of otherwise puzzling forms. Hennig (1941) investigated it in the Diptera Pupipara, concluding that the slit occurs in at least some genera of the Streblidae, Nycteribiidae 26 Volume XXXII and Hippoboscidae and that all three families are therefore modified Calyptrata. According to him, whenever present the position of the slit marks the true dorsal surface of the second antennal segment, regardless of its shape. The occurrence of the slit has actually been investigated in only a few hippoboscid genera. Jobling (1926) describes and figures the antennal furrow ( AF ) for Pseudo- lynchia canariensis (- L. maura), where it is well developed and extends to mid-length of the segment along the dorsal side of the appendage (Fig. 11F). He found it also in Hippobosca, but states that it is lacking in Melophagus. Hennig (1941) regards it as a primitive characteristic of the Hippoboscidae and explains its ab- sence in certain genera as the result of closing up following the progressively stronger sclerosis of the integument. Jobling (1926) points out that the hippoboscid antenna is greatly modified no doubt as a sequel to permanent ectoparasitism. He considers that the modifications serve to protect the most important parts of the antenna, namely the third segment and its arista, when the flies move amidst the feathers or hairs of the host. The protec- tion is ensured by the deep antennal pits and the retraction of the third segment within the second. The bristly appendage of the second segment, which occurs in the genera with partly open anten- nal pit or pits, may also shield the arista from contact with the feathers or hairs. As the main function of the third segment and arista is believed to be olfactory, I suggest, in addition, that the protection is mainly against the exudates and particles of epidermis and dirt which often gather on the surface of the host ’s skin. Such extraneous matter might easily clog up the openings of the olfactory receptors when the fly feeds. I suggest furthermore that the re- markable development and hairiness of the second segment do not merely protect the third segment, but that they also have an active function. The second antennal segment of insects is known to con- tain the peculiar organ of Johnston, apparently an important tactile sense organ, registering movements of the distal part of the antenna. In most Hippoboscidae it is eminently suited for this purpose, its surface being enlarged, produced forward and beset with many long and short tactile setae. In the dark recesses of a bird’s feathers or a mammal ’s hairs a fly must rely mainly on the tactile sense to direct its movements. In comparison the sense of smell may be relegated to second place in the life of the fly, accounting for the great reduc- tion of the third antennal segment. The mouth-parts (Fig. 7A-C ; after Jobling) are modified into an extremely efficient piercing and sucking proboscis, which, how- 27 ENTOMOLOGICA AMERICANA CIP Fig. 7. A-D, P seudolynchia canariensis (Macquart) ; mouth-parts and adjoining structures, modified after Jobling (1926) : A, longitudinal section with retracted proboscis ; B, the same with fully extended proboscis ; C, cleared preparation with proboscis partly everted; D, cross-section through middle part of haustellum: CIP, cibarial pump; CL, clypeus; FE, frons; FTJ, fulcrum; H, hyoid; H A, haustellum; EY, hypopharynx; LA, labella; LEP, labrum- 28 Volume XXXII ever, is duplicated elsewhere in the Diptera, though not exactly in the same manner. The soft basal rostrum membrane ( EM ; basi- proboscis), which may be invaginated within the head capsule, bears the several sclerites of the tentorium, called by Jobling the fulcrum ( FTJ ), the hyoid ( H ), and the stipites (ST). The presence of a hyoid (Frey’s theca), a small sclerite connecting the fulcrum with the base of the labium (praementum), is considered by Frey (192], p. 208) a decisive character of calyptrate Muscoidea, which he calls therefore the Thecostomata. The Muscoidea Acalyptrata generally lack a hyoid, for which reason Frey calls them the Haplostomata or Athecostomata. It is somewhat open to question, however, whether the mere presence or absence of a hyoid has such a fundamental im- portance. Jobling regards its presence as an adaptive feature, correlated with the protractility of the proboscis. The matter is discussed at some length by Hennig (1941, p. 239). In any case, the hyoid is present in the Hippoboscidae, while, according to Job- ling, it is present in the Nycteribiidae and absent in the Streblidae. On the sclerites of the tentorium are inserted the muscles of the ros- trum which protract and retract the proboscis. The two single- jointed palpi (PA ; probably the maxillary palpi) protrude from the rostrum beyond the anterior margin of the head. They are almost always well developed and compressed, with con- cave inner surface, their chief function being to form a protecting sheath around the haustellum. As they usually bear prominent sensorial setae, they have no doubt also a tactile or chemoreceptive function. They are, however, very short and possibly functionless in Allobosca, of Malagasy lemurs, and vestigial in E chesty pus, of African antelopes. The haustellum (HA), or sucking tube of the proboscis, is long and narrow, strongly sclerotized. At rest its needle-like distal part lies concealed in the palpal sheath ; its bulbous, slightly ovoid base may be protracted or retracted, being withdrawn far within the rostrum membrane and the head capsule when the proboscis is not in use. The haustellum consists of the modified labium (LAB), which encloses the labrum-epipharynx (LEP) and the hypopharynx (HY). These three parts are closely joined (Fig. 7D) and enter the skin as a unit, when the fly bites. The labium ends in two epipharynx; LG, labial gutter; P, pharynx; PA, palpus; pts, ptilinal suture; EM, rostrum membrane; SD, salivary duct; SIV, salivary valve; ST, stipes; TH, theca; V, vertex. E, Glossina palpalis (Robineau-Desvoidy) ; cross-section through middle of narrow part of haustellum, modified after Jobling (1933) ; lettering as in Fig. 7D. F, Olfersia coriacea Macquart, female; frontal area and antennae. 29 ENTOMOLOGICA AMERICANA lobelia (LA; possibly modified labial palpi), provided at tbe tip with a double crown of prestomal teeth (PUT ; Figs. 6A-B), which act as the cutting organs when the proboscis bores in the skin. Ac- cording to Frey (1921) and Hennig (1941), true prestomal teeth are also characteristic of the Muscoidea Calyptrata. In all Hippo- boscidae the labella are very short as compared with the remainder of the haustellum. The labium is divided over its entire length into two parts, a ventral theca (TH) and a dorsal labial gutter (LG), as shown externally by a distinct furrow running on each side along the needle-like part of the labium. The two side walls of the labial gutter are curved upward over the labrum-epipharynx which they enclose more or less completely. The labrum-epipharynx is stylet- shaped, with sharp point, and is a nearly closed tube, with a longitu- dinal lower (or inner) slit obliterated by the adjacent hypopharynx, thus forming a complete suctorial tube. The outer lateral walls of the labrum-epipharynx are covered with minute teeth, which inter- lock with similar teeth on the inner walls of the labial gutter (Fig. 7D). The hypopharynx is very slender, but somewhat flattened. Its median, thicker portion is perforated over the entire length by the salivary duct (SD),. which opens in the labial gutter on the dorsal side of the hypopharynx at its narrow distal end. The struc- ture of the proboscis and palpi shows relatively little variation within the family, except in the size of the parts and in minor de- tails. These organs may therefore be regarded as inherited without much change from a common ancestral stock, before the family began to break up into a number of subsidiary groups or sub- families. The structure of the mouth-parts is the same in both sexes of the Hippoboscidae, as both suck blood. This is true also of the other Pupipara, the Glossinidae and the non-pupiparous blood- sucking Muscoidea. The foregoing description of the head capsule and mouth-parts follows in the main Jobling (1926), who reviewed, corrected and completed the earlier work of Dufour (1825; 1844; 1845), Muggen- burg (1892), Massonat (1909), Roberts (1927), and others. Job- ling studied mainly Pseudolynchia canariensis (= Lynchia maura) and Melophag us ovinus. Gouin (1949, pp. 244-248) recently exam- ined again Hippobosca equina and Melophagus ovinus. As men- tioned before, the area which Jobling had called “ f ronto-clypeus ” (a terminology adopted in my former papers) is not a composite region, as he thought, but represents the frons only (Figs. 6E-F). As in the Muscoidea, the true clypeus (CL) is the very small sclero- 30 Volume XXXII tized area between frons and rostrum membrane, which Jobling formerly called the torma. (See also Snodgrass, 1935, p. 322; and 1943, pp. 36-38 and p. 46, fig. 171). The depressed thorax, more than any other single feature, gives the Hippoboscidae their characteristic louse-like or tick-like ap- pearance. In side view it is often only one-third as high (or thick) as the width of the mesonotum, in contrast to the higher Muscoidea, where it is at least as high as the mesonotal width. In the process of flattening, the sides of the thorax have folded outwardly at two levels, the intervening area of the mesopleura being flat or even con- cave. The upper or dorsal fold, at about the level of the wing and more or less horizontal, is the stronger and forms a rounded edge or blunt ridge. The lower or ventral fold, at about the upper level of the fore and mid coxae and apparently near the position of the anepisternal suture (as) of other Diptera, is often very weak and usually slanting. As a result of the upper fold, the dorsal, horizon- tal surface of the thorax comprises, in addition to the true notum, the upper portion of the mesopleurites, which in the higher Mus- coidea occupy the vertical sides, while the prothoracic spiracles are placed more dorsally. The sternal area also is deeply altered by the flattening. In the higher Muscoidea (Fig. 9B), the sternum and particularly the mesosternum are relatively unimportant, the coxae of each pair being inserted close together, while the legs are strictly ventral. In the Hippoboscidae (Fig. 9A), the sternum is very ex- tensive and the coxae of each pair are widely separated, so that the legs are placed ventro-laterally, allowing them to stretch out and move in a near-horizontal plane. To further complicate matters, some of the sutural lines commonly used as landmarks, but already greatly modified in the higher Muscoidea, have become obliterated. To trace homologies under such conditions is fraught with hazards and may be sometimes pure guesswork. Dorsally (Fig. 8A) the pronotum has a very short median sclerite, or protergum ( PT ), usually hidden by the occipital margin. In some cases the pro-mesonotal suture ( pms ) is very superficial or obsolete. In the Ornithoicinae and Hippoboscinae (Fig. HE), the protergum is short, but dorsally visible between head and meso- scutum. Laterally the propleuron (PP), on which the fore coxa articulates, is sometimes partly visible from above, when the head is free from the thorax; more often it is covered by the projecting angle of the humeral callosity (HC ; humerus or postpronotum) . This callosity is as a rule separated, at least partly, from the pre- scutum by the posthumeral suture (phs) and bears posteriorly the 31 ENTOMOLOGICA AMERICANA Fig. 8. A, Ornithoctona erythrocephala (Leach), female; thorax from above. B, Glossina fusca Walker, female; thorax from above. ACO, axillary cord; AES, anepisterimm ; EPI, epipleurite; lib, humeral bristles; SC, humeral callosity; hdb, posterior dorso-central bristle; idb, posterior acrostichal bristle; mns, medium notal suture; mpb, mesopleural bristles; MS, mesoscutum; msb, discoscutellar bristle; nob, notopleural bristles; NP, notopleuron ; pab, postalar bristles; pb, presutural bristles, PC, postalar callus; PCL, lower calypter; pits, posthumeral suture; POS, prothoracic spiracle; PS, prescutum; PSC, parascutellum ; PT, protergum; sab, supraalar bristle; scb, scutellar bristles; SCTJ, scutellum; ss, scutoscutellar suture; tms, transverse mesonotal suture. 32 Volume XXXII conspicuous, more or less dorsally placed prothoracic spiracle ( POS ). Whether the humeral callosity includes in addition some other propleural sclerite (episternum or epimeron) is open to question. The mesonotum occupies the major part of the dorsum. In Ornithoica , Ornithomyia, and Ornithoctona (Fig. 8A), where it is most similar to that of the Muscoidea, it is divided behind mid- length by a straight, transverse mesonotal suture ( tms ), often somewhat interrupted medially, into an anterior prescutum {PS) and a posterior mesoscutum {MS). It may also show a longitudi- nal superficial median notal suture (mns), running from the ante- rior margin to close to the scutellum. A narrow, wedge-shaped notopleuron { NP ), on each side before the mesonotal suture, is set off by deep grooves from the prescutum and mesopleuron. In other genera the mesonotal suture undergoes various changes or dis- appears. In Lipoptena (Fig. 11A), the median notal suture is indicated only anteriorly; instead the mesonotum bears a pair of superficial longitudinal intrascutal grooves ( lig ). Likewise the notopleuron may be partly or completely fused with the prescutum, its position being then indicated only by the insertion of the noto- pleural bristles {nob ; posthumeral bristles) ; in Hippobosca (Fig. 11E), Lynchia, and Olfersia the fusion is complete; it is as yet in- complete in some species of Lipoptena (Fig. 11A) and Stilbometopa. Postalar calli {PC) are rarely or only superficially set off at the outer hind corners of the mesoscutum, although postalar bristles {pab) are often present. The large dorsal sclerite, on each side between the prothoracic spiracle and the base of the wing, is the upper portion of the mesopleuron, or anepisternum (AES), which has moved to a horizontal position. My former interpretation of this sclerite in Lipoptena as the notopleuron (1942a, p. 9 and fig. IE, NP) was erroneous and is corrected in Fig. 11 A. The stiff setae near its hind margin or scattered over the surface, correspond to the upper mesopleural bristles {mpb) of the Muscoidea. The two small sclerites often found between the hind margin of the anepis- ternum and the root of the wing, may be modified epipleurites {EPI). Exceptionally, as in Melophagus, all mesonotal sutures are nearly obliterated. The scutellum {SCTJ), often large, is limited anteriorly by a scuto-scutellar suture {ss), usually deep and complete. In Melo- phagus, where the scutellum is small, it is nevertheless set off by a shallow suture. It generally shows a longitudinal depression, sometimes widened triangularly. Laterally it continues in an ante- 33 ENTOMOLOGICA AMERICANA rior and a posterior ridge, enclosing on each side a more or less grooved area, the parascutellum (PSC). In most Hippoboscidae the anterior ridge is little developed or is fused with the hind mar- gin of the postalar callus ; it is well defined in Ornithoica. The postscutellum ( POTJ ; postnotum), lies behind and below the scutel- lum from which it is rarely separated by a narrow subscutellum (SSL) in the Hippoboscidae; but its two lateral areas, the pleuro- tergites (PTG) of the metanotum, are sometimes much swollen or bear characteristic pleurotergal processes . Below the postscutellum lies the metanotum (MTN), usually developed on the sides only or fused with one of the adjoining sclerites. The reduction of the sternal area in most Diptera, particularly pronounced in the calyptrate Muscoidea, and the resulting near- contiguity of the coxae in each pair are the extreme development of a peculiarity common to all Neuropteroidea. In these insects part or most of the primitive or true sternum, with the paired inter- nal sternal apophyses, are inflected on the middle line by so-called cryptosterny, producing a forked endoskeletal apodeme, the furca , usually traced outwardly as a median sternal or fur cal suture ( ful , fu2 , fu3). In the more primitive winged insect thorax, the bases of the sternal apophyses are often connected by an internal sterna- costa, forming externally a sternocostal suture which divides the sternum proper into a presutural basisternum (BSl, BS2 , BS3) and a postsutural fur cast ernum ( FSl , FS2, FS3 ; sternellum) • anterior to the basisternum, an additional small median sclerite, the pre- sternum (PSl, PS2, PS3), is sometimes detached. In the Diptera, however, these several areas have undergone many shifts; some of the original sutures have disappeared and new ones have second- arily taken their place, so that it is extremely difficult and perhaps even impossible to trace the true sternal homologies. This will be realized from a comparative study of the sternum of Glossina fusca (my Fig. 9B), Callipkora (Snodgrass 1935, p. 171, fig. 95B), and Musca (Hennig, 1941, pi. 9, figs. 1-2). In all these figures a partly developed longitudinal suture runs forward on each side from the insertion of the mid coxae (weakest in Musca). This may possibly be a remnant of the original division between the sternum and the pleura or laterotergites. It should also be noted that the mid coxae are more widely separated from each other in Glossina than in either Musca or Calliphora , no doubt owing to the slight flattening of the thorax of the tsetse-fly. Neither Snodgrass (1935) nor Crampton (1942) attempt to trace the homologies of the sternal regions of the Diptera. Cramp- 34 Volume XXXII Fig. 9. A, Ornithoctona erythroceyhala (Leach), female; thorax from below. B, Glossina fusca Walker, female; thorax from below. BS1, basister- num of prothorax; BS2, basisternum of mesothorax; BSS, basisternum of metathorax; COl, C02, COS, fore, mid and hind coxae; FSS, furcasternum of metathorax; fu2, f ureal suture of mesosternum; fu3, f ureal suture of meta- sternum; HAL, halteres; HC, humeral callosity; PCL, lower calypter; PP, propleuron; PS1, presternum of prothorax; PSL, prosternal lobes. 35 ENTOMOLOGICA AMERICANA ton merely labels each of the three sternal divisions of Ornithoctona as basisternnm (bs) on a side view of the thorax (1942, fig. 6E). He also distinguishes as furcasternum (fs) a posterior area of the prosternum ; I am unable to find a suture separating such a region in Ornithoctona erythrocephala. The only study of the hippoboscid sternum now available is that by Hennig (1941). His suggestions as to the internal changes involved in dipterous cryptosterny and their relations to the later, secondary widening of the sternum in the Pupipara deserve serious consideration. He does not, however, propose a terminology that might be used for the several areas, some of which are of great importance for the taxonomy of these flies. It is rather unfortunate that he selected Hippobosca for his study, as this is one of the more specialized of the Hippoboscidae. The thorax of the Ornithoicinae (Ornithoica) , the most primitive of the Recent Hippoboscidae, is less depressed than that of the other subfamilies, and the three main divisions are more clearly defined. On the sternum seen from below (Fig. 12B), some of the pleural areas are visible on each side, and, in the mesothorax, the lower portion of the anepisternum (AES) is separated from the sternum proper by a slight depression, but without any real suture. The prosternum consists of a single, moderately wide, trapezoidal, intercoxal basisternnm ( BSl ), without longitudinal depression; in front of it the cervical membrane has no additional sclerite. The mesosternum comprises a single sclerite, presumably the combined basisternum ( BS2 ) and furcasternum ( FS2 ), the lateral portions of the latter being set off as triangular areas in the hind corners near the mid coxae by very weak transverse depressions (without su- ture) ; a median fur cal suture (fu2) runs the whole length of the mesosternum. Young (1922) regards the area here called the mesosternum as corresponding to the lower portions of the episterna (sternopleura) , which may have crowded out the true basisternum and furcasternum. This view may be morphologically correct and would fall in line with Hennig ’s (1941) contention that, when the coxae of the Hippoboscidae moved apart, they merely shifted laterally, presumably within the pleural sclerites, all the true sternal areas having become permanently absorbed in the endoskeletal furca. The metasternum consists of two sclerites: the anterior, larger area, or basisternum ( BS3 ), is divided throughout by a deep median f ureal suture ( fu3 ) ; the posterior, very small portion, or furcasternum (FS3), is undivided. The sternum of the other Hippoboscidae fits in the same general pattern, as a comparison with Ornithoctona (Fig. 9A), Hippobosca 36 Volume XXXII Fig. 10. A, Ornithoctona erythrocephala (Leach), female; thorax in side view. B, Glossina fusca Walker, female; thorax in side view. AES, anepis- ternum; as, anepisternal suture; B S2, basisternum of mesothorax; BS3, basis- ternum of metathorax; COl, C02, COS, fore, mid and hind coxae; HAL, halteres; HC, humeral callosity; hpb, hypopleural bristles; KES, katepister- num ; MAP, metapleuron ; me, membranous cleft of pleuron ; MES, metathoracic spiracle ; mpb, mesopleural bristles ; MPL, meropleurite ; MS, mesoscutum ; MTN, metanotum; NP, notopleuron ; POS, prothoracic spiracle; POTJ, post- scutellum; PP, propleuron; PEC, precoxal area; ps, pleural suture; PS, prescutum; PTG, pleurotergite ; SCJJ, scutellum; SEP, supraepimeron ; SF, subalifer; SSL, sub scutellum ; stb, sternopleural bristles. ENTOMOLOGICA AMERICANA Fig. 11. A-B, Lipoptena depressa (Say), female; thorax from above (A) and below (B) : aab, anterior acrostichal bristle; adb, anterior dorso-central bristle ; AES, anepisternum ; BS2, basisternnm of mesothorax ; BS3, basisternum of metathorax; FS3, furcasternum of metathorax; fu2, f ureal suture of meso- sternum; f%3, furcal suture of metasternum; leb, latero-central bristles; lig, longitudinal intrascutal groove; mns, median notal suture; MS, mesoscutum; 38 Volume XXXII (Fig. 11D), Lipoptena (Fig. 11B), and Melophagus will show; the mesosternum extends farther to the sides, thus hiding the pleural areas. The prosternum and metasternum also widen and there are various other minor changes. In Lipoptena and Melophagus, the prosternum is deeply emarginate anteriorly; in Ornithoctona and other Ornithomyiinae, it is divided into a pair of lobes or prongs, the prosternal lobes ( PSL ) , directed forward. Hippobosca is pecul- iar in that the hind lateral areas of the mesosternum (sides of the furcasternum) are separated from it by a suture and fused with the basisternum of the metathorax. The pleura of the Hippoboscidae are best understood by com- paring them with those of the higher Muscoidea, in which they are moreover already greatly modified from those of more primitive winged insects. The following areas may be recognized in Glossina fusca (Fig. 10B). The propleuron (PP) is very short and incom- pletely divided from the mesopleuron. In the more primitive winged insects, a pleural suture runs a fairly simple, oblique course from the pleural wing process to the pleural coxal process and di- vides the mesopleuron into an anterior episternum and a posterior epimeron. In the Muscoidea, however, the pleural suture ( ps ) fol- lows a zigzag course, with two angular bends, dividing it into three sections: the first (anterior) bend sends forth a slightly slanting anepisternal suture (as), which divides the mesepisternum into an anterior (upper) anepisternum (AES) and a posterior (lo’vyer) hat episternum ( KES ), the latter fused with the lower precoxal area (PRC); at the second (posterior) bend, an upward running suture divides the mesepim,eron also into an anterior (upper) su- praepimeron (SEP; anepimeron) and a posterior (lower) infra- epimeron, the latter fused with the meron of the mid coxa to form a meropleurite (MPL). A vertical membranous cleft (me), in front of the upper section of the pleural suture, extends as far NP, notopleuron ; PC, postalar callus; PP, propleuron; PS, prescutum; psb, prescutellar bristles; PSL, prosternal lobes; PT, protergum; SCTJ, scutellum; tms, transverse mesonotal suture. C-E, Hippobosca equina Linnaeus, female; thorax in profile (C), from below (D) and from above (E) : as, anepisternal suture; BS1, basisternum of prothorax; COl, C02, C03 fore, mid and hind coxae ; hdb, posterior dorsocentral bristle ; HC, humeral callosity ; me, mem- branous cleft of pleuron; MES, metathoracic spiracle; PCL, lower calypter; pms, promesonotal suture; POS, prothoracic spiracle; ps, pleural suture; PS1, presternum of prothorax; PIG, pleurotergite ; SF, subalifer; ss, scuto-scutellar suture; other lettering as in Fig. 11A-B. F, Pseudolynchia canariensis (Mac- quart) ; left antenna from the side, after Jobling (1926) : AA, antennal ap- pendage ; AF, antennal furrow ; AP, antennal pit ; AR, arista of third segment ; IAS, 3AS, 3AS, first, second and (outline of) third antennal segments. 39 ENTOMOLOGICA AMERICANA down as the anepisternal suture. Some small sclerites are wedged in the upper corner of this cleft, between the pleural suture and the base of the wing; the most conspicuous of these is the subalifer (SF). The metapleuron (MAP), below the metathoracic spiracle (MES), is fused anteriorly with the meropleurite. The combined metapleuron, meron of mid coxa and infraepimeron of the meso- pleura of the Muscoidea form the area usually called the hypo- pleuron by dipterists and bearing the hypopleural bristles (hpb). The insertion of the halter (HA) marks the separation of the meta- notum (MTN) from the pleurotergites of the postscutellum. In Ornithoctona erythrocephala (Fig. 10A), most of the sutures and areas of the pleura of Glossina may be recognized, but are some- what modified. The two bends of the pleural suture are more angu- lar, particularly the lower one; while the anepisternal suture is nearly vertical, extending the first section of the pleural suture downward. The metapleuron appears to be very superficially di- vided from the meropleurite of the mesopleura. The subalifer (SF) is greatly developed and wedged in between the membranous cleft and the anepisternal suture. In most other genera of Hippo- boscidae, for instance in Hippobosca (Fig. 11C), the sclerites of the pleura are more difficult to trace, some of the sutures becoming obsolete, particularly in the subapterous forms. In Melophagus nearly all dorsal and lateral sutures of the thorax have disappeared, although a few are retained on the sternum ; the dorso-ventral flat- tening has even affected the metapleura, so that the metathoracic spiracles also are in a dorsal position. It would seem meaningless to trace any homologies of parts or areas in such a highly special- ized complex. The legs (Fig. 12C-E) are robust, with rather swollen femora, flattened tibiae and short, compact tarsi. They vary in length and are on the whole shorter and stouter in the mammal-inhabiting forms than in those of birds. In the latter the legs are adapted to gliding and scurrying about swiftly, forward, backward, as well as sideways, amidst the soft feathers ; in the former they are built more for grasping and adhering to the skin and to the coarse hair of the pelt. The coxae (COl, CO 2 and COS) are large and swollen; those of the fore legs are more or less balloon-shaped and freely movable, so that they may be directed downward as well as side- ways; but the mid and hind coxae are flattened, horizontal and sometimes nearly immovable. The trochanters (TB) of the fore legs are small, as usual in the Diptera; those of the mid and hind legs are larger and movable on both coxa and femur, obviously to 40 Volume XXXII compensate for the relative fixity of their coxae. The femora (FE) and tibiae ( TI ) show no unusual features. The tibial spurs ( TSP ) are sometimes difficult to- trace amidst the apical bristles. The tarsi {TAR) consist of five short segments and a terminal pretar- sus (de Meijere; Ockler’s “Krallenglied”). The basal segment or basitarsus {BTA) of the hind tarsi is sometimes more elongate than the others; the fifth segment or distitarsus {BTA) is always broad and depressed. The pretarsus consists of a membranous, elastic base (Ockler’s “ abschliessende Haut”), which bears a median, slender, often hairy or feathered empodium ( EMP ; Ockler’s “ Streckborste” ; de Meijere ’s “ Sohlenf ortsatz ? ’ or “Sohlenbor- ste”), flanked on each side by a broad, flap-like pulvillus {PU; ‘ ‘Haftlappchen” of Ockler) and a claw {UN; unguis; Ockler’s “Kralle”). A small, transversely striate, median, sclerotized un- guitractor plate ( UTR ; Ockler’s “ Streckplatte ” ) lies in the mem- branous base on the under side of the pretarsus; attached to its proximal end is the tendon of the retractor muscle of the claws. In all the Hippoboscidae, of mammals as well as of birds, two claws are present on all legs. They are unusually developed and consist of a broad, flattened base or heel {HE), followed by one or two teeth {TO). The heel has a basal protuberance hinged to the dorsal apical margin of the distitarsus on either side of a median unguifer process {UP ; Ockler’s “ Krallengelenkhocker ”) ; its apical portion is prominent and sometimes prolonged, simulating an ad- ditional tooth. Simple claws (Fig. 12F), ending in one tooth, oc- cur in all mammal-infesting flies (Hippoboscinae, Melophaginae, Ortholfersiinae, and Alloboscinae) and in the bird-flies of the sub- family Ornithoicinae ; their claws are sometimes said to be bidentate or bifid (Meigen, 1830, p. 230; Kieffer, 1900, p. 338), owing to the greatly developed, tooth-like basal heel, which is strongly sclerotized and darkened along the inner edge. The remaining bird-flies (Or- nithomyiinae) have claws with two separate teeth, in addition to the basal heel (Fig. 12K) ; as the basal heel is even more elongate than that of the simple claws, the claws of the Ornithomyiinae are sometimes called by error tridentate or trifid (Kieffer, 1900). Ac- tually, the additional tooth is due to the cleaving of the basal heel ; in some of the flies with simple claws the basal heel shows a deep longitudinal groove. The claws are mobile and each of them can move independently in a pair. The pulvilli of the Hippoboscidae also differ markedly from those of other Diptera. As Ockler (1890, pp. 33 and 36) pointed out, they show peculiar adaptations to ectoparasitic life amidst fur 41 ENTOMOLOGICA AMERICANA Fig. 12. A, Myiophthiria lygaeoides Rondani, female; wing. B, Ornithoica vicina (Walker), female; thorax from below: AES, anepisternum ; BS1, BS2, BS3, basist.ernum of pro-, meso-, and metathorax; COl, C02, COS, fore, mid and hind coxae; ES2, FS3, furcasternum of meso- and metathorax; fu2, fu3, furcal suture of meso- and metasternum; EC, humeral callosity; PP, propleura. C-F, Eippobosca equina Linnaeus, female; C-E, fore, mid and hind legs: CO, 42 Volume XXXII and plumage. The fully-extended pulvillus is a lengthened, tender and soft flap inserted on a long, narrow, sclerotized auxiliary plate ( AUP ; Ockler’s “ Stiitzplatte ” ) attached to the membranous base of the pretarsus. When the pulvilli function they are projected forward between the claws, which simultaneously are raised and spread out, their bases being thrown back over the dorsal apex of the distitarsus. The claws are lowered when in use and at the same time the pulvilli are folded back beneath the apical area of the distitarsus where they lie spread out flat, side by side, and are pro- tected by the prominent, broad apical portion of the heels of the claws. In some, but not in all louse-flies of mammals, the pretarsus is asymmetrical, the claws being unequal in ' each pair, one being thicker or shorter ( J. Bequaert, 1942a, p. 68 ; fig. 5E, inner and outer claw of fore leg of Lipoptena cervi) . The unequal claws were first described by Meigen (1830, p. 230) for Lipoptena cervi (= Or- nitkobia pallida). In most hippoboscids with unequal claws the pulvilli also are unequal in each pair, as de Meijere (1901, pp. 446 and 467) noted for Lipoptena cervi, where the larger pulvillus ad- joins the shorter claw. This is true also in most species of Hippo- bosca (such as H. equina); in others one of the pulvilli is rudi- mentary. Hippobosca camelina is unique in the genus in that the very long claws are equal in each pair, while both pulvilli are com- pletely aborted. The wings vary in size and shape. In the majority of the genera (15 out of 20) and species (107 out of 124), they are well developed and functional, being relatively long and broad for the size of the body. Such fully-winged flies occur on birds as well as on mam- mals, and both types of hosts also harbor species with more or less reduced wings, no longer used for flight. The normally subapterous forms belong to 4 genera, with some 15 species. Melopkagus, con- taining two species on mammals, is the only genus with completely atrophied wings, reduced to minute, elongate, setigerous knobs. In 3 of the winged genera, with 18 species on mammals, related to Melopkagus, the newly emerged fly has fully-developed and func- coxa; FE, femur; TAB, tarsus; TI, tibia; TB, trochanter; TSP, tibial spur; UN, claw; F, smaller claw and adjoining pulvillus of fore leg: UAP, auxiliary- plate; EE, heel; PU, pulvillus; TO, tooth. Or, Stenepteryx hirundinis (Lin- naeus), female; wing. H, Crataerina pallida (Latreille), female; wing. I, Allobosca crassipes Speiser, female; wing. J, Ornitheza metallica (Schiner), female; base of wing. K, Ornithomyia fringillina Curtis, female; distitarsus and pretarsus: AUP, auxiliary plate; DTA, distitarsus; EMP, empodium; HE, heel of claw; PU, pulvillus; TOl, T02, double teeth of claw. 43 ENTOMOLOGICA AMERICANA tional wings, which break off close to the base after the final host is reached. When it is most complete, as in Ornithoctona and Ornitho- myia (Fig. 13A), the venation is similar to that of the Myiodaria Calyptrata, except for two minor peculiarities. The normal mus- coid venation is itself greatly reduced, as compared to that of the lower Nematocera. The resulting difficulty in tracing the homolo- gies of the veins is particularly felt in the Ilippoboscidae, and is even enhanced in Lipoptena, Neolipoptena, and E chesty pus, which show the greatest reduction of veins in a fully-developed wing of any of the Cyclorrhapha. For descriptive purposes I prefer to a so-called morphological terminology one of the purely conventional terminologies of veins and cells of the older authors. I find this less confusing than the now popular notations derived from the Comstock-Needham system, based on the real or supposed homolo- gies of the dipterous venation with that of other insect orders. Comstock applied his notations in their present form to a hippo- boscid for the first time in 1924 (p. 874, fig. 1118, labelled Olfersia sp. ; actually of a Lynchia, presumably L. americana) . Soon after- ward, Tillyard (1926, p. 378, fig. W73) offered a modification shown for the wing of an Australian species of Ornithomyia which he called 0. australasiae, but which was probably not Fabricius’ spe- cies of that name. In this, the 4th longitudinal became M1 (instead of M1 + 2), the 5th longitudinal M3 + Cux (instead of if3 + 4), and the 6th longitudinal Cux + lAn (instead of 1st A). Ferris and Cole (1922, p. 196) stated that their interpretation was in accord with the Comstock-Needham system, presumably as first published with a different type of notations by J. H. and A. B. Comstock in 1895 (p. 488, fig. 595) ; but Comstock’s vein 1st A becomes Cu2 in their fig. 12 supposedly of Lynchia americana (more probably L. fusca of California) and 2nd A + Cu2 in their fig. 18 of Stilbometopa impressa, where in addition the cross-vein closing the anal cell is labelled as part of Cu2. In 1925 (p. 418, fig. 4) Ferris used again Cu2 + 2nd A in the wing of Ornitheza metallica for Com- stock’s 1st A, while the axillary vein behind it is labelled 3rd A (instead of 2nd A). More involved changes of the Comstock nota- tions were proposed recently by Lower (1951, p. 81, fig. 22, wing of Ornithomyia sp.). It would seem pointless to discuss more in detail the several emendations of the Comstock system proposed in this country and abroad, since my remarks are not intended as criticism, but merely show the lack of agreement in the matter. Nevertheless, I have inserted in the following list of terms the 44 Volume XXXII equivalent notations of the original Comstock system and some of the suggested changes, so that the reader may use them, if he wishes. Costa ( CO in Fig. 13A; C of Comstock), strongly developed along the anterior margin and ending at the tip of the 3rd longitu- dinal vein or slightly beyond it. The apical and posterior margins never show a trace of an ambient vein. The costa always bears many setae, some often very long, particularly toward the base. A costal spine or bristle, at the termination of the subcosta, is rarely present (in Ornithoica) . Subcosta (SC in Fig. 13A and of Comstock; mediastinal or auxiliary vein), sometimes not continued to the costa. A short humeral cross-vein (h in Fig. 13 A; h or hm of Comstock; Speiser’s “basale Querader”; Townsend’s first subcostal vein) connects it with the costa a short distance from the base. First longitudinal vein (1 in Fig. 13A; R1 of Comstock; R1 + 2 of Lower, 1951; Speiser’s Subcostalis). Second longitudinal vein (2 in Fig. 13A; R2 + 3 of Comstock, 1924; R3 of Lower, 1951; radial vein; Speiser’s Radialis), some- times running close to or partly fused with the costa, apicallv (in Ornitheza , Fig. 14D). Third longitudinal vein (3 in Fig. 13A ; R± + 5 of Comstock, 1924 ; R* of Lower, 1951; cubital vein; Speiser’s Cubitalis), sometimes more or less confluent with the costa apically (in Ornithoica) . Fourth longitudinal vein (4 in Fig. 13A; M1 + 2 of Comstock, 1924; Mt of Tillyard, 1926; R5 + M1 of Lower, 1951; discal vein; Speiser’s Discoidalis). This is always connected with the 3rd longi- tudinal, usually about mid-length in the wing, by means of a short anterior cross-vein ( r-m in Fig. 13A and of Comstock, 1924; r4-m of Lower, 1951 ; first basal, median, middle, outer, or small cross- vein ; Speiser ’s ‘ 1 kleine Querader ’ ’ ; Tillyard ’s radio-median cross- vein; Ferris’ radial-medial cross-vein). Fifth longitudinal vein (5 in Fig. 13A; Ms + Cut of Comstock, 1924; M3 + 4l of Tillyard, 1926, and of Lower, 1951; Speiser’s Posti- calis). In some genera this is connected, by means of the anterior basal cross-vein (M3 in Fig. 13A and of Comstock, 1924; Jf3 + 4 of Lower, 1951; basal or inner cross- vein; Speiser’s “hintere Quer- ader”), with the 4th longitudinal, basad of the anterior cross-vein. Sixth longitudinal vein (6 in Fig. 13A; Cux + 1st A of Com- stock, 1924; anal vein). In some genera this is connected with the 5th longitudinal by means of the posterior basal or anal cross-vein (ac in Fig. 13 A; given no notation by Comstock, Tillyard and Lower; Cu2 of Ferris and Cole, 1922; Speiser’s “ Analquerader ”). 45 ENTOMOLOGICA AMERICANA Fig. 13. A, OrnitTiomyia fringillina Curtis, female; wing: ac, posterior basal cross-vein; ALU , alula; AN, anal cell; IB and 2B, first and second basal cells; CO, costa; h, humeral cross-vein; M3, anterior basal cross-vein; r-m, anterior cross-vein; SC, subcosta; 1, 2, 3, 4, 5, and 6, first, second, third, fourth, fifth and sixth longitudinal veins; Lb, basal costal cell; Is, second costal cell; 11, subcostal cell; III, marginal cell; IV, submarginal cell; V, first posterior cell; VI, second posterior cell; VII, third posterior cell; VIII, axillary cell. B, Lipoptena cervi (Linnaeus) ; stalk of halter, after Brauns (1939). C, OrnitJiomyia biloba Dufour; halter, after Brauns (1939): Jen, knob; sea, scabellum; sta, stalk. D, Ornithoctona erythrocephala (Leach), female; right dorsal half of thorax and base of wing:. ACO, axillary cord; ALU, alula; 46 Volume XXXII Axillary vein ( 2nd A of Comstock), usually absent or vestigial, except at the thickened extreme base. The 4th, 5th and 6th longitudinals are generally weak and dis- colored beyond the cross-veins and never reach the hind margin. The veins are usually bare, except the costa; in some species one or more longitudinals may be partly or mostly setulose on the upper side. The anterior basal cross-vein frequently shows a dis- colored stretch, or bulla (BU), which in Lynchia and Olfersia is so extensive that the vein appears to be obliterated. There may be a similar bulla basally on the 4th longitudinal near the middle of the 2nd basal cell and the vein may be bent upward at this point to run for some distance closer to the base of the 3rd longitudinal. A small bulla is rarely indicated at the upper end of the anal cross- vein. Bullae appear to be absent in most higher Muscoidea; but in Glossina there is at least a trace of one in the anterior basal cross-vein. Costal cell (I in Fig. 13A; C of Comstock; mediastinal), be- tween costa and subcosta, divided by the humeral cross-vein into a basal costal ( lb in Fig. 13A) and a second costal (Is in Fig. 13A). Subcostal cell (II in Fig. 13A; Sc of Comstock), between sub- costa and 1st longitudinal. Marginal cell (III in Fig. 13A; Rx of Comstock), between 1st and 2nd longitudinals. Submarginal cell (IV in Fig. 13A; R3 of Comstock; cubital), between 2nd and 3rd longitudinals. First posterior cell (V in Fig. 13A; R5 of Comstock), between 3rd and 4th longitudinals. The anterior cross-vein ( r-m ) cuts off the basal portion as the first basal cell (IB in Fig. 13A; R of Com- stock; upper or anterior basal cell; Speiser’s “vordere Basal- zelle”). Second posterior cell (VI in Fig. 13A; M2 of Comstock), always combined with the discal cell in the Hippoboscidae. When present, the anterior basal cross-vein (M3) cuts off the basal portion as the second basal cell (1 2B in Fig. 13A ; middle or posterior basal cell ; Speiser’s “hintere Basalzelle”) . AX, fourth axillary; BA, third axillary; BAC, basicosta; CO, costa; DCL, upper calypter; EPI, epipleurite; h, humeral cross-vein; I A, second axillary; INC, axillary incision; NO, first axillary; PCL, lower calypter; PWP, posterior wing process; SC, subcosta; sv, stem-vein; TG, tegula; 1, 2, 3 and 4, -first, second, third and fourth longitudinal veins. E, Pseudolynchia canariensis (Macquart) ; longitudinal section of salivary duct through valve, after Jobling (1926) : NYC, hypodermic cells; SB, salivary duct; SIV, salivary valve; SVM, salivary valve muscles; TA, taenidia. 47 ENTOMOLOGICA AMERICANA Third posterior cell ( VII in Fig. 13A; Cu of Comstock). When present, the anal cross- vein (ac) cuts off the basal portion as the anal cell (AN in Fig. 13A; lAn of Comstock; lower or third basal cell). Axillary cell (Till in Fig. 13A; 2 An of Comstock). The chief venational differences between all Hippoboscidae and most higher Muscoidea are: (1) the lack of a discoidal cross-vein (m of Comstock; se or serial vein of Lower, 1951 ; medial cross-vein of Ferris, 1950; discal, posterior, medio-cubital, postical, or lower marginal cross-vein) between the 4th and 5th longitudinals apicad of the anterior basal cross-vein, so that no discal cell is closed off from the apical portion of the 2nd posterior cell; (2) the straight 4th longitudinal, which is never curved or bent forward at the tip into a subapical or hind marginal cross-vein setting off an “ apical or subapical cell. ’ ’ The axillary lobe or alula (ALU; alar lobe; Speiser’s “Anal- lappen”; proximal alar lobe of Snodgrass, 1935, p. 227) is usually well developed, more rarely reduced or lacking ; when present, it is divided from the axillary cell by a deep axillary incision (INC) of the hind margin. The membrane of the wing is either totally bare or partly or almost wholly covered, either on the upper or the under side or on both, with minute hairs or microtrichia. The extent of these hairy areas often provides useful specific characters, although it tends to vary somewhat within specific limits. Since microtrichia occur sporadically throughout the family, regardless of host associations, it is difficult to surmise what use, if any, they may have. The major differences in the venation, such as the number of the longitudinal and cross-veins, are usually of generic value. Some of the main types are shown in Figs. 12A, 12G-I, 13A, and 14A-F. Minor differences, particularly in the position of the cross- veins and of the apices of the longitudinal veins, are often unreli- able even as specific characters. The Melophaginae show the great- est reduction of the venation, although in the newly-emerged adults, or volants, wings are functional until the permanent host is reached. In Hippolosca, the wings are sturdier than usual; the veins are thick basally and anteriorly, and the membrane beyond them is closely wavy or corrugated, many delicate, more or less parallel or bifurcate wrinkles running toward the hind margin. Both features seem to be adaptations to life among the coarse hair of Ungulates and Carnivora, the chief hosts of Hippolosca , which has some of the most active fliers in the family. 48 Volume XXXII Fig. 14. Wings of Hippoboscidae : A, Hippobosca rufipes v. Olfers; B, Ortholfersia phaneroneura Speiser; C, Austrolfersia ferrisi J. Bequaert; D, Ornitheza, metallica (Schiner) ; E, Olfersia sordida Bigot; T, Lynchia ameri- cana (Leach). 49 ENTOMOLOGICA AMERICANA In many Diptera and particularly in the higher Muscoidea, the base of the wing membrane is connected with the posterior ridge of the parascutellum by means of two scale-like lobes, the calypteres (squamae or squamulae). (Snodgrass, 1935, p. 227, calls the com- bined upper and lower calypteres “ alula”). The one nearest the alula, from which it is separated by a notch, is the distal or upper calypter ( DCL ; squamula alaris or superior) ; the other, inserted on the posterior side ridge of the parascutellum, is the proximal or lower calypter ( PCL ; squamula thoracalis or inferior). At rest, when the wings are directed backward or lie on top of each other over the abdomen, the distal calypter folds over the proximal one, so that the upper surface of the former comes to lie against the upper surface of the latter. Ad. Lutz, Neiva and da Costa Lima (1915, p. 174) first noted that calypteres are not completely lacking in all Hippoboscidae, as is sometimes claimed. In the majority of genera they are reduced in size and particularly very narrow. There is no trace of them in the winged Melophaginae, which also have no alula, as well as in the equally winged Ortholf ersiinae ; they are rudimentary in the winged Ornithoicinae. On the other hand, in certain fully-winged Ornithomyiinae, such as Ornithoc- tona (Fig. 13D), the lower calypter is fairly large, translucent but stiff, bordered by a thick axillary cord ( ACO ) bearing many long cilia; between it and the alula, separated at either end by distinct, broad notches, lies a very narrow membranous upper calypter, also briefly ciliated at the thin edge. In Hippobosca, the relative devel- opment of the two calypteres is reversed, the upper calypter being large and membranous (in fact more squamula-like than the lower calypter of Ornithoctona) , while the lower calypter is very narrow. The difference between the two genera suggests that the reduction of the calypteres in the Hippoboscidae is not a primary condition, possibly retained from some common Muscoid ancestral stock, but rather a secondary feature, which recurred several times independ- ently and not always in the same manner in the course of evolution. Most probably the ancestral stock of all louse-flies was a calyptrate muscoid type, possibly one in which the calypteres were not as fully developed as in most Eecent higher Muscoidea and particularly as in the Glossinidae. Axillary sclerites (alar ossicles or p ter alia) may also be traced in the winged Hippoboscidae (Figs. 12 J and 13D). The tegula or epaulet ( TG )', at the extreme upper base of the wing, is scale-like and usually setulose. This is followed by the short and bare basi- costa (BAC ; subepaulet or parategula), on which the swollen basal 50 Volume XXXII section of the costa is inserted. Immediately behind the tegula lies the first axillary (NO; notopterale), often partly divided by trans- verse sutures, and, between this and the stem vein of the first longi- tudinal, the second axillary (I A; intraalare). Behind these, two closely associated sclerites form the fourth axillary (AX; adanale), at the tip of the posterior wing process (PWP). The third axillary (BA; basanale) is a strong sclerite connecting the fourth axillary with the thickened base of the axillary vein. Halteres (HA) are present in all winged Hippoboscidae, even those with reduced or functionless wings; they are absent in the two completely apterous species of Melophagus. They are rela- tively short for the fly’s size and about the same size in both sexes. Contrary to published statements (Massonat, 1909, p. 79 ; Fraenkel, 1939, p. 75), they are not appreciably reduced in the subapterous species. Brauns (1939) gives the length (in mm.) of those he exam- ined as follows : Ornithomyia avicularia, J 0.479, $ 0.571 ; 0. fringil- lina , 5 0.504, $ 0.445 ; O. biloba , J 0.420, £ 0.437 ; Lipoptena cervi, 5 0.378 to 0.471, £ 0.377 to 0.462; Stenepteryx hirundinis, 5 0.420, <$ 0.395 ; Crataerina pallida , 5 0.496, J1 0.487. They differ little in shape from those of other Diptera and occupy the same position as in the higher Muscoidea, but are never hidden from view by the calypteres. Each halter (Fig. 13C) consists of a moderately widened, conical base or scabellum (sea), followed by a short, slen- der stalk (sta) bearing a relatively broad and short, hammer-shaped knob (kn). According to Roberts (1927) and Brauns (1939), the basal portion of the stalk is provided with a pair of sclerotized in- tegumental rods, one anterior, the other posterior, which are con- nected by a row of transverse braces, giving it a striated appear- ance (Fig. 13B). This structure is peculiar to the ectoparasitic Pupipara and Brauns believes that it prevents further injury to the base of the halter in case the knob breaks off while the fly moves in the plumage or pelt of the host. For flight the fully developed wings spread horizontally; at rest they lie flat over each other on the back of the abdomen, like closed scissor-blades, a characteristic position they share with those of the tsetse-flies. In species where they are reduced and no longer used for flight, they are immovable and cover the halteres, which they may help protect. The integument of the abdomen is mostly soft and extensible, the hardened sclerites being dorsally more or less reduced in size or number and ventrally small or almost lacking. Adult Hippo- boscidae may be said to be physogastric from the time they emerge, 51 ENTOMOLOGICA AMERICANA the soft areas of the integument merely stretching as feeding pro- ceeds or as the larva grows in the uterus. The true first tergite ( TEBl .), nearly always present, forms a pair of laterotergites, bearing the first pair of abdominal spiracles (SP1) near the base, the corresponding median area having seemingly disappeared. These laterotergites are either free or fused in part or wholly with a basal median tergal plate (true 2nd tergite; TEB2) or with the second laterotergites, when either of these are present. The sclero- tized portions of the succeeding tergites are generally reduced to the median plates, varying in number, relative size and shape and thus offering reliable specific and even generic characters. The maximum number of median tergal plates observed in the family is five (in Ornithoica) , the basal one being the true second tergite as shown by its relation to the second pair of spiracles ( SP2 ). At the apex a few small sclerites occur near or around the anal and genital openings, both dorsally and ventrally. A distinct ventral sclerite or first sternite (STB) is often present at the base of the abdomen. In some genera, such as Hippobosca and Myiophthiria, all sclerites, except the basal ones are greatly reduced or lacking, even dorsally, the original segmentation of the abdomen being then indicated only by the position of the spiracles. The spiracles being often difficult to trace, some authors, such as Duf our, were led to believe that the Hippoboscidae lacked all segmentation of the abdomen; in the more generalized Ornithoica the dorsal segmentation is as distinct as in many Muscoidea. It should be kept in mind that in a newly emerged hippoboscid the soft, membranous integument of the ab- domen is greatly contracted or shrivelled, so that the sclerotized parts are crowded together or even overlap, and may appear to cover most of the surface. The soft areas expand, spreading out the several sclerites, after feeding in both sexes, and even more in the female as the larva grows in the genital tract. The appearance of the body is thus completely altered, sometimes deceiving the ob- server into thinking he has to do with a different species. More- over, the hardness and color of the sclerotized areas may also change in the same individual during life. The terminal abdominal segments, or terminalia , are poorly dif- ferentiated and relatively unimportant for taxonomic purposes in this family. In the unfed, non-pregnant female the anal opening ( ANO ), at the hind extremity of the body, occupies the center of the anal tergite (ANT), a broadly elliptical or subcircular sclerite at least partly surrounded by a raised rim. Immediately beyond lies the vulvar opening (YTJO; female goilopore), usually a trans- Volume XXXII verse slit between two small, finely setnlose, sclerotized genital plates (GEP), one anterior, the other posterior. Since, according to Crampton (1942), the female gonopore is located between the eighth and ninth segments, the anterior genital plate may be the modified eighth and the posterior genital plate the modified ninth abdominal sclerite. The male copnlatory organs or external geni- talia were first described and figured by Reaumur (1738, p. 159; PL 11, fig. 4) for Stenepteryx. They are usually placed ventrally and simpler than those of most higher Muscoidea, being reduced to five pieces, possibly representing the gonocoxites, penis valves and penis of other Diptera, although the homologies may be disputed. For my present purpose I adopt Michener’s (1944) terminology, without expressing an opinion as to the validity of his homologies. The gonocoxites ( GCO ; basal segments of the outer claspers; Cole’s genital styles; paralobes of Zumpt and Heinz) are a pair of sclero- tized, bristly, lateral lobes or knobs, either sessile or stalked and sometimes very short, inserted on a small, median, horseshoe-shaped sclerite (ninth sternite), beyond the anal opening. They are peri- phallic (or pseudophallic) organs and merely cover or protect the aedeagus, which when not in use may be retracted almost com- pletely within the notch of the horseshoe. They reach their greatest development in Olfersia and are atrophied in Melophagus. The aedeagus (AED) comprises the true phallic (or euphallic) organs, derived from appendages of the tenth segment. It consists of a basal ring or common stalk, the caulis ( CATJ ) or theca (Hennig’s “Gabelplatte”), supporting three elongate pieces of nearly equal length: a median, cone-shaped or rod-like penis proper (PE; intro- mittent organ or phallus; Cole’s aedeagus; penis sheath of Roberts) and two hard, thickened and pointed penis valves ( PEV ; parameres or gonapophyses of authors; Cole’s interior forcipes; claspers of Roberts), one to each side of the penis. The terminal portion of the penis is hinged to a broad supporting piece (SUP; Hennig’s “Tragplatte”), itself connected basally with the caulis. While morphologically speaking the vulvar opening and the male terminalia are behind the anal opening, they are almost always in a ventral position in the Hippoboscidae, so that in a ventral view they appear to lie basad of the anal sclerite. When the abdomen is not too distended by food or by a devel- oping larva, the terminal median sclerites are often retracted in a deep apical notch, so that the body appears to end in two broad lobes. This is sometimes rather striking, though not truly charac- 53 ENTOMOLOGICA AMERICANA teristic either of the family or of any of its constituents ; nor can it be relied upon to distinguish the sexes. The number and position of the abdominal spiracles (8 Pi to 8P7, Figs. 16A-B) often are given incorrectly for the Hippobos- cidae, although Lyonet (1829; 1832) recognized 7 pairs in Melo- phagus ovinus more than a century ago. Massonat (1909, p. 174), for instance, credits the entire family with 5 pairs only, while other authors have given various numbers for the several genera. Rob- erts (1927, p. 17) gives 5 pairs for Hippobosca, but I was able to locate 7 pairs in this genus. F. R. Cole (1927, pp. 452-453) found 7 pairs in Lynchia americana, Lipoptena mazamae, L. gracilis (= L. traguli ), and Melophagus ovinus. Schuurmans Stekhoven (in litt., 1952) located 7 pairs in Pseudolynchia canariensis. In all the genera where I attempted to trace them, I have found 7 pairs, though sometimes with difficulty. Although I have not examined all the known genera, I feel certain that 7 pairs is the normal num- ber throughout the family. The position of the spiracles and their connection with the tracheal system are shown for Melophagus ovinus by Webb (1945), two of whose » excellent drawings I have copied with the author’s permission (Figs. 16A-B). Possibly some of the abdominal spiracles may be functionless in certain genera, as they are often very small and seem to lie sometimes just beneath the integument, apparently without outside opening. In such cases particularly, they are apt to be removed with the tracheal system when the abdomen is cleared by maceration and washing, preparatory to mounting on a slide. The first two pairs lie some- times free on each side in the ventral integument ; more often one or both of them are included in the sclerotized first and second lat- erotergites, which makes them particularly difficult to find. The succeeding pairs are usually placed in the membranous sides of the dorsum ; the last two pairs are often close to tergal sclerites or one of them may even be included in them (/. i., in Hippobosca) ; but the 6th and 7th pairs never occur together in the 6th tergite, as is sometimes the case in the higher Muscoidea. Malloch (1929, p. 553) pointed out that, in the Glossinidae and Gasterophilidae, the abdominal spiracles 1 to 7 lie in the membrane between the tergites and the sternites; in other calyptrate Diptera at least spiracles 2 to 5 are located in the sclerotized laterotergites. In view of the close relationship of the Glossinidae and Hippoboscidae, it is interesting to note that, in Orhithoctona, spiracles 1 to 7 also lie in the membranous sides of the abdomen, although this is not true for all spiracles in other genera. Moreover, it is doubtful whether the 54 Volume XXXII location of the spiracles either in the membrane or in the sclerites has much significance, since most of the original laterotergites have clearly been lost in the Hippoboscidae, thus placing the spiracles secondarily again in the membrane. The hairiness or pilosity depends on the number of setae and hairs, as well as on their length and texture, three factors which do not necessarily work in the same direction to produce the final ap- pearance. The most hirsute hippoboscid is the apterous Melopha- gus, which is fairly evenly setose all over ; even here the body setae are not actually more numerous than in many higher Muscoidea ( Glossinidae, Calliphora, Musca, etc.) ; they are merely longer and particularly stiffer. In other subapterous and flightless genera, large areas, particularly of the thorax, are completely bare and the hirsuteness results from the overdevelopment and stiffness of the remaining setae. The fully-winged genera differ from these mainly in that the setae are only moderately long. In most cases, however, the setae of the thorax and legs are stiff and bristle-like, while those of the abdomen are softer and shorter. The general stiffness of the setae often obscures the orderly arrangement, or chaetotaxy, of the longer bristles (so-called “ fixed setae” or macro- chaetge), which is such an important feature of the Myiodaria. Some of the customary bristles may nevertheless be recognized in most genera. On the head, the innermost area of the inner orbit bears one or more rows of orbital bristles (orb ; frontal bristles of authors), while near each outer corner of the postvertex are placed at least one, rarely two or more vertical bristles (vb; post-vertical bristles of authors) ; a row of small bristles may be present ven- trally also along the oral margin and there may be a short median row or group of setae on the gula close to the occipital foramen. On the dorsum of the thorax, a few acrostichals (idb) and dorso- centrals ( hdb ) are sometimes present, as a rule posteriorly; or there are instead transverse rows or groups of later o -centrals ( leb ) or prescutellars ( psb ), as for instance in Lipoptena (Fig. 11A). Notopleurals (nob)., postalars (pab) and supradlars (sab) can often be traced; the presuturals (pb), when present, are usually placed far forward. The scutellum bears a preapical row of scutellars (scb). The humeral bristles (hb), on the humeral callosities, may be few (as in Lipoptena) or many, in the latter case showing no definite pattern. The mesopleurals (mpb) either are placed dor- sally on the anepisternum in one or more irregular rows before the base of the wing, or there are one or two only at the outer hind edge. The sides of the thorax are usually almost bare, except sometimes for 55 ENTOMOLOGICA AMERICANA a posterior downward row of mesopleurals ( mpb ) and a few setae seemingly corresponding to the posterior sternopleurals ( stl ?), though not in any definite arrangement. There may be even a group of setae on the sclerite between the hind coxa and the metathoracic spiracle, where the hypopleurals ( hpb ) are located in many Muscoidea; these are never placed in rows and are possibly not homologous with true hypopleurals. The swollen pleurotergites often bear setae, sometimes in a conspicuous vertical row (as in Hippobosca) . The sternum is either nearly bare or more or less setulose and there is often a tendency for the setae to form longitu- dinal or transverse patches or to be crowded toward the sides, par- ticularly near the coxae. The legs are usually also very hirsute, but show only exceptionally outstanding characteristic bristles. In most Hippoboscidae the abdomen is fairly uniformly and densely covered with short or long setae; some species have characteristic bare areas, which in Lynchia are densely covered with a microscopic transverse striation. In addition some parts of the abdomen may bear more prominent bristles or there may be definite numbers of setae in rows or groups on the well-defined sclerites. In most genera only the costa of the wing is setulose. In a few ( Olfersia , Hippo- bosca), some other longitudinal veins, either the third or the fourth, are partly setulose ; but the stem vein, or common base of the 1st, 2nd and 3rd longitudinals, never bears setae. The body hairs are never erect, but either slant more or less backward or are closely appressed against the integument, an arrangement allowing the fly to glide swiftly within the plumage or pelt, while keeping it from dropping out when the host is in motion. The setae are always stiff and occasionally even spine-like ; they are never flattened nor arranged in regular and dense comb-like rows (ctenidia), such as are found in many Streblidae and Nycteribiidae. The nearest ap- proach to such an arrangement is on the apical scutellar margin of Stilbometopa. All bristles or setae of the Hippoboscidae, whatever their size or stiffness, are set in sockets provided with a nerve ending, so that they function as tactile sensoria. True hairs, or purely cu- ticular productions of the outer body wall, are rare, sometimes at the outer margins of the wings or scattered as microtrichia over the wing membrane. Ferris (1930, p. 70) called attention to peculiar patches of dull pollinosity on the head and thorax of Lynchia , Pseudolynchia, and Olfersia , which appear as if dusted with gray powder. With the proper magnification the patches may be re- solved into closely packed, minute hairs, which produce the polli- nose appearance by dulling the reflected light. 56 Volume XXXII In the Hippoboscidae the integument of head and thorax is un- usually tough, strongly sclerotized, leathery and resilient, so that the body can withstand much pressure without harm. These features are particularly pronounced in Hippobosca and even more so in Melophagus, the latter being almost impossible to crush with the fingers. Coloration. The body colors of most louse-flies are dull and form poorly defined markings. Their various shades and patterns are of less diagnostic value than in almost any other group of Dip- tera. Unfortunately, most of the older descriptions are based on color alone, often ignoring completely the true structural charac- ters, so that many species cannot be recognized without a study of the types. Color varies not only individually within the same popu- lation ; it may also change in the same specimen under the influence of extraneous factors. Usually the integument is fairly uniformly pale gray or testaceous in the newly emerged fly; soon afterward some areas of head and thorax, as well as the sclerotized plates of the abdomen, darken to various shades of brown, blackish or mahog- any-red. Some species gradually turn darker all over, either through aging or on certain hosts, possibly due to damper or cooler local climatic conditions, or to differences either in the type of blood available as food or in the color and texture of the hosts’ plumage or pelt. Thus Ornithomyia fringillina tends to be lighter on passerines and decidedly darker on certain other hosts, notably on gallinaceous birds. There is no evidence, however, that these individually acquired color differences are in any way inherited or of racial value. With the very low degree of host specificity of 0. fringillina, some of the offspring of the darker flies of game birds may be expected to reach eventually some song bird, on which they will revert to the paler type. Much could be done in this particular field by controlled experiments on a sufficiently large scale. Cowan (1943, p. 182) noted the color changes of individual Lipoptena depressa in British Columbia : “In the spring and early summer it is possible to distinguish the survivors of last year’s brood from the new brood, even pregnant females, by the more intense pigmentation of the older individuals, — they look brown and weather-beaten be- side the new batch.” It should be emphasized that Cowan found both light and dark keds living side by side on the same individual host. In most species of certain genera, the integument is very dark, either reddish-brown ( Hippobosca ) or black ( Ornitheza , Olfersia, and Lynchia), sometimes with a light metallic greenish or bluish 57 ENTOMOLOGICA AMERICANA sheen. Hippolosca is, in addition, unusual in the family in being ornate with ivory-white or pale-yellowish spots on head and thorax, the resulting regular pattern being to some extent specific, though not always fully reliable. Roberts (1927, p. 12) remarked that in newly-emerged H. equina the whole color scheme is much lighter throughout, the markings being distinct, but that these become less so as the insect gradually assumes a darker hue. A more unusual peculiarity of certain Hippoboscidae is the dull grass-green color they show over the paler areas of the integument during life and for some time after death. The green coloring matter resides not in the integument, but in the circulating haemo- lymph, so that it appears only where the cuticula is not too heavily pigmented. The physiological aspects of this feature will be dis- cussed later in connection with the circulatory system. The green eventually fades away after death in dried flies and disappears even more rapidly from specimens kept in a fluid. It persists longer in the legs and is sometimes seen also in the lighter stretches of the wing veins. It has not been observed in any of the louse-flies of mammals and, among the bird-flies, is known thus far only from the following genera of Ornithomyiinae : Ornithomyia, Ornithoctona, Crataerina, Myiophthiria, and Stenepteryx. The first mention of the dull green color in a hippoboscid was by Reaumur (1738, p. 158) for Stenepteryx hirundinis of European swallows. He states in the “Explanation” of the figures of his “spider-fly of swallow’s nests,” readily recognizable as S. hirun- dinis, that the abdomen and thorax are usually dark green.2 The greenish tinge of this species was apparently not mentioned since in print. Mr. G. E. Woodroffe, in England, who observed both S. hirundinis and Crataerina pallida alive, informs me (in litt., 1952) that unfed adults of these species bred in the laboratory never showed any green color upon emerging or afterward when kept unfed. On the other hand, adults collected from nests were often greenish, particularly on the femora; some specimens were more strongly colored than others and these may have been flies that had fed more recently. Slabber (1768) seems to have been much impressed by the color differences of the three specimens of Crataerina pallida (his “ge- vleugelde zesendertig-tengelige vogel-luis,” as is evident from the figures), which he kept alive for some time. One was pale-green (“licht-groen”) over parts of the abdomen, thorax, head and legs; 2 “ Communement leur corps et leur corcelet sont d’un vert fonce. ” 58 Volume XXXII another showed only a trace of greenish color ; the third lacked it altogether. The green color of C. pallida, the common parasite of European chimney-swifts, was mentioned by several later authors : Massonat (1909, p. 322), Austen (1926, p. 353), Falcoz (1926, p. 36), Kemper (1951, pp. 232-234), and no doubt others. In describ- ing another species of the genus, C. oblusipennis, Austen (1926, p. 356 ) mentioned that a greenish tinge was sometimes distinctly visi- ble here and there in the legs. It may safely be assumed that the greenish color appears in all members of the genus Crataerina. Myiophthiria, with which I unite Brachypteromyia, is a close relative of the two preceding genera and occurs on the same type of hosts. It may be expected to show greenish color also ; owing to the difficulty of obtaining live specimens, the color has been observed in one species only. W. Beebe (1949, Zoologica, New York, 34, pt. 2, 61) noted in Venezuela that the abdomen of M. ( Brachypteromyia ) neotropica was “bright sage green” in life. Green has most often been reported for species of Ornithomyia and is probably universal in this genus. Degeer (1776, p. 285) first noted it in 0. avicularia, stating that body and legs were green, except for the black and shiny thorax. The specific names of two of the synonyms of avicularia refer to the greenish color : Ornitho- myia viridis Latreille (1805) and Ornithomyia viridula Meigen (1830). This feature is mentioned in both descriptions and the authors, unaware of its sporadic and transitory nature, seem to have been unduly influenced by it and by Linnaeus ’ failure to mention it when describing avicularia. Curtis (1836) described his 0. fringil- lina, in England, as “ochreous, inclining to bright green”; Say (1823), in describing his Ornithomyia pallida from North American specimens of fringillina, noted that the tarsi were tinged with green. The specific name of Bergroth’s Ornithomyia chloropus (1901), an- other synonym of fringillina, again refers to the verdigris legs ; it is stated, in addition, that the head was mostly pale green beneath and the abdomen greenish-yellow. Dufour’s (1827) original descrip- tion of O. loiloba likewise stated that the abdomen had a greenish tinge, which disappeared by drying, and that the legs were a livid greenish-gray. Schiner (1864) said of his 0. tenella, which I re- gard as a synonym of loiloba, that the thorax was greenish or yellow- ish and the legs horn-yellowish or pale green. The green color has been so often noted in the foregoing three Old World species that it seems superfluous to cite more references. The South American 0. remota Walker (1849) may also be partly greenish. E. C. Reed (1904) emphasized that in his Ornithomyia chilensis, a synonym of 59 ENTOMOLOGICA AMERICANA remota, “green prevails in life and black after death, because the green occurs in the soft integument between the coriaceous plates and is more apparent before the insect shrivels and dries. Green is more extensive on the under side, as well as on the legs. The extent of this tinge varies greatly among the specimens.” Bigot (1885) mentions the greenish color in the description of his 0. fuscipennis, a species supposedly of Colombia, but as yet unrecognized. In Ornithoctona, the greenish color is a well-known feature of the common American 0. erythrocephala, mentioned by Leach (1817) in his original description for the under side of the head, the sternum and the legs beneath (“sordide viridescente testacei”). However, in his type, at the British Museum, the greenish areas have now turned to a dirty pale-yellow. Newstead (1909, p. 467) noted the vivid green tinge of this fly in life. The descriptions of the following synonyms of erythrocephala mention greenish areas : Ornithomyia laticornis Macquart (1835) and Ornithomyia costaric- ensis Swenk (1916). The second common American species of the genus, 0. fusciventris, shows the same color feature, the legs being originally described by Wiedemann (1830) as mostly green. It was also mentioned in the descriptions of the following synonyms of fusciventris: Ornithomyia parva Macquart (1843), Ornithomyia varipes Walker (1849), Ornithomyia synallaxidis Lynch- Arribal- zaga (1881), and Ornithomyia pirangae Swenk (1916). The Indo- Malayan 0. plicata also is often blotched with green, although this was not mentioned by v. Olfers (1816) in the original descrip- tion. It was noted, however, by later authors who redescribed the same species under new names: Leach (1817) for his Ornithomyia nigricans ; Wiedemann (1824) for his Ornithomyia columbae ; Guerin-Meneville (1831) for his Ornithomyia. australis ; Boisduval (1835) for his Hippobosca sitiens; Macquart (1851) for his Ornitho- myia asiatica; Walker (1858) for his Hippobosca viridipes; Walker (1861) for his Ornithomyia doreica and Ornithomyia batchianica. The African 0. platycera also may be partly greenish, as Macquart (1843) mentioned in the original description. One of the specimens of Ornithoctona ( Ornithopertha ) nitens I have seen from Costa Rica had the femora distinctly greenish several years after it was collec- ted and had been kept dry. INTERNAL ANATOMY AND PHYSIOLOGY I. Internal Appendages of the Integument. Various folds or ingrowths of the body-wall form the rods, ridges and knobs of the endoskeleton, which either strengthen the exoskeleton or act as levers 60 Volume XXXII or apodemes for the muscles. In the Hippoboscidae such ingrowths are unusually developed in the thorax, because its dorso-ventral flattening has resulted in a great extension of the exoskeleton. They should not be mistaken for external features when macerated and cleared specimens are examined in slide mounts. The internal appendages of the head are particularly important, as they consti- tute the hyoid and tentorium, which act as levers for the projector and retractor muscles of the proboscis. II. Muscular System. The head contains ptilinal muscles, which retract the ptilinal sac within the head after the fly emerges ; as well as antennal, proboscidal, suction (or cibarial), and salivary valve muscles. In the winged Hippoboscidae the thoracic muscles are all mesothoracic : six pairs of longitudinal dorsal muscles over the whole length of the dorso-central area (dorsad of the proventri- culus) and three pairs of vertical or slightly oblique sterno-dorsal muscles along the sides of the dorsals. Both sets act indirectly upon wing motion by changing the size and shape of the thorax. There are also many smaller muscles moving the neck (cervicals) and the abdomen (thoraco-abdominals), as well as powerful coxal muscles moving the legs as a whole. In the wingless Melopkagus sterno-dorsals are wanting, while dorsals are either wanting (ac- cording to Cuenot and Mercier) or much reduced and functionless (according to Massonat). Inside the leg segments, sets of muscles move the several pieces, either flexing or extending them, or draw- ing them forward or backward. Most Hippoboscidae are powerful and rapid runners, but rather slow fliers, even when the wings are fully developed. The abdominal muscles are relatively unim- portant in these flies and mostly associated with the respiratory" movements, the digestive tract and the reproductive system, partic- ularly the terminalia. Webb (1945a) describes them in detail for Melopkagus ovinus in connection with the respiratory mechanism. The muscular system was studied in Melopkagus by Dufour (1845), Massonat (1909), Cuenot and Mercier (1922), Jobling (1926), and Webb (1945) ; in Lipoptena by Massonat 1909) and Mercier (1924) ; in Pseudolynckia by Massonat (1909) and Jobling (1926) ; in Ornitkomyia, Crataerina, and Hippokosca by Massonat (1909). III. Nervous System. The nervous system of adult Hip- poboscidae agrees in general with that of the Myiodaria. The central nervous system comprises : ( 1 ) a large cepkalic ganglion, or brain, perforated by the oesophagus and innervating the structures and sense organs of the head; (2) a voluminous tkoraco -abdominal 61 ENTOMOLOGICA AMERICANA ganglion , in the anterior region of the thorax close to the meso- sternum, connected anteriorly with the cephalic ganglion by the cephalo -thoracic cord ; it sends nerves to the inner and outer parts of thorax and abdomen. In addition, a splacknic system , consisting of a proventricular ganglion and connected with the cephalic gang- lion, innervates the viscera. The sense organs comprise: (1) organs of sight , the compound eyes and ocelli, described before. Little is known of visual percep- tion in the Hippoboscidae. In the normal life of the fly it seems to play a minor role, perhaps discriminating only degrees of lumi- nosity. Negative phototropism may to some extent induce the flies to hide in the feathers or hairs, although this behavior is chiefly conditioned, it would seem, by tactile stimuli. Gravid females definitely search for dark places in species leaving the host for larvi- position. On the other hand, when alarmed or forced to leave a dead host, the flies are positively phototropic, tending indoors to fly toward window panes. Perception of moving objects, only a response to differences in light intensity and a rather crude method of orientation, may sometimes guide the flies toward a potential host, for instance after they emerge from puparia dropped at random. There is no evidence that louse-flies distinguish definite objects, either by shape or by color. (2) The organs of smell or olfactory pores are in most insects located mainly on the third antennal segment and palpi. As I have pointed out, the greatly reduced and concealed third antennal seg- ment suggests that the sense of smell is of relatively little impor- tance in the life of the Hippoboscidae. K. M. Smith (1919, p. 64, fig. 43) found that the third antennal segment of Ornithomyia avicularia has no true sense-pits, such as are found in the Muscoidea, but only slight depressions in the surface with a few thin sensory processes. These insects also have no known scent glands and give off no odor detectable by man. (3) In the Muscoidea, the organs of taste are usually located on the labella of the mouth-parts and on the tarsi. It is doubtful that they are much used by louse-flies, which live under conditions where only one type of food, namely blood, can be reached by the mouth-parts. Any preliminary probing of the skin, before actual feeding, is probably regulated mainly by touch and perhaps partly by smell. (4) The organs of touch consist of many tactile setae on the palpi, the second antennal segment, the dorsum of the thorax, the legs, the costa and various other parts of the body. To judge by the 62 Volume XXXII abundance and size of such receptors, it would seem that touch out- weighs all other senses in normal hippoboscid life. According to Schellhase (1921) and Hase (19275, pp. 199-205), when Hippobosca equina is deprived of sight and smell by decapita- tion, it survives for some time, as long as circulation and digestion function. In Hase’s experiments such a fly remained alive away from a host at 22° to 25° C., for 3 days in a dry atmosphere and up to 9 days in a moist chamber, survival comparing favorably with that of a normal insect. It did not attempt to fly and remained practically motionless, being deprived of means of orientation, but held on firmly to the support by the claws. When turned over, it resumed the normal position by throwing the body forward over the anterior margin of the thorax, showing that the ventral surface is positively thigmotropic. It is most interesting that it reacted to a heated glass rod, moved near the wings or hind legs, by lively cleaning motions applied to these parts. The Hippoboscidae have a keen temperature sense, evidenced by their quickly leaving the cooling body of a dead host. Martini (1916, Lehrbuch Medizinischen Entomologie, 3rd Edition, p. 268) and Hoffman, Roth and Lindquist (1950) noted that at a higher am- bient temperature Melophagus ovinus migrates away from the skin of sheep to the outer wool. Herter (1952, p. 144—145 and 151-152) determined experimentally what he calls the preferred temperatures of Melophagus ovinus, Hippobosca equina, Lipoptena cervi, Ornitho- myia avicularia, 0. biloba, and Crataerina pallida , using unfed flies away from the host. They were on the whole close to the normal skin temperatures of the normal hosts, the slight differences observed having scarcely the ecological importance which the author seems to claim for them. Thus the preferred temperature of M. ovinus was 37.89° C., while the surface skin temperature of sheep is 36.5° to 38.5° C. His experiments show that a temperature sense is highly developed in these flies. Prouty and Coatney (1934) found that Pseudolynchia canariensis (= P. maura) has a decided humidity sense, the presence of moisture being very repellent. That the flies, if kept away from a suitable host, survive longer in a moist than in a dry atmosphere, was observed for Hippobosca equina by Hase (19275) and for Melophagus ovinus by Sweet and Seddon (1917). The sensory mechanism of heat and moisture stimulation is as yet little known in insects. A few species seem to have special thermo- receptors and hygroreceptors ; but no such sensoria have been located in any of the louse-flies. Prouty and Coatney (1934) studied various other tropisms of 63 ENTOMOLOGICA AMERICANA Pseudolynchia canariensis. They found that this fly is positively thermotropic and thigmotropic, and negatively phototropic and hydrotropic. According to Cowan (1943), newly-emerged, winged Lipoptena depressa display a strong negative geotropism, always moving upward; also a pronounced positive phototropism, second- ary however to the negative geotropism, but no positive thigmotro- pism. On the other hand, dealated keds, which have fed for some time on a deer, show a strong positive thigmotropism, that causes them to cling to any small object and to each other. (5) Chordotonal and campaniform (or dome-shaped) sense organs are present on some of the wing veins, both on the upper and under side, as well as on the halteres, of all Diptera, even of those that have lost the power of flight. The sensoria on the basal portion of the costa and on the halteres of Lipoptena depressa , which Mc- Indoo (1918) interpreted as “ olfactory pores,” certainly belong in the present group. The innervation of the tactile hairs and chordotonal organs of the wing of Crataerina pallida was carefully studied by Zacwilichowski (1934). He showed that, even in the atrophied and functionless wing of this fly, sensoria are as fully developed and as completely innervated as in some other Diptera with functional wings. Brauns (1939) found in Ornithomyia, Stenepteryx, Crataerina , and Lipoptena numerous dome-shaped sensoria on the base of the halteres, as well as on the swollen and slender portions of the stalk, the latter also bearing so-called Hicks ’ papillae ; the knob itself usually lacks dome-shaped receptors, but bears instead a few short sensorial setae (Fig. 13C). According to Roberts (1927), the knob of the halteres of Hippobosca equina bears two tufts of sensorial setae. The tactile setae and other sensoria of the wings and halteres of the Diptera appear to be proprioceptive organs, stimulated by the movements and internal strains of the body, and to influence also equilibrium during flight. This can hardly be disputed in the case of Diptera with fully-developed and functional wings. As I have pointed out, in 85 per cent of the species of Hippoboscidae the wings are fully developed and used for flight. In these the halteres have no doubt the same function as in other winged Diptera. There is, however, much difference of opinion as to the sensorial mechanism involved. It is particularly difficult to explain why the removal of the halteres impairs the faculty of flight in some Diptera, but not in others. Moreover, total amputation of one or of both halteres sometimes affects the ability to use the legs for walking or for rest- ing in a normal position. The several theories offered in this con- 64 Volume XXXII nection and the experiments carried out to prove or disprove them, have been ably reviewed by Brauns (1939; 1951), Melin (1941), Pringle (1948), and Eggers (1950). Pringle gives preference to the older view and regards the halteres as primarily organs of spe- cial sense, which by their vibrations act as alternating gyroscopes, thus helping maintain equilibrium in flight ; they warn the fly specifically of unsteadiness and deviations from the right course. Evidently this explanation is hardly satisfactory for the subapter- ous, flightless Hippoboscidae ( Crataerina , Stenepteryx, Myiopthi- ria, and Allobosca) , in which the halteres not only show no great reduction, but are well provided with the usual types of sensoria.3 The theory which regards the halteres chiefly as stimulatory organs, contributing to the energy or tonus of muscular movements, might perhaps better fit their case. Melin generalized much the same concept by attributing to the halteres a general controlling nervous function in the physiology of flight, releasing what he calls “myo- dynamic reflexes.” Eggers supposes a nervous coordination be- tween the motions of the wings and of the halteres, which seems hardly more than a variant of the ‘ 4 * * stimulatory ’ 7 theory. It might perhaps be suggested that in the subapterous hippo- boscids of birds, which are long-legged, unusually speedy runners, capable of swift changes of direction, forward, backward and side- ways, the halteres regulate and coordinate the movements of the legs. It is remarkable that, in these flies, enough of the wing is preserved to form a protective covering for the halteres, so that these organs would seem to be essential for the normal life of the insect. On the other hand, the short-legged and completely wing- less Melophagus, which also lacks halteres, is sluggish and merely crawls in the fur of the host. The “subapterous” Allobosca has short, swollen legs and well-developed halteres, and is a slow crawler ; as all known specimens have been taken on the host, I sus- pect that its wing pads may be basal remnants of fully-developed, functional wings present in the newly-emerged fly. • The Hippoboscidae appear to have no functional organs of hearing, nor any devices or organs producing sound either in flight 3 Only the two completely wingless species of Melophagus lack halteres. It should be noted that the species of Lipoptena, Neoli- poptena, and E chesty pus are fully-winged and active fliers upon emerging from the puparia, and cannot be regarded as having reduced wings. I do not, therefore, include them among the subap- terous hippoboscids. Allobosca is provisionally regarded as subap- terous. 65 ENTOMOLOGICA AMERICANA pr at rest. At any rate, the normal flight of the winged forms is noiseless to the human ear. Hase (19276, p. 193), however, noticed that Hippobosca equina buzzes when held by the legs in such a manner that the wings can move freely. As in other insects, much of the so-called instinctive behavior consists of reflexes, tropisms and more purposeful activities in res- ponse to various external stimuli. For normal efficient life it is important that the sensorial receptors, through which the stimuli reach the central nervous system, be kept perfectly clean. Foreign matter should also be removed from legs and wings, if these organs are to function properly. The cleaning motions of the winged Hippoboscidae are similar to those of most other Diptera. Hase studied them in Hippobosca equina (19276, pp. 196-198), which is very active in this respect, no doubt because life in a mammalian pelt exposes it continually to dirt. All the legs, in various combina- tions, are used to clean the legs themselves, the wings and every part of the body within reach. Particular attention is paid to the head, especially the eyes, and the wings. Shortly before feeding ends, the fly always makes some cleaning motions over the eyes and wings, indicating that it is about to leave the site of the bite. Other- wise cleaning is mostly in response to the presence of foreign mat- ter on a given area. Heinz (1949, p. 359) found that the spontane- ous cleaning motions of Stenepteryx hirundinis are of the same type, but very swift and skilful. This fly cleans itself as a rule immediately after being removed from the bird’s plumage; the process is not repeated spontaneously later, but may be induced by powdering the body. Kemper (1951, pp. 253-255) described the cleaning process of Crataerina pallida. Observations by Freund and Stolz (1928), Heinz (1949, p. 358), and Kemper (1951, pp. 255-256) show that Melophagus ovinus is very indifferent to dirt: it does not clean itself spontaneously, except on the mouth-parts; even stimulation by heat or concentrated acetic acid only induces cleaning of the proboscis, exceptionally also of the eyes, with one or both fore legs, the proboscis or head usually bending then toward the moving leg. The nervous system and sense organs were studied by Dufour (1844), Massonat (1909), and Mclndoo (1918) in Melophagus ; by Massonat (1909) in Crataerina, Pseudolynchia, and Lipoptena; by Dufour (1844), Massonat (1909), and K. M. Smith (1919) in Ornithomyia ; by Dufour (1844; 1845; 1851), Ciaccio (1884), Massonat (1909), and Hase (1927) in Hippobosca. The sensoria of the wings and halteres were described by Hicks (1857; 1860), 66 Volume XXXII Weinland (1890), and Mclndoo (1918) for Hippobosca ; by Mc- Indoo (1918) and Branns (1939) for Lipoptena; by Bans (1936 ; his “ Mellophagus cervi” was Melophagus ovinus, according to Branns) for Melophagus ; by Mclndoo (1918) for Lynchia americana (as Olfersia americana) ; by Bans (1936) and Branns (1939) for Ornithomyia and Stenepteryx; by Weinland (1890), Zacwilichow- ski (1934), Bans (1936), and Branns (1939) for Crataerina. IV. Digestive and Excretory Systems. The digestive tract, very similar in all Hippoboscidae, consists of the nsnal main three regions (Fig. 15). 1. The fore-gut (stomodenm) comprises the month, the hyoid, the cibarial pnmp, the pharynx and the oesophagus. In the month- parts, the mouth proper, or buccal cavity, is the hollow tube of the haustellum. The pharynx (P; Fig. 7A-D), or anterior, tubular portion of the oesophagus, runs through the cephalic ganglion in the head; an angular bend, with the apex directed forward, separates it from the cibarial pump ( CIP), which is worked by special dilator muscles, drawing up the blood through the haustellum and pushing it back into the oesophagus. The cibarial pump connects with the hollow tube of the haustellum by means of an intervening tubular part of the buccal cavity, with sclerotized walls, comprising the hyoid (H). The oesophagus ( OE ) continues the pharynx as a slender tube, through the neck to about mid-length of the thorax. Here it opens in the mid-gut through a dilatation containing the cardiac valve ( CV ). Just before the valve, a tube opens on the ventral side of the oesophagus and extends backward into the basal portion of the abdomen, where it widens in a bilobed diverticulum (D), corresponding to the reservoir stomach or crop of other Diptera. The diverticulum should be considered somewhat in detail, be- cause of its possible role in feeding. It is greatly developed in the free-living hematophagous Diptera, such as the stable-flies and tsetse-flies, which use it to store large quantities of blood at one meal. It was overlooked by Dufour (1845) in Melophagus, Hippo- bosca, and Ornithomyia, and also by Boubaud (1909, p. 408, fig. 86) in Melophagus. Its existence in the Hippoboscidae was first recog- nized by Massonat (1909, pp. 159-164), who noted that it was more developed in the flies of birds than in those of mammals. In Hippo- bosca camelina and Lipoptena cervi, the canal is very narrow and the crop small (0.2 mm. long, 0.1 mm. high and 0.17 mm. wide in L. cervi) ; the canal is much wider in Melophagus ovinus and the crop is 0.25 mm. long; blood wras never seen in the crop in these 67 ENTOMOLOGICA AMERICANA genera. On the other hand, in the bird-flies Pseudolynchia canari- ensis (= Lynchia maura ), Ornitkomyia avicularia, and Crataerina pallida , the canal and bilobed pouch are much better developed (particularly in Crataerina) and blood is actually stored in the diverticulum. Massonat states that he saw frequently blood cor- puscles in the crop of Pseudolynchia. Hoare (1923) confirmed these findings for M. ovinus j Theodor (1928, p. 289, fig. 6) for Lipoptena capreoli (= L. caprina), H. camvlina, and Hippobosca equina; Adie (1915) for P. canariensis. These authors did not see blood in the diverticulum ; Theodor noted that in L. capreoli it is sometimes filled with a yellowish liquid. According to unpublished observations by Mr. I. B. Tarshis (in lift., 1951), in living Stilbo- metopa impressa the bilobed diverticulum is rather large, extremely dilated, with smooth, elastic, not plicated outer walls, and is at least partly filled with blood after each meal. Evidently it serves as an additional food reservoir in this species, which, nevertheless, uses also the expansible duodenum for the same purpose. 2. The mid-gut (mesenteron) consists of the proventriculus, ventriculus and duodenum. The proventriculus ( PV ), abruptly wider than the oesophagus, lies in the posterior portion of the thorax. It is essentially a cardiac valve (CV), being nearly filled with the thickened folds of the wall. Its structure is more complex in some genera than in others and its functions are not completely understood. The ventriculus (YE) extends over the hind half of the thorax into the base of the abdomen. The abdominal portion of the mid-gut is the duodenum (DU) and starts at the base of the abdomen as an abruptly widened, sausage-shaped, expansible por- tion, where blood is stored while feeding. This is followed by an unusually long tubular portion (6 to 7 times the length of the body), with post-imaginal growth in the Hippoboscidae ; it describes several loops and opens in the hind-gut, in the distal region of the abdomen, just before the insertion of the Malpighian tubules. At the bend of the right-side loop, Melophagus shows a widened section with an area where the wall is thickened and contains a mycetome (my) of enlarged cells filled with symbiotic micro-organisms. Simi- lar mycetomes have been found in all Hippoboscidae examined for the purpose. Their possible significance will be discussed later in connection with the feeding behavior. Aschner (1931, p. 394) men- tions incidentally that the mid-gut contents of Pseudolynchia canariensis (= Lynchia maura) are enclosed in a peritrophic mem- brane. The presence of such a membrane in the Hippoboscidae was to be expected in view of its common occurrence in the Muscoidea 68 Volume XXXII Fig. 15. Melophagus ovinus (Linnaeus), female; digestive tract and ex- cretory system, modified after Hoare (1923&) and Anigstein (1927) : ANO, anus; C, colon; CV, cardiac valve; D, diverticulum; DU, duodenum; JET, hyoid; MT, Malpighian tubules; my, mycetome of mid-gut; OE, oesophagus; P, pharynx; PY, pyloric valve; E, rectum; SD, salivary duct; SG, salivary gland; VE, ventriculus. 69 ENTOMOLOGICA AMERICANA (including Glossina ), although it has been generally overlooked. The functions of the mid-gut are simultaneously those of digestion and absorption, the digested food products passing by dialysis through the walls of the gut into the body cavity. The process has not been studied in the Hippoboscidae. In Glossina , Lester and Lloyd (1928) and Wigglesworth (1929) describe a division of labor among the several sections of the mid-gut, the apical two-thirds only containing a powerful coagulin. One of the initial changes of diges- tion seems to be the rapid draining of water from the ingested blood, apparently by the Malpighian tubules. However, conditions observed in the tsetse-flies may not be strictly comparable to those of the louse-flies, at least in the species that do not use the divertic- ulum of the fore-gut as a true food reservoir, but store blood instead in the expansible basal portion of the mid-gut. Zacharias (1928, pp. 687-688) observed that, in engorged Melophagus ovinus, blood stored in the basal (or anterior) portion of the mid-gut retains its original color and fluidity. In the next two coils the blood is scarcely modified, only acquiring a more brownish-red tinge. Be- ginning with the right-side coil, where the mycetome is located, the blood is strongly altered, very dark brown to blackish, and clearly in the process of being digested. 3. The hind-gut (proctodeum) is relatively short. It starts with a funnel-shaped pyloric valve (PY), or ileum, where the Malpi- ghian tubules open, followed by a tubular straight colon ( C ) widened posteriorly into a rectum (P), the inner wall of which bears four rectal papillae, or pulsating glands, jutting into the cavity. The rectum ends in the anal opening or anus ( ANO ). The function of the hind-gut is excretory, its wall extracting waste products from the blood of the fly’s body cavity and diffusing them into the proctodeum. Very little material seems to reach it from the mid-gut after digestion. The rectal papillae have also a car- diac or circulatory function. Pour mgin excretory organs, the Mal- pighian tubules (MT), are arranged in two pairs, one on each side, but without true common duct; each tubule is very long and thin, closed at the free end, and twisted throughout the adipose tissue of the body cavity. The contents of the hind-gut are always fluid. The digestive and excretory systems were investigated in Melo- phagus by Ramdohr (1809; 1810; 1811), Lyonet (1829; 1832), Dufour (1844; 1845; 1851), Massonat (1909), Hoare (1923), Engel (1924), Anigstein (1927), and Zacharias (1928) ; in Lipoptena by Massonat (1909), Engel (1924), and Theodor (1928) ; in Hippo- bosca by Dufour (1825; 1844; 1845; 1851), Massonat (1909), Pat- 70 Volume XXXII ton and Gragg (1913), and Roberts (1927) ; in Ornithomyia by Dufour (1844; 1845; 1851), Massonat (1909), and Zacharias (1928) ; in Crataerina by Massonat (1909) ; in Pseudolynchia by Massonat (1909) and Adie (1915). At present they are being stud- ied most carefully in Stilbometopa impressa by Mr. I. Barry Tar- shis, whose results will be published in the near future. V. Adipose System. Dufour (1825, p. 320) first noted that the fat body of Hippobosca equina differs markedly in appearance from that of most other insects. Ilis observation was confirmed and extended to other hippoboscids by v. Wielowiejski (1886, p. 522, for Melophagus) and others. According to Oektay ’s (1951) recent work, in the adult female of Melophagus ovinus, multinucleate cel- lular elements, arranged in loose moniliform or beaded strands, form a mesh throughout the body cavity of the head, thorax and abdomen. The cellular elements, each usually with an even num- ber of nuclei, are of two kinds in the strands, most of them being filled with stored fat particles, but a few being without them. The adipose tissue of the male is mostly similar; the sexual organs are surrounded by a thin, continuous, one-layered sheath of cellular fat-containing elements of the more usual type. Oektay found much the same structures in Melophagus rupicaprinus, Lipoptena cervi, and Hippobosca equina , the differences being mainly in the number of nuclei in each fat-bearing cellular element. Massonat (1909) states that, in Melophagus ovinus and in dealated specimens of Lipoptena cervi, most of the longitudinal dorsal muscles of the thorax are replaced by adipose tissue. In addition to being used for storing superfluous fat, the adipose cellular elements may also have an excretory function and possibly other functions as yet unrecognized. The adipose system was investigated in Hippobosca by Dufour (1825), Massonat (1909), and Oektay (1951) ; in Melophagus by v. Wielowiejski (1886), Berlese (1899), Massonat (1909), and Oektay (1951) ; in Lipoptena by Massonat (1909) and Oektay (1951) ; in Ornithomyia and Crataerina by Massonat (1909). VI. Glandular System. Besides the excretory organs men- tioned in connection with the digestive system (Malpighian tu- bules) and the circulatory system (pericardial cells), various or- gans secreting substances useful to the metabolism or the life of the insect occur throughout the body. They are connected either with the integument or with internal organs. The most important of these are the salivary glands (SG; Fig. 15), built on the same general plan throughout the family. A pair 71 ENTOMOLOGICA AMERICANA of blind, variously shaped expansions in the basal portions of the abdomen, just beneath the diverticulum, one on each side of the ventriculus, are the excretory glands (EG). Each sends off an- teriorly a slender salivary tubule (STU), which, after entering the thorax, widens into an elliptical salivary reservoir (SB; A die’s goblet organ), after which it becomes tubular again. The two sal- ivary reservoirs lie directly above the thoraco-abdominal ganglion, shortly behind and ventral to the cardiac valve. The distal por- tions of the salivary tubules next unite beneath the oesophagus into a common, usually coiled salivary duct ( SD ), which enters the neck. This finally connects at the bulbous base of the haustellum with the very fine tubular channel that runs the whole length of the stylet-like hypopharynx, opening close to its tip in the lumen of the haustellum. The general plan is the same as that of the salivary system of Glossina (Jobling, 1933, pp. 483-487, fig. XI, for G. pal- palis ), except that the tsetse-fly lacks the salivary reservoirs in the thorax. As in the case of the tsetse-fly, there is no true salivary pump in the Hippoboscidae, where the muscles of the walls of either the salivary reservoirs or the excretory glands, or of both, compensate for its absence f the contraction of these muscles squeezes the saliva through the ducts. The structure of the salivary glands was studied most com- pletely in Stilbometopa impressa by Mr. I. B. Tarshis, who has al- lowed me to quote the following details from his unpublished ac- count. The excretory glands are definitely sausage-shaped, about 650 /x long and 80 /x wide. They seem to be attached to the wall of the under side of the diverticulum and are difficult to dissect out without severing their connection with the tubules, which are very narrow, about 8.23 y in diameter. The thoracic salivary reservoirs are elliptical, with an average length of 453 y and a greatest diam- eter of 190 y. The distal portion of the tubules is convoluted and the common duct also pursues a rather winding path. The diam- eter of the common terminal duct is approximately 4 y. The details of the several parts vary somewhat in the other Hip- poboscidae examined so far. The excretory glands of Melophagus ovinus are spherical expansions, according to Dufour (1845) and Massonat (1909, p. 149). Theodor (1928, p. 291, fig. 7) found them also spherical, about 0.25 mm. in diameter, in Lipoptena capreoli (= L. caprina). In the other genera, they appear to be sausage-shaped, as Massonat (1909) and Theodor (1928, p. 292, figs. 9-10) describe them for Hippobosca equina and H. camelma, Theodor adding that they are coiled around the mid-gut. Both 72 Volume XXXII Massonat (1909) and Adie (1915, p. 674) found them sausage- shaped in Pseudolynchia canariensis (= L. maura). Massonat also examined them in Ornithomyia and Crataerina. Of Crataerina pallida (1909, p. 149) he states that the excretory gland (“tube excreteur”) is in the thoracic region, strongly convoluted instead of straight; most probably he saw only the basal portions of the salivary tubules, the true excretory glands in the base of the ab- domen being overlooked because they were severed in dissection. Massonat (1909, p. 148, figs. 35 and 48) first noted that, in Hippobosca camelina, the common salivary duct bends shortly after entering the head, after which it suddenly widens and its wall now acquires inner spiral taenidia. The constriction at the bend has several muscles attached and functions as a salivary valve ( SIV ; Fig. 13E), regulating the outflow of the saliva from the duct into the hypopharynx: upon contracting, when the haustellum is pro- truded for feeding, they stretch the wall of the valve and open the lumen; upon relaxing, when the haustellum is retracted, the wall of the valve folds inward and closes the lumen. Cornwall (1923, pp. 1005-1007, figs. 14-15) describes this valve also for Hippobosca. According to Jobling (1926, p. 336, fig. Ill), the salivary valve of Melophagus ovinus is like that of Hippobosca; in Pseudolynchia canariensis (= L. maura) the basal portion of the duct, behind the valve, is also provided with spiral taenidia. Theodor (1928, p. 291, fig. 8) describes the salivary valve of Lipoptena capreoli (= L. ca- prina) as similar to that of Pseudolynchia. The glands of internal secretion, corpora allata and corpora cardiaca , small cellular bodies in the head, generally at the sides of the fore-gut behind the brain and closely associated with the nervous system, are of special interest. M. F. Day (1943) found that Melophagus ovinus has two corpora allata in both adult and larva, lying one on each side of the dorsal aorta near the anterior part of the thoraco-abdominal ganglion in the adult and near the cerebral ganglia in the larva ; they reach their maximum size during late larval development (third instar) and are considerably re- duced in both pupa and adult. The corpus cardiacum is a single median organ not well developed in the larva, decreasing in the pupa and practically absent in the adult. No differences were found in the corpora allata of male and female, nor in females with and without developing larva. I have described these glands some- what in detail because Day points out that in Melophagus they dif- fer from those found in the higher Muscoidea, where the corpus allatum is a single median unpaired structure in the larva (the 73 ENTOMOLOGICA AMERICANA “ring gland” or “Weismann’s ring,” surrounding the aorta), as well as in the adult. In Lucilia sencata, the corpus allatum is very inconspicuous in the larva, increasing in size in the pupa and the adult. Whether these and other differences are merely functional and correlated with those of the reproductive processes, or whether they may have some deeper, evolutionary significance, is open to question. It will be necessary to study first the incretory glands in other, more generalized Hippoboscidae and in the pupiparous, free-living tsetse-flies. The true function of the corpora allata in insects is as yet obscure ; although there is some evidence that, in Calliphora , Drosophila , and Musca, the larval ring gland is the source of the hormone which induces the formation of the pupa- rium. VII. Respiratory System. The only complete account of the tracheal system of a hippoboscid is that of Melophagus ovinus, recently published by Webb (1945), from which the following is condensed (Figs. 16A-D). The system comprises two main dorsal longitudinal tracheae extending forward from the posterior end of the abdomen into the thorax, where they open into a series of air sacs, and of a subsidiary pair of laterally disposed spiracular tracheae connecting the abdominal spiracles of one side with each other and with the dorsal longitudinal trachea. It is identical for both sexes in the head and thorax, but shows some slight sexual modifications in the abdomen, as was first noted by Massonat (1909, p. 175). There are two pairs of thoracic air sacs. The large anterior pair (AT A) occupy the greater part of the thorax and the pro- thoracic spiracles ( POS ) open directly into their anterolateral angles. They join anteriorly to form a short median trunk entering the head, where it gives rise to four branches further subdivided for the tracheation of the head. Three pairs of tracheae pass di- rectly from the anterior sacs to the respective pairs of legs. The metathoracic spiracles (MES) open directly into the posterolateral angles of the anterior sacs, dorsal to the tracheae of the hind legs. The very small posterior thoracic air sacs ( PTA ) extend upward to the dorsal surface of the thorax and are joined basally by means of a short tracheal trunk to the anterior thoracic air sacs. The two dorsal longitudinal tracheae of the abdomen, upon entering the thorax, open into the posterior thoracic air sacs and pass on to join the anterior thoracic air sacs at the level of the mid legs. In the abdomen of the male (Fig. 16B), the two dorsal longi- tudinal tracheae ( DLT ) arise from the 6th pair of abdominal 74 Volume XXXII spiracles ( SP6 ) and pass forward on either side to join the thoracic air sacs. A dorsal commissure ( TDC ) usually connects both longi- tudinal tracheae. Immediately after leaving their point of origin, each dorsal trachea bifurcates and forms two trunks running side by side. These reunite at the level of the 4th abdominal spiracle ( SP4 ) and continue forward as a single trachea receiving in turn a branch from each of the first five abdominal spiracles ( SPl to SP5). Each of the two subsidiary spiracular tracheae ( SPT ) arises from the 7th abdominal spiracle ( SP7 ), bifurcates almost immedi- ately, runs beneath the double dorsal longitudinal trachea and re- unites just behind the level of the 4th abdominal spiracle. It then diverges into a lateral position, receives branches from the first five abdominal spiracles and finally joins the dorsal longitudinal trachea near the level of the 1st abdominal spiracle. A branch ( TVM ) arises from the 1st abdominal spiracle of each side, run- ning downward and backward to supply the ventral muscles. The passage of air from the thoracic air sacs to the abdominal tracheal system is largely controlled by an intra-tracheal valve (at INV ), inside the anterior region of each of the dorsal longitudinal tra- cheae. The valve consists of a flat sclerotized sheet attached to the floor of the trachea at the point of entry of the branch from the 2nd abdominal spiracle and extending forward as a horizontal lamina almost to the base of the posterior thoracic air sac. This structure allows air to pass freely from the longitudinal trachea to the tho- racic air sacs, but will cut off most of the flow of air in a reverse direction. The abdominal tracheal system of the female (Fig. 16 A) differs from that of the male mainly as follows. The subdivision of the two dorsal longitudinal ( DLT ) and the two spiracidar tracheae (SPT) in the posterior half of the abdomen is greatly increased, and an anastomosing mass of large tracheal trunks (TAN) is formed lying one above the other on either side of the posterior dorso-ventral muscles. The 1st abdominal spiracle (SPl) remains connected with the dorsal longitudinal trachea, but is severed from the spiracular trachea. The branch from each of the 1st abdominal spiracles, which in the male supplies the dorsal muscles (TVM), breaks up into a larger mass of tracheoles now associated with the anterior region. of the uterus. In addition, a similar branch aris- ing from the spiracular trunk midway between the 2nd and 3rd ab- dominal spiracles, and a third, somewhat smaller branch, from the 7th spiracle, supply the lateral and posterior regions of the uterus respectively. ’ 75 ENTOMOLOGICA AMERICANA All Hippoboscidae have two pairs of thoracic and seven pairs of abdominal spiracles. Keilin (1944) has shown that the anterior pair of thoracic spiracles are prothoracic ( POS ), not mesothoracic as had been supposed, since they are placed in the first thoracic seg- ment of the dipterous larvae. The posterior pair are metathoracic ( ME 8 ), being placed in the third thoracic segment. The location of the seven pairs of abdominal spiracles ( SPl to SP7) was dis- cussed in connection with the morphology of the abdomen. In Melophagus ovinus, the prothoracic spiracles are larger than any others and situated mostly dorsally on the antero-lateral sur- face, above and slightly behind the fore, coxae. According to Webb (1945a), the external aperture ( EAP ) is almost circular and opens immediately into a cylindrical atrium (Fig. 16D). The walls of the atrium (AT) are beset with upwardly curving, chitinous spines. At the proximal end of the spiracle, the walls are turned inward to form a stout, sclerotized ledge surrounding the elliptical internal aperture ( IAP ), which may be closed by a diaphragm (DM). The diaphragm is composed of two folds of soft chitin lying on either side of the internal aperture and rigidly fixed to the lower surface of the stout ledge of the inner aperture. An occlusor muscle (MOC) passes from the anterior edge of the diaphragm to an apo- deme (APO) arising from the inner surface of the thorax. The contraction of this muscle stretches the diaphragm longitudinally, approximates the two lips and closes the inner aperture. When the the occlusor muscle relaxes, the diaphragm returns to the open posi- tion by the elasticity of two sclerotized bars (cba), situated in the diaphragm on either side of the aperture. The smaller metathoracic spiracles are placed in the furrow between the thorax and abdomen, facing posteriorly. In structure they differ from the prothoracic spiracles only in minor details (Fig. 16C). The abdominal spiracles differ greatly from those of the thorax, but are alike in all seven pairs. The atrium comprises three suc- cessive, intercommunicating chambers, with smooth inner walls ; the outermost chamber is the largest and opens by a circular aperture ; the second and third are rather short and partially invaginated into the first. The innermost chamber communicates with the trachea by a short, narrow, thickly sclerotized tube, inserted into the side of which is a stout sclerotized rod upturned at its distal end to form two alate processes at right angles to the shaft. From the inner surfaces of these processes an occlusor muscle passes forward and embraces the sclerotized tube on which it becomes inserted. The contraction of this muscle operates on the sclerotized rod constrict- 76 Volume XXXII ing the channel in the sclerotized tube. Below the insertion of the rod, the connecting tube widens slightly before entering the trachea and is here lined with long hairs. From an analysis of the structure of the respiratory system and experiments on the rate of entry of ground derris dust, Webb (1945a; 19455 ) concluded that mechanical ventilation of the tracheae normally takes place in M. ovinus and results from pumping move- ments of the abdomen. Contraction of the abdomen compresses the abdominal tracheae and forces the air they contain over the intra- tracheal valves into the thoracic air sacs, where a slight positive air pressure is then maintained. This pressure is held on the one hand by the closed diaphragm of the thoracic spiracles, and on the other by the intra-tracheal valves which prevent the return of air into the abdominal tracheae. When the muscles relax and the abdomen ex- pands, the abdominal tracheae resume their normal shape and air is drawn in through the abdominal spiracles. When this process has been completed, the diaphragms of the thoracic spiracles open and the positive pressure in the air sacs is released. The dia- phragms close before the next contraction of the abdomen. Prior to Webb’s work, the respiratory apparatus of Melophagus ovinus had been partially and mostly incorrectly described by Lyo- net (1829; 1832), Dufour (1831; 1844; 1845; 1851), Krancher (1881), Massonat (1909), and Hassan (1944). There are also a few data on the respiratory system of Hippohosca by Dufour (1825; 1831; 1844; 1845; 1851) and Massonat (1909), of Ornithomyia by Dufour (1831; 1844; 1845; 1851), of Pseudolynchia by Massonat (1909), and of Crataerina by Krancher (1881) and Massonat (1909). All these accounts are too fragmentary for an intelligent comparison with Melophagus. Massonat was inclined to believe that the respiratory system was more important in the apterous and subapterous than in the fully-winged forms. VIII. Circulatory System. The blood of the Hippoboscidae consists of the usual haemolymph carrying a few suspended cells or haemocytes. A qualitative and quantitative investigation of its organic and inorganic constituents has not been undertaken. The haemolymph is colorless in most species; but in some Ornithomyiinae of birds it contains a pale-green coloring matter, visible through the lighter areas of the integument. The occurrence of this color in certain bird-flies was discussed before. Kemper (1951, pp. 232-234) studied it more carefully in Crataerina pallida. It would seem that it appears in flies only after they have fed : at first the imbibed red bloQd shows distinctly through the integument 77 ENTOMOLOGICA AMERICANA POS ATA SP1 DLT TDG SP6 i I \ r i Pig. 16. Melophagus ovinus (Linnaeus) ; respiratory system; modified after Webb (1945a). A, tracheal system of female in profile from the mesial aspect; B, tracheal system of male from the dorsal aspect; C, longitudinal section of metathoracic spiracle; D, longitudinal section of prothoracic spiracle, APO, apodeme; AT, atrium; ATA, anterior thoracic air sac; cb'a, sclerotized bars of diaphragm; ex, sectioned connections with left thoracic air sacs; DLT, dorsal longitudinal tracheae of abdomen; DM, diaphragm; DAP, external aperture of spiracle; IAP, internal aperture of spiracle; INV, location of intratracheal valve; ME 8, metathoracic spiracle; MOC, occlusor muscle of spiracle; POS, prothoracic spiracle; PTA, posterior thoracic air sac; SP1, 78 Volume XXXII of the anterior part of the abdomen, while the caudal half is at least partly deep bluish-green ; one or two days later, the anterior por- tion of the abdomen also turns deep-green. The changes are some- what more pronounced in the male than in the female. Kemper quotes unpublished work of H. Remmer, showing that the green color of Crataerina is a degradation product of the red haemoglobin of the host’s blood, a so-called “verdoglobin.” Normally the green color passes later in the general body cavity, first in the hind legs, spreads then to the other legs and the soft parts of the head near the proboscis, and sometimes enters the base of the wings and certain areas of the thorax. It was observed twice that a half-grown larva voided prematurely by a greenish female was likewise tinged with green. It is doubtful whether the dull-green color of the Hippo- boscidae is due to the same chemical substance as that found in some other Diptera, such as certain Stratiomyidae and Tabanidae. Ad. Lutz (1912, pp. 79-81) showed that the green color of certain American tabanids {Tab anus limpidapex, T. mexicanus, and others) also resides in the haemolymph, as shown by the blood currents fol- lowing the pulsations of the heart. He states that the green tinge of these flies is a constant feature of the species and not the result of a previous blood meal. That its occurrence in tabanids is independ- ent from the blood diet is further shown by its presence in both sexes, though the males of these flies never bite. A comparison of the chemical properties and origin of the green coloring matter of the blood in the Hippoboscidae and Tabanidae might be of unusual interest. In Hippobosca equina the main blood vessel, or heart, lies dor- sally on the middle line of the abdomen, immediately beneath the exoskeleton. It is a muscular tube comprising five chambers, each opening by a pair of lateral ostia , but without inner dividing septa. The anterior prolongation of the heart into the thorax, or aorta, is much narrower, simple and without lateral ostia. The heart is con- tained in a pericardial cavity (or dorsal sinus) separated from the body cavity below by the pericardial or dorsal diaphragm ; on either side it is joined by the branches of five wing-like muscles and many tracheal ramifications. The remainder of the pericardial cavity is filled with peculiar pericardial cells (or nephrocytes), which appear to have an important excretory function. According to Hase (1927, SP2, SP3, SP4, SP5, SP6, SP7, first, second, third, fourth, fifth, sixth and seventh abdominal spiracles; SPT, longitudinal spiracular tracheae; TDC, dorsal commissure, sectioned at the connection with the left half of the tracheal system in Fig. A; TVM, tracheal branch from 1st abdominal spiracle. 79 ENTOMOLOGICA AMERICANA pp. 205-211), the rate of beat of the heart varies greatly in Hippo- bosca equina (from 60 to 200 pulsations per minute) and the beat is interrupted now and then by pauses. It also changes continually in response to outside and inner factors. As in many other insects, circulation in the appendages is con- trolled or activated by accessory pulsating organs. Hase (1927) found such organs in Hippobosca equina on the legs, one in each tibia, its pulsations being observed at the soft membranous joint between femur and tibia, on the under (or inner) side of the base of the tibia. In the wing an accessory heart occurs at the extreme base of the second basal cell, where the elbow-shaped common origin of the 4th and 5th longitudinals is thickened and discolored. It is interesting that the accessory hearts of legs and wings do not always pulsate in unison with the main abdominal heart and that the rate of beat is not always the same in all legs. Circulation in the wings is probably similar to that observed in some Muscoidea, where the blood enters from the thorax by the costa and adjacent anterior veins and returns to the body by the posterior veins (Thomsen, 1938, p. 419, fig. 1, showing the direction of the blood stream in the wing of Musca domestica) . The only published studies of the circulatory system of the Hip- poboscidae seem to be those by Massonat (1909) and Hase (1927) for Hippobosca. Schuurmans Stekhoven and his co-workers will publish shortly observations on the circulation in Pseudolynckia canariensis. IX. Reproductive System. The peculiar reproductive organs are an outstanding characteristic of the Hippoboscidae. A. Internal Male Organs. In Melopkagus (Fig. 17A) and Hippobosca, the pair of testes ( T ) are simple, slender tubes of considerable length (in some cases 4 to 5 times the length of the body when uncoiled), each closely convoluted into an oval tangle, from which the somewhat club-shaped blind end protrudes. Each appears to be a single sperm tube, in which the male germ cells develop into spermatozoa. It continues posteriorly as a very short, slightly swollen vas deferens (YD), which opens directly (without intervening vesicula seminalis) into the single, median ductus ejaculatorius ( DE ). This is moderately long, slightly swollen at its origin and connected through the gonopore with the endophallic chamber of the aedeagus. The apical portion of the ductus ejacula- torius makes a loop to the left around and behind the rectum. There are two pairs of unusually long vehicular or accessory glands (AG) , each pair opening by a short common duct in the correspond- 80 Volume XXXII ing vas deferens at the point where the latter enters the ductns ejaculatorius. In Ornithomyia, according to Holmgren (1903), there are five accessory glands, the anterior pair being replaced by three glands. The male organs of the Hippoboscidae differ greatly from those of most Muscoidea in the lack of a differentiated vesicula seminalis, in the unusual development and tubular shape of the testes and of the accessory glands, and in the apical loop of the ductus ejaculato- rius around the rectum. They agree in these peculiarities with those of the Glossinidae (Fig. 18 A), which, however, have only one pair of accessory glands. The male genital organs of the Nycterib- iidae and Streblidae have been too little investigated for an intel- ligent comparison; but, in the Nycteribiidae, they appear to com- bine some features of the Muscoidea and of the Hippoboscidae. B. Internal Female Organs (Fig. 17B). The female genital tract lies ventrally in the posterior part of the abdomen, close to the ventral body wall. In a newly-emerged fly, it is short and flattened, occupying only a relatively small part of the shrivelled abdomen. As the development of a larva progresses, the uterus increases greatly in length and thickness and eventually the genital organs fill most of the greatly expanded abdomen. A pair of ovaries (0), one directed to the right and the other to the left, lie dorsally to and a short distance behind the anterior or proximal extremity of the uterus ( U ). The ovaries are ovoid and strikingly dissimilar in size, because they alternate in producing a mature ovum, the out- standing feature in the reproduction of the Hippoboscidae and other Diptera with integral viviparity (so-called pupiparity). Each ovary narrows into a very short lateral oviduct ( LD ), which combines with its fellow to form a rather spacious, but short, com- mon or median oviduct ( MD ), sharply set off from the uterus. In Melophagus and Lipoptena, the junction of the two lateral oviducts with the median oviduct forms a somewhat dilated atrium (AT ; preuterus), which functions as a reservoir for the sperm. Accord- ing to Graham and Taylor (1941), in M. ovinus sufficient sperm is thus stored from a single mating to last during the female ’s lifetime, which is not so surprising in view of the few larvae that are pro- duced meanwhile. The median oviduct joins the genital chamber, or uterus (Z7), somewhat behind the latter’s latero-ventral end. In the newly-emerged fly, the uterus is a broad, depressed tube, about half as long as the abdomen, attached by strong muscles to the dorsal and ventral inner surfaces of the integument ; but when it contains a growing larva, it stretches considerably and eventually occupies 81 ENTOMOLOGICA AMERICANA half or more of the cavity of the abdomen. The uterus narrows posteriorly into a short vagina ( VA ), which opens externally through a crescentic, slit-like vulva ( VU ) at the postero-ventral end of the abdomen (behind the anus), between two genital plates. The foregoing arrangement of the parts is that of Melophagus ovinus, as described by Pratt (1899) and others. The few other genera exam- ined thus far differ only in minor details. According to Harden- berg (1927 ; 1929), Lipoptena is very similar; but the atrium lies to the right of the uterus (not dorsally on it, as in Melophagus) and the left lateral oviduct is much longer than that on the right side. He also states that in Hippobosca variegata (=H. maculata), H. equina , H. rufipes, Crataerina pallida , and Stenepteryx hirundinis, there is no atrium, the two lateral oviducts opening directly in the median oviduct, while the latter is not sharply set off from the prox- imal end of the uterus. The dorsal anterior (or proximal) end of the uterus receives two sets of glands. In Melophagus ovinus , where there are two pairs, the glands of each pair first combine in a common duct and these two ducts then unite in a single very short duct opening in the uterus. The hind pair are two large, extensively branched, tubular structures, evidently modified accessory (or fecundatory) glands. They have acquired a nursing function seemingly in all Hippo- boscidae, secreting a milk-like fluid on which the larva feeds, so that they can properly be called milk-glands {MG) . The forward glands ( FG ) vary in number, size, structure and function in the family. Their homology has been disputed ; but there is now little doubt that they are modified spermathecae (receptacula seminis), since they occupy the same position and open at the same point of the uterus as the functional spermathecae of the higher Muscoidea (such as Glossina, Fig. 18B) . In Crataerina pallida and Stenepteryx hirun- dinis, according to Hardenberg (1927 ; 1929), there are three such glands (a frequent number in the Muscoidea, though Glossina has two only), which are short, unbranched and actually function for the storage of the spermatozoa after mating. The same author de- scribes the forward glands of Hippobosca as long and branched (also observed by Dufour, 1845, p. 92; pi. 3, fig. 35, H. equina) ; he found three of them in H. rufipes, but only a pair in H. variegata {= H. maculata) and H. equina ; he also states that they no longer function as spermathecae, the spermatozoa being stored instead in the oviducts, since there is no atrium. Hardenberg suggests no other function for the branched forward glands of Hippobosca ; I surmise that they serve to some extent as milk-glands. In Melopha- 82 Volume XXXII Fig. 17. Melophagus ovinus (Linnaeus). A, male organs: AG, accessory- glands; DE, ductus ejaculatorius ; T, testes; VD, vas deferens. B, female organs from above: AT, atrium; EG, forward pair of glands; LD, lateral oviduct; MD, median oviduct; MG, milk glands (drawn on one side only); 0, ovary; V, uterus; V A, vagina; VV, vulva. C, longitudinal section of female abdomen and of larva developing in uterus: ANO, anus of adult; AL, anus of larva; EG, forward gland; HG, hind gut of larva; MD, median oviduct; MGD, duct of milk-gland; MT, Malpighian tubules of larva; 0, ovary; OE, oesophagus of larva; PE, proventriculus of larva; TJ, uterus; VE, ventriculus of larva; TV, vulva. All modified after Pratt, Roubaud and others. 83 __AG — SPE 01 Fig. 18. A-C, Glossina palpalis (Robineau-Desvoidy). A, male genital organs, modified 'hfter Minchin (1905, Proc. Boy. Soc. London, B, 76, p. 542, fig. 5). B, female genital organs, modified after Roubaud (1909, p. 430, fig. 91) and Newstead (1924, p. 5, fig. 1). C, inner structure of ovaries, modi- fied after Newstead (1909) and Hagan (1951). D, Melophagus ovinus (Lin- naeus) ; inner structure of ovaries, modified after Pratt (1899). AG, accessory glands; ANO, anus; AT, atrium; BE, ductus ejaculatorius ; DMG, common duct of milk glands; FOl, F02, F03, F04, F05, F06, F07, F08, follicles in the order in which they produce ova ; GF, germarium ; LB, lateral oviducts ; MB, median oviduct; MG, milk glands; NTJ, nurse cells; 01, 02, right and left ovaries; OOC, oocyte; OV11, 0VI2, OVI3, OVI4, ovarioles in the order in which they produce ova; PES, peritoneal sheath; B, rectum; sp, sperm; SPE, spermathecae ; T, testes; TTJ , tunica propria of each ovariole; U, uterus; V A, vagina; VU, vulva. 84 Volume XXXII gus and Lipoptena, however, they are reduced to a pair of short, thick, simple tubes, which do not store spermatozoa and may be rudi- mentary and functionless. Zacharias (1928, p. 685) observed that in M. ovinus the forward glands occasionally contain some sper- matozoa, which he believes are resorbed there. Each of the two ovaries is encased in a peritoneal sheath of un- usual thickness, forming an elastic sac. In Melophagus ovinus (Fig. 18D), the only hippoboscid in which the inner structure has been completely described (Pratt, 1899), each ovary contains two ovarioles ( OVI ), each enclosed in its own tunica propria ( TTJ ), and each with a germarium ( GE ) at the bottom end, followed by two successive follicles ( FO ). In each ovary also, the two ovarioles function alternately, so that the ovary contains only one developing ovum (OF) at any time. Hardenberg states that the ovaries of Lipoptena, Hippobosca, Grataerina, and Stenepteryx agree with those of Melophagus ; unfortunately he describes the inner structure of none of these genera. A comparison with the female genital tract of Glossina (Fig. 18B) discloses that its structure is fundamentally the same in the Hippoboscidae and the Glossinidae, and is, moreover, essentially that of the other higher Muscoidea. Glossina differs in the follow- ing particulars, according to Roubaud (1909, p. 430, fig. 91), New- stead (1924, p. 5, fig. 1), and Hagan (1951, pp. 112-118), the differ- ences being all seemingly of minor importance. (1) Each ovary comprises only a single ovariole, instead of two. (2) The single ova- riole contains four or five follicles at a time, instead of two, so that the total number of follicles in each ovary is about the same. The two foregoing differences are based on the assumption that Pratt’s interpretation of the inner structure of the ovary of Melophagus is correct and that the other genera of Hippoboscidae agree with this. Personally I feel that the matter calls for further investigation. Whether the ovaries of Glossina open directly in the common oviduct or by means of intervening lateral oviducts, seems to be a matter of definition. In Melophagus and Lipoptena, an atrium intervenes to differentiate more clearly the lateral oviducts from the median ovi- duct ; but in other Hippoboscidae, which lack the atrium, I can see little difference from the arrangement of Glossina. (3) The two ducts of the accessory or milk-glands unite into a long common duct before opening in the uterus. (4) The two fully functional sperma- thecae are globular, firmly united by a transparent tunica propria, but each with its own long, slender tube connecting it with the uterus; in the Hippoboscidae they are clearly degenerated, func- 85 ENTOMOLOGICA AMERICANA tionless in some genera, or possibly with a secondarily acquired nutritive function in others. From what little is known of the female genital tract of the Nycteribiidae and Streblidae, these insects differ mainly in the lack of a differentiated oviduct, the two ovaries opening directly in the uterus, in the presence of a pair of short, simple, functional spermathecae, and in the more elongated common duct of the two ramified milk-glands. It may also be noted that while the maturing ovum of Glossina contains 15 nurse cells in addition to the oocyte (as in most Muscoidea), only 7 nurse cells are present in the Hippoboscidae (at any rate in Melopliagus ovinus, the only species examined in this respect). According to Harden- berg (1927 ; 1929), the Nycteribiidae have 7 nurse cells (in Cyclopo- dia) and the Streblidae 15 (in Nycteribosca) .4 The reproductive system of Melophagus was studied repeatedly during the past century, beginning with v. Siebold (1837) and Dtifour (1844; 1845; 1851) ; Leuckart (1854; 1855; 1858) was the first to interpret it correctly and he was followed by Pratt (1899), Berlese (1899), Holmgren (1903), Roubaud (1909), Patton and Cragg (1913), Hardenberg (1927; 1929), Zacharias (1928), and Zavattari (1928). It was described by Reaumur (1742), Dufour (1825; 1844; 1845; 1851), Massonat (1909), Patton and Cragg (1913), Hardenberg (1927 ; 1929), and Zavattari (1928) for Hippo- bosca; by Hardenberg (1927 ; 1929) for Lipoptena ; by Dufour (1844; 1845; 1851) and Holmgren (1903) for Ornithomyia; by Hardenberg (1927; 1929) for Crataerina; by Hardenberg (1927; 1929) and Huzimatu (1938) for Stenepteryx. Hagan (1951, pp. 159-169) discussed the female genital tract of the Hippoboscidae from the literature and compared it with that of the Glossinidae. DEVELOPMENT AND EARLY STAGES Together with other so-called “pupiparous Diptera” (Nycteribi- idae, Streblidae and Glossinidae), the Hippoboscidae are endowed with integral viviparity of a peculiar, adenotrophic or incubating type (H. R. Hagan, 1948, p. 64 ; 1951, p. 59 ; formerly called by him, 1931, p. 22, intussuctio-viviparity), not known at present among other insects. In these flies the egg contains sufficient yolk to nour- 4 In the text of his paper Hardenberg (1927, p. 36 ; 1929, p. 532) states definitely that Cyclopodia ferrarii (cited by error as C. plen- octena in 1927) has 7 nurse cells, like the Hippoboscidae; but in the summary (1927, p. 42; 1929, p. 538) he attributes 15 nurse cells ta both the Nycteribiidae and the Streblidae. 86 Volume XXXII ish the embryo until the first instar larva hatches, after which spe- cialized maternal organs feed the offspring in the uterus throughout larval life. In the Hippoboscidae (Fig. 18D), each of the two follicles of each of the two ovarioles contains a developing ovum; so that at any one time the ovary shows at most four developing ova, or a total of eight for both ovaries of a fly. Normally the two ovaries alternate in producing a mature ovum and so do the two ovarioles in each ovary. One of the ova develops therefore far ahead of the other seven, so that the two ovaries are strongly asymmetrical in size. When fully mature, the ovum moves by way of the oviduct to the uterus, where it eventually hatches into a first instar maggot. An- other ovum now matures in one of the ovarioles of the ovary on the opposite side ; it does not move to the atrium until the preceding larva is fully developed and has been voided by the fly. After an ovum has left an ovariole, the germarium produces a new follicle, thus restoring the original number of eight ova. The ovaries thus engender a series of mature ova, one after another, until the pro- ductivity of the germarium is exhausted or the fly dies. As a con- sequence, the total number of ova produced by a female is very small as compared with the numerous offspring of most other insects. According to Dr. Hagan (in lift., 1951) : “ Oocyte formation prob- ably follows that of related Diptera; therefore, in the germarium ( GE ), a cell from the germal mass, at intervals, undergoes 3 divi- sions, thus producing 8 daughter cells. Of these, one will be oocyte (00(7), the remaining 7 cells will be nurse cells (NU). This proc- ess does not subtract 8 cells from the reserve in the germarium with each ovulation, but only one. The original supply then would seem more than ample to last the entire life of the female.” Lassmann (1936, p. 398) found on two occasions in Melophagus ovinus that the uterus contained two eggs, both with complete egg envelopes and micropyles. As he never found twin larvae in the uterus he as- sumed that one of the larvae is aborted as soon as it hatches or suc- cumbs due to lack of nourishment.5 Egg. Upon leaving the ovary, the mature egg is typically muscoid, about 1.16 mm. long and 0.3 mm. wide, elongate cylindri- cal, tapering at the poles and blunter posteriorly, very slightly con- cave dorsally and somewhat convex ventrally. It is enclosed in a 5 Abnormal voiding of twin larvae has been observed on rare occasions in the tsetse-fly, Glossina palpalis. It is therefore not im- possible that it might sometimes occur in the Hippoboscidae. 87 ENTOMOLOGICA AMERICANA membranous two-layered chorion, at the cephalic end with a funnel- shaped depression, in which the pores of the micropyles open. The depression is soon filled with spermatozoa, impregnation occurring either in the atrium or in the oviduct. The embryonic develop- ment was rather fully investigated in Melophagus ovinus by Leuck- art (1858), Pratt (1893; 1897; 1900), Hardenberg (1927; 1929), and Lassmann (1936) ; their observations have recently been criti- cally reviewed by Hagan (1951, pp. 169-186), to whom the reader is referred for the details. Hardenberg (1927; 1929) also published fragmentary observations on the gastrulation of Lipoptena cervi, Stenepteryx hirundinis , and Crataerina pallida. The maturation processes of the egg were partly described for M. ovinus by Lassmann, but actual fertilization was not observed. Polyspermy seems to be the rule in Melophagus, each egg containing six sperms in the early maturation stages. According to Cooper (1941), in Melophagus ovinus both male and female gonial plates show a diploid set of 18 chromosomes, the largest number yet reported in the Diptera Cyclorrhapha. Lass- mann’s earlier observation of only 8 haploid somatic chromosomes in the oocyte of the sheep-ked was erroneous. On the other hand, Cooper (1942; 1944a) found that the diploid set of male Olfersia lisulcata, as determined from spermatogonia and spermatocytes, has only 8 chromosomes. I am under obligation to Dr. K. W. Cooper for calling my attention to several other features of chromo- somal behavior in these flies. In both acalyptrate and calyptrate Muscoidea, no cases are known of a regular occurrence of chiasmata connecting the pairing chromosomes at meiosis in the male and in this respect the meiosis of Olfersia and Melophagus is like that of other Muscoidea. In first spermatocytes the sex chromosomes of the Muscoidea have only a small point of association in pairing and this is also true of Olfersia ; but in Melophagus the sex chromosomes do not pair at all and their mode of segregation is of a peculiar type. Furthermore, in both Olfersia and Melophagus, at meiosis in the male the autosomes, like the sex chromosomes of most muscoid flies, pair only by small segments ; in this regional localization of meiotic conjunction in the autosomes of males, they differ from the Muscoi- dea (as well as from the Streblidae and Nycteribiidae investigated thus far), in which the autosomes pair along sizeable lengths so far as known. Larval Instars. The following account refers to Melophagus ovinus, the larva of which is best known. The first larval instar, hatching from the egg, is more or less maggot-like, with superficial 88 Volume XXXII traces of segmentation (12 segments according to Leuckart), 8 pairs of rudimentary lateral spiracles, a posterior anal opening, and ante- riorly a small, nipple-like projection bearing the month between two minute lateral papillae. There is no trace at this stage of a cephalo- skeleton nor of mouth hooks. A posterior stigmatic plate, at the distal end, bears a single pair of stigmata. After the first moult, the maggot-like shape disappears, the second larval instar being a plump ellipsoid. In the course of intra-uterine development there are two moults in all, between the three larval instars. The first moult occurs soon after the egg hatches and the second when the second instar is about 2.7 mm. long. The third larval instar is barrel-shaped, slightly flattened at both ends and ventrally, 3.5 to 3.7 mm. long, about 1.9 mm. wide and about 1.6 mm. thick, when fully grown. The mouth is directed cephalad in the female ’s body, facing the median oviduct. The posterior end, with the anus and respiratory openings, faces the vulva. The larva (Fig. 17C) lies free, without connection with the mother’s tissues and separated from the inner wall of the uterus by the exuviae of the earlier instars. The only external traces of segmentation are two curved rows, one on each side both dorsally and ventrally, of 7 shallow depressions (mistaken for spiracles by Bonnet, 1776) mark- ing the insertions of dorso-ventral respiratory muscles which run on either side of the digestive tract. These are the only cutaneous muscles developed in the larva. Anteriorly the integument shows the first indications of the circular and semicircular seams or weakened lines, typical of all Cyclorrhapha, along which the pupa- rium will break open for the escape of the adult. According to Pratt, the first and second larval instars of Melopltagus show no traces of these seams, which appear only after the second larval moult. A tubular heart extends nearly the whole length of the body, dorsad of the mid-gut. Adipose tissue is unusually abundant in the general cavity (Oektay, 1951) and the first indications of the sexual organs may be traced. The nervous system is similar to that of the muscoid maggots, but the several ganglia are less concentrated in a central mass. There appear to be no sense organs. The respiratory system of the third larval instar is metapneustic, on the whole normally developed, although showing some unusual features correlated with intra-uterine life. There are two main longitudinal, dorsal tracheal trunks, with several transverse con- nectives and many side branches ramifying throughout the body. Of the eight pairs of rudimentary lateral spiracles of the first stage 89 ENTOMOLOGICA AMERICANA larva, the six hind pairs are completely lost ; the two anterior pairs, prothoracic and metathoracic, persist as microscopic, fnnctionless apertures of the cuticle, just before the circular seam and above the horizontal seam. The main tracheal trunks do not open ante- riorly and posteriorly, as in most Muscoidea, but only in a pair of highly modified distal stigmal plates, the “polypneustic lobes” of Newstead (1918). In Melophagus each of these lobes is furnished with a deep, cup-shaped pit, near the bottom of which and occupying a sub-central position is the stigmal opening which communicates directly with one of the main tracheal trunks ; near the rim or pe- riphery of the pit are two large stigmata : one toward the venter, the other toward the dorsum ; in addition there is also a very minute pore-like stigma; and outside the pit an outer-lateral stigma ren- dered most conspicuous by its large and strongly sclerotized peri- treme. All the stigmata, with the possible exception of the very minute one, are connected by a thick-walled air sac or trunk, which latter can be easily traced through the integument. The stigmal plates of the larva of Melophagus were also studied by Keilin (1944, p. 55, fig. 48a). Buxton (1924) and Ferris and Cole (1922) state that, in the larva of Lipoptena, each polypneustic lobe bears three curved lines of spiracular pores, those of each line opening in a tracheal branch, the three tracheal branches uniting in a spherical chamber from which starts the rather thick-walled main tracheal trunk. A variation of this arrangement was described and figured by Ferris (1923) for Ornithoctona fusciventris (= 0. strigilecula) . As Buxton suggested for Lipoptena , the three groups of spiracular pits are evidently derived from the three slits in the anal stigmal plates of the third stage maggot of many Muscoidea. The more aberrant arrangements of Melophagus and other genera are further specializations. Exchange of respiratory gases is exclusively through the genital opening and vulva of the mother. The digestive tract is voluminous. The functional mouth, open- ing through a conical papilla, leads to an irregular cavity, followed by the invaginated true cephalic segment ( cephalic pouch), which is moved by a set of muscles in rhythmic contractions, so that it acts as a sucking pump to take up the liquid food. On each side of the buccal papilla, dorsally to the horizontal arch seam, there is a smMl papilla. The cephalic pouch contains the pharynx and its walls are sclerotized into a “ pharyngeal skeleton’’ or “ bucco- pharyngeal armature,” as in other Cyclorrhapha. However, ac- cording to Keilin (1916, p. 401), the armature is reduced to the innermost, basal or pharyngeal sclerite, the ventral wall of which 90 Volume XXXII is smooth, the distal pieces and particularly the mouth hooks being absent.6 There are also no salivary glands. The cephalic pouch continues in the tubular, horizontal oesophagus ( OE ), which bends upward abruptly into a tubular, vertical proventriculus (PR), it- self opening in the very large, balloon-shaped ventriculus (YE), closed behind, which fills most of the body cavity. The hind-gut (H G) is a short narrow tube, blind anteriorly where it receives four Malpighian tubules (MT ; 2 ventral and 2 dorsal), and ending in a ventrally placed anus (AL). The peculiar structure of the digestive tract of the hippoboscid larva is almost duplicated by that of the larval tsetse-flies. In both cases it is clearly conditioned by the intra-uterine feeding with the highly nutritious and directly assimilable secretion of the mother’s milk-glands, which has induced profound changes in the physiology of digestion and absorption. According to Roubaud (1919), in the tsetse-flies, the food received in the ventriculus of the larva under- goes no further digestive changes, but only a selective absorption. The epithelial cells remove mainly the adipose parts of the food to the larval body cavity. Nearly all the proteins remain unchanged inside the ventriculus and become available to the general metabo- lism only during the pupal stage. The continued accumulation of proteins in the ventriculus throughout larval life accounts for the enormous size of this portion of the digestive tract. There is also very little formation of nitrogenous waste products, thus making their steady elimination from the body unnecessary, which explains why the ventriculus is closed off from the hind-gut. The peculiar digestive processes of the larvae of the pupiparous Diptera no doubt have also far-reaching effects upon other phenomena in the life of these flies, such as the histological changes during pupation, the duration of pupal development, the resistance of the puparia to ad- verse conditions, and consequently the rate of reproduction, as may be seen by referring to Roubaud ’s work. In Melophagus ovinus, the fat entering the body cavity of the larva is stored in the adipose tis- sue, which, according to Oektay (1951), differs from that of the adult fly ; it forms irregular lobes of one to three layers of cells and there are also isolated fat cells; it is therefore more of the usual type found in most insects. 6 Keilin states that he examined the larvae of Melophagus ovinus and Hippobosca equina, but unfortunately does not describe nor figure their buccal armature. He gives a figure only of the pharyn- geal armature of Glossina and infers that in Melophagus and Hippo- bosca it is similar. 91 ENTOMOLOGICA AMERICANA The nature of the larval food is of great interest. From the position of the larva in the uterus and the rhythmic movements of its cephalic pump, it should take up any material, either liquid or in suspension, that fills the common median oviduct. In Melophagus, Leuckart, Pratt and most other authors believe that this material is essentially, if not solely, the secretion from the milk-glands of the female. Berlese (1899), on the other hand, claims that the larva feeds on the abundant sperm and secretion from the accessory glands of the male, both being injected in the uterus at the frequent matings. Berlese also states that the secretion from the so-called milk-glands of the female has no nutritive function, but serves only to cover the newly laid larva with the sticky substance that glues the puparium to the fleece. The two views may perhaps be recon- ciled, as the larval food may possibly include, at least in the early larval stages, some sperm and secretion from the male accessory glands, which might explain the unusual development of the male internal organs. Zacharias (1928), however, found a few sperma- tozoa in the lumen of the forward pair of glands opening anteriorly in the uterus of female Melophagus, so that some of the superfluous sperm may perhaps be resorbed in these organs. I agree with Bou- baud (1909), moreover, that the main larval food of the Hippobosci- dae is the secretion from the female milk-glands. Most probably the viscose substance surrounding the freshly voided larva of Me- lophagus is also secreted by the milk-glands, as Berlese believed, since no other glands are known that might produce it. In other Hippoboscidae, however, which do not glue the puparium, but deposit clean and fairly dry larvae, the milk-glands are fully as well developed as in Melophagus. Huzimatu (1938) has given a rather complete description of the three larval instars of Stenepteryx hirundinis (= Lynchia nipponica Kishida). He dwells particularly upon the tracheal system of the third instar. Imaginal Disks. Pratt (1900, correcting his earlier papers of 1893 and 1897) found that, in Melophagus ovinus, the several imaginal disks, or rudiments from which the adult organs will be built up during pupation, appear at different times in the ontogeny. The cephalic and thoracic disks can be traced in the embryo, before the egg hatches. Three cephalic disks, one ventral and two dorsal, appear first, when the intestinal, tracheal and nervous systems are as yet rudimentary. The thoracic disks appear later, about the time the primitive embryonic stomodeum is involuted within a projecting papilla or sucking mouth and after the head disks are 92 Volume XXXII fully formed. In all, three ventral and three dorsal pairs of disks, pro-, meso-, and metathoracic, can be traced in the embryo. It is interesting that, although adult Melophagus has only the merest traces of wings and lacks halteres completely, the imaginal disks of both these organs are fairly well developed and persist in the larva, as Stange (1907) also found. The abdominal disks are not developed in the embryo, but appear later, probably in the early larval instars. They were only briefly described by Pratt in his earlier paper (1897, pp. 194-197) ; they comprise first two pairs of anal disks, placed in a transverse row in front of the anus, which Pratt thought develop later into the sexual external appendages; seven ventral and two dorsal pairs of groups of cells, regularly distributed in the epidermis, constitute the epidermal disks; there is also a ring of imaginal epithelial cells around the anus. Puparium. All three successive larval instars develop in the uterus of the female fly. The third instar is voided by the mother only when it is full-grown. It is practically motionless, as it lacks true muscles of progression. Once outside the mother’s body it takes no food of any kind. At first it is usually dirty-yellowish or white, except for the much darker, brown or blackish posterior polypneustic lobes ; the larval integument gradually turns chestnut- brown to black, while the cuticle hardens and becomes rather brittle. It thus transforms rapidly into a puparium, within which is formed the true pupa, as in all Cyclorrhapha. The soft third larval instar, in the process of being deposited, may therefore be called a prepuparium. In the puparium, the papillae and the closed buccal slit are barely visible at the anterior end; while posteriorly the two polypneustic lobes protrude somewhat more than in the larva. Dorsally and ventrally the two rows of 7 de- pressions of the larva are retained, each row placed in a slight longitudinal groove. The seams of the anterior cap, through which the adult will escape, are more or less apparent, in Melophagus difficult to see. The circular seam runs across and around the body (in the higher Muscoidea it is placed in the apparent maggot seg- ment 5). The semicircular seam passes over the top to the sides, forming a bow slightly dorsad of the buccal papilla and ending in the circular seam. When about to emerge, the adult pushes blood, by contracting thorax and abdomen, into the soft-walled, reversible ptilinal pouch. The increasing pressure of the pouch on the inner wall eventually ruptures the cap at the semicircular seam into two valves, which break off at the circular seam. The opening of the 93 ENTOMOLOGICA AMERICANA puparium by means of the ptilinum was first described by Dufour (1845, pp. 85-86) for Melophagus ovinus. Because as a rule a full-grown third instar larva and not a true puparium is deposited by the female, it has been contended that the term “pupiparous” as applied to the Hippoboscidae is a misnomer. The larva when voided is, however, at least a potential puparium, since it merely needs to harden the integument. In Stilbometopa impressa, according to unpublished observations by Mr. I. B. Tarshis {in lift ., 1952), larviposition proceeds by stages: at first the larva is only partly protruded, being carried about for some time by the female, much like the ootheca of certain cock- roaches ; meanwhile the larval integument hardens and turns amber- colored and shiny, before the prepuparium is actually deposited. Moreover, as shown later in the discussion of pupation, in some species the larva occasionally remains in the uterus until after some of the internal pupal transformations have been initiated. In any case, the integral incubating viviparity of Hippoboscidae differs fundamentally from the partial viviparity of certain other so-called “larviparous” Diptera, which deposit larvae in an early stage of development, that feed, grow further and sometimes even moult outside the mother’s body.7 For these reasons it would certainly be more of a misnomer and completely misleading to call the Hippo- boscidae merely “ larviparous. ” Those who wish to be unduly meticulous might perhaps use the term ‘ ‘ larvi-pupiparous. ’ ’ In all cyclorrhaphous Diptera that have been studied for the purpose, the epidermal cells become detached from the cuticle over the whole or part of the inner surface of the third larval in- star, during the process of hardening into a puparium. This de- tached epidermis secretes a new, very thin and delicate mem- branous cuticle, which may be called the prepupal cuticle. To the inside of this sheath adhere the remains of the tracheal system of the third larval instar. Within this complete or incomplete fourth larval instar or prepupa, the true pupa is now formed by another 7 Most of the usual “ larviparous” Diptera may be regarded as ovoviviparous (Hagan, 1951, p. 59), since the first instar larva is voided shortly after the embryo hatches from the egg. However, in cases such as the Neotropical Mesembrinella and Chortinus, the larva undergoes one or more moults in the mother’s uterus and is of fair size when it is eventually deposited. Evidently it must have done a certain amount of intra-uterine feeding, so that these flies would seem to exhibit incomplete or partial adenotrophy. 94 Volume XXXII moult, producing- a much thicker pupal cuticle, later left behind in the puparium when the fly emerges. The production of a pre- pupal cuticle was first clearly recognized in acalyptrate Muscoidea : in Bhagoletis (Trypaneidae) by Snodgrass (1924, Jl. Agric. Res., 28, pp. 22-23, PL 5, fig. C ; figure reproduced by Wigglesworth, 1950, p. 58, fig. 47), in Drosophila (Drosophilidae) by Robertson (1936, Jl. of Morphology, 59, p. 355), and in Psila (Psilidae) by Ashby and Wright (1946, Trans. Ent. Soc. London, 97, p. 375). Its occurrence in the higher, calyptrate Muscoidea was established by Fraenkel (1938 b) for Calliphora, Lucilia, Phormia, and Sar- cophaga, so that it seems to be a regular feature of all Cyclor- rhapha. It could therefore have been anticipated in the Hippobos- cidae. I was delighted to hear recently from Dr. J. H. Schuurmans Stekhoven ( in lift., 1952) that he observed in Pseudolynchia cana- riensis a moult within the puparium, producing a thin membrane, which generally lies very close to the thick casing ; it may be visible under water, because there is often a bubble of air between the membrane and the thick cuticle of the third instar. The true pupa at first shows externally only the abdomen and thorax, with the wings and legs well formed within their sheaths. At this stage the head lies invaginated inside the thorax. The body of the pupa shows wave-like peristaltic movements, gradually in- creasing in strength, until eventually one of them suddenly evagi- nates the head, which now becomes part of the pupa (Fraenkel, 1938a). Gabler (1935) found on the newly formed inner pupa of Lipoptena cervi a pair of anterior functional spiracles (thoracic spiracular horns). In the pupa of Melophagus ovinus, however, according to de Meijere (1902), they form only two minute rudi- ments, placed rather far back of the head, scarcely projecting and appearing only as scars, without buttons or pits. In the majority of the Hippoboscidae the outer surface of the completed and hardened puparium is bare; but in Olfersia it is covered with stiff, erect hairs, simple in some species, hooked in others. The bare puparia may be smooth and shiny, as in Melopha- gus and Lipoptena. More often the surface is engraved with a mesh of microscopic lines, though otherwise shiny, as in Stenepte- ryx and Pseudolynchia ; sometimes it is in addition more or less punctate or minutely pitted, as in Stilbometopa and Ornitheza. In Lynchia, the engraved lines are deeper and form such a close net- work that the surface becomes alutaceous and dull. The puparium of Hippobosca appears dull and finely punctate under a hand lens ; 95 ENTOMOLOGICA AMERICANA but a higher magnification discloses that the “punctures” are ac- tually raised, very short, triangular spinules. After the adult emerges there is sometimes a certain amount of imaginal growth in the abdomen, at least of the female. The con- siderable increase in size after engorging and during pregnancy may be due entirely to the stretching of the folds of the integument. There may be in addition a true growth of some of the internal or- gans. Theodor (1928, p. 285) observed that, in Lipoptena capreoli (= L. caprina), the mid-gut of a newly emerged ked is 0.9 to 1 cm. long, but reaches 4 to 5 cm. in a ked 14 to 20 days old. Growth is restricted to the mid-gut and is due to the enlargement of existing cells of the wall, not to cell division. The milk-glands likewise grow in size after emergence. NATURAL HISTORY As the term implies, the section on “ Natural History” attempts to bring together what is known of the history of the Hippoboscidae in nature, as distinguished from the structure and functions of their organs. Due to the scarcity of reliable observations on wild flies, it is based to a large extent on information obtained from the relatively few parasitic flies of domestic animals and on the results of laboratory experiments. It should be emphasized that conclu- sions drawn from this type of observation do not necessarily apply to flies living under natural conditions on wild hosts. “Natural history” includes, besides the topics usually covered by the ‘ ‘ ecology ” or “ bionomics ’ ’ of some writers, certain types of instinctive behavior which perhaps are not always nor fully ex- plained as direct responses to the environment. Regardless of whether or not the foregoing three terms are strictly synonymous, I shall use them interchangeably, much as I prefer the more old- fashioned word. While the interrelations of organisms among themselves and with the environment show a bewildering variety, they are all ex- pressions of the three fundamental organic needs for food, protec- tion and reproduction, the first two regulating mainly the survival of the individual and the last that of the species. All three needs are always closely integrated and this is particularly true for in- sects with total viviparity, such as the Hippoboscidae, whose prob- lems of feeding and protection are mostly solved at the same time for the adult and the offspring. All three aspects of hippoboscid life are dominated by the host-parasite relation, the final goal of their activities, which is correlated with nearly every detail of their 96 Volume XXXII morphology and physiology. It will be both rational and conven- ient to group first the various aspects of their natural history under the three customary headings of nourishment, shelter and reproduc- tion, and to conclude with a discussion of their host-parasite rela- tion. I. Food The Blood Diet. It should be made clear at the start that the Hippoboscidae feed exclusively on fresh, fluid blood of warm- blooded vertebrates (homoiotherms) . Under natural conditions, they are not known to drink water or to imbibe surface moisture either on or away from the host. Ormerod’s (1900) statement that Hippobosca equina feeds on the perspiration given off by cattle in the summer, could not have been based on actual observation, since the mouth-parts of Hippobosca are scarcely built to absorb moisture from the surface of the skin. Adult louse-flies remain almost permanently on the food supply and do therefore not live under the same conditions as the tsetse-flies, although their feeding habits are essentially the same.8 When they stray from a live or dead host, either by accident or intentionally, they soon succumb for lack of food and shelter, unless they can reach a suitable new host. Both sexes of the Hippoboscidae suck blood. Prouty and Coatney (1934) attempted feeding various sub- stances artificially to Pseudolynchia canariensis (= P. maura), sim- ilar experiments having been successful with several other blood- sucking arthropods. Pigeon-flies confined in a glass cylinder were allowed access to a feeding tube closed with a membrane and filled with a variety of substances. No fly ever tried to suck cold food. Only the following materials were actually imbibed : fresh pigeon blood, kept at 94° F., on which one fly fed for 7 minutes, the ab- domen becoming engorged and red ; raisin juice ; whole milk ; bovine serum ; and saturated sucrose solution. None of the flies fed arti- ficially survived for a second meal. They died within from 2 to 117 8 It is generally agreed that vertebrate blood is the only normal food of the tsetse-flies and that it is essential for their reproduction. The claim that some species of Glossina may at times add to this in nature either water or vegetable juices, is based on very few obser- vations, which, moreover, are far from convincing. It is true, nev- ertheless, that some tsetse-flies may be made to imbibe water from an artificial feeding apparatus, so that it is not unthinkable that such free-living flies might under unusual circumstances be driven by hunger to attempt abnormal feeding. 97 ENTOMOLOGICA AMERICANA hours, their longevity being about the same as when kept com- pletely starved under similar conditions of temperature and moisture. The restricted nature of the food, at all stages of development, places the Hippoboscidae in a select group of strictly blood-sucking Arthropoda, which includes also the other pupiparous Diptera (Glossinidae, Streblidae and Nycteribiidae), certain small groups of Heteroptera (Polyctenidae, Cimicidae and Triatominae), the sucking lice (Anoplura), the ticks (Ixodoidea), and a few of the parasitic mites. No doubt it is significant that integral intra- uterine viviparity (or so-called pupiparity) occurs only among arthropods with a strict blood diet and is, moreover, confined to in- sects that feed mainly or exclusively on warm-blooded vertebrates. An exclusive diet of warm blood must have favored in some way the evolution of pupiparity to its present perfection, even though it did not perhaps induce it at the start. Vertebrate blood is a concentrated type of food, hence one of the most desirable nutritive substances available. In addition to the highly nutritious essential components (plasma, red and white corpuscles), it is loaded with other organic and mineral ingredients, hemoglobin, vitamins and more subtle biotic substances, continually renewed and carried to all parts of the body. It is kept fluid and under pressure in walled vessels, some of which form a dense maze close to the outer skin, where it is readily accessible and easily in- gested by means of properly constructed mouth-parts. The preva- lence of hematophagy among several unrelated groups is sufficient proof that the Arthropoda have succeeded repeatedly, not only in discovering and reaching this choice source of food, but also in solving the more delicate physiological problems of assimilating it for their own metabolism. A strict diet of fresh vertebrate blood calls for a suitable struc- ture of the mouth-parts, as well as for special anatomical and physi- ological arrangements insuring the free inflow of blood and its proper digestion and assimilation. The description previously given of the proboscis leaves little doubt that the louse-flies are well equipped for blood-sucking. How this came about by evolution is not difficult to understand if we assume, as seems reasonable from the available evidence, that the Hippoboscidae are modified Mus- coidea. Their ancestors acquired from the muscoid stock mouth- parts adapted to a liquid diet, calling only for further specializa- tion. As a result, hematophagous habits originated several times independently among the Muscoidea, probably as soon as vertebrate 98 Volume XXXII blood became available. The original incentive to puncture the skin, in order to tap this new and rich supply of food, may have been the more primitive dipterous habit of sponging either .perspira- tion or other normal secretions of the skin, as well as blood and plasma oozing from wounds. Moreover, the labella of many non- biting Muscoidea bear asperities ( prestomal teeth ), sometimes sharp enough to scratch the skin and thus give the fly its first taste of blood. It should be pointed out that the very efficient, needle-like haustellum of the Hippoboscidae is duplicated almost exactly in the Glossinidae. These two families differ mainly in that the haus- tellum is not retractile within the rostrum membrane in the tsetse- flies, as it is in the louse-flies. This raises the possibility that both groups are rather closely related and that the ancestral Hippo- boscidae may have acquired the present perfected mouth-parts, as well as the exclusive blood diet, before they became permanent ecto- parasites. The more intricate adjustments of digestion, assimilation and general metabolism needed by the change from a vegetable or mixed to a strictly sanguine diet, are not so easily explained. Perhaps they were gradually forced upon the flies during a transitional stage, when the ancestral forms fed at the same time on blood and on plant juices, as some other blood-sucking Diptera do to this day. The physiological peculiarities allowing a strictly blood-sucking insect to thrive on a diet of blood alone, are as yet scarcely under- stood, so that the following discussion is mainly an exposition of unsolved problems. Only two of the several mechanisms involved seem to have been investigated, though not in the Hippoboscidae. When a blood-sucking insect feeds, it mixes with the blood its own saliva containing one or more enzymes, one of which is an anti- coagulin which prevents clotting. This process seems to be a re- finement of an ancestral habit, retained by many other Diptera, of ejecting saliva on solid food in order to change it into a liquid suit- able for ingestion. The precise constituents and action of the saliva of the Hippoboscidae are not known ; but they may be analogous to those of the tsetse-flies, studied by Lloyd (1921) and Lester and Lloyd (1928). When the salivary glands were removed from liv- ing Glossina tachinoides , the flies survived for some time (usually from 7 to 14 days, in one case up to 58 days), meanwhile feeding on man and digesting normally, sometimes even producing healthy larvae. Glandless flies fed on fowl did not thrive so well. Even- tually, however, such flies found it difficult or impossible to feed and particularly to digest, as the blood remained in the proboscis 99 ENTOMOLOGICA AMERICANA and clotted in the crop, fore-gut and anterior portion of mid-gut, causing death by starvation. The experiments showed that injec- tion of saliva in the host’s tissues is not an essential preliminary to the act of feeding, but is rather incidental, since the salivary secre- tion and the blood mix at the very tip of the haustellum. The function of the saliva and of the anticoagulin it contains is to pre- vent the clotting of blood in the lumen of the haustellum, crop, fore- gut and anterior part of the mid-gut, so that the food remains fluid until it reaches the region of the mid-gut where digestion occurs. Saliva plays no part in digestion proper, as is shown also by its lacking digestive enzymes (Wigglesworth, 1929; 1950, p. 329). In the tsetse-flies and presumably also in the Hippoboscidae, digestion and absorption are carried on in the posterior two-thirds only of the mid-gut, where the epithelium secretes a powerful co- agulent and various digestive enzymes. A first question is whether the mid-gut enzymes alone are adequate for digesting blood or must be supplemented for this purpose from some other source. What digestive enzymes are produced by the louse-flies is again unknown. In Glossina, according to Wigglesworth (1929; 1950, p. 331), protease, which digests proteins (some of the main constituents of blood), is very active in the mid-gut, while carbohydrases are absent except a very feeble amylase in the same region. It should be stressed that digestion is rapid in the pupiparous Diptera. Even tsetse-flies, which imbibe If to If times their own weight of blood at one meal, usually feed again within 30 hours. A second problem is how an exclusive diet of blood may provide an insect with all the organic substances required for growth, reproduction and energy production, including the energy consumed in locomotion and in flight by the winged species. It is, of course, evident that the Hip- poboscidae and other pupiparous Diptera thrive precisely on this type of diet; but exactly how do they achieve this? To what ex- tent and by what method the various constituents of vertebrate blood are broken down for assimilation by insects have scarcely been investigated ; although it is known that in certain blood-suck- ing insects hemoglobin disintegrates only partially, the remaining product being excreted directly. It is now generally recognized that sterile vertebrate blood, the only food directly available to the Hippoboscidae and other pupi- parous Diptera, is an inadequate or incomplete diet, since it is de- ficient in certain accessory substances, particularly certain vitamins, which are essential for growth and sometimes for normal reproduc- tion (Wigglesworth, 1950, p. 348; 1952, p. 66). In some blood- 100 Volume XXXII sucking insects these substances seem to be produced by micro- organisms living freely in the intestinal tract. It is difficult to conceive how the gut of the Hippoboscidae could become infected with outside bacteria or yeasts; such organisms might presumably contaminate the labella, when these come in contact with the surface of the skin, and then be taken up with the blood. Actually the lumen of the mid-gut of certain louse-flies harbors extracellular, presumably anaerobic and saprophytic micro-organisms. Rickett- sia melophagi Noller (1917), a unicellular, spheroidal or rod-like organism, which has been the subject of many investigations (see J. Bequaert, 1942a, pp. 195-196, for references), occurs in most sheep-keds, in Europe and North America, on the inner surface of the mid-gut, and is transmitted from the adult to the larva with the secretion of the milk-glands, surviving pupation. It does not seem to affect the health of the sheep-ked adversely, but on the other hand is not known to play a role in digestion. Aschner (1931, pp. 397- 400) found a somewhat similar Rickettsia- like organism under the same conditions in Lipoptena capreoli ( = caprina ). He also described (1931, pp. 394-395) from the mid-gut of Pseudolynchia canariensis (= Lynchia maura) slender, thread-like bacteria, mostly free between the peritrophic membrane and the epithelium, but some also inside cells of the muscular coat surrounding the wall. His account of these organisms is rather confused and that of simi- lar bacteroids in Hippobosca camelina (1931, pp. 395-397) is even less convincing, since he states that in this species they occur al- most always inside epithelial cells. I suspect that these intestinal bacteroids of Pseudolynchia and Hippobosca might be only intra- cellular symbionts, such as will be discussed later, seen at a tem- porary extracellular stage. Steinhaus (1943; 1946, p. 44, fig. 15, and p. 245) described as Cory neb act erium lipoptenae rod-like bacteria found chiefly in the lumen of the intestinal tract of all of 10 Lipoptena depressa taken from a deer near Hamilton, Montana. Present evidence is altogether too slight to assume that the extra- cellular micro-organisms found normally in the mid-gut of certain louse-flies either assist in digestion or produce accessory food sub- stances. Intracellular Symbionts and Nutrition. In several types of insects every individual contains specific micro-organisms packed in specialized cells, so-called mycetocytes, usually of a characteristic shape. These cells are often grouped to form mycetomes, always located in or near the same organs in every species. Almost in- variably some special mechanism assures the transmission of the or- 101 ENTOMOLOGICA AMERICANA ganisms from one insect generation to the next. Because of these and other peculiarities, the intracellular organisms of the myceto- cytes are now regarded as more than tolerated, innocuous parasites, but rather as messmates or mutualistic symbionts (symbiotes; endo- symbionts), useful somehow to the insect. It is noteworthy that they occur generally in the strictly blood-sucking Pupipara and that the mycetomes of these flies are usually associated with the wall of the mid-gut. Since they have been found in all genera of the Hippoboscidae examined for the purpose, it is safe to assume that they occur throughout the family; indeed, Buchner (1952, p. 148) regards their presence as one of the characteristic adaptations of these flies, inherited from their early ancestors. Mycetomes were first recognized by Sikora (1918) in Melophagus ovinus, where they have been frequently studied ( Jungmann, 1918 ; Hertig and Wolbach, 1924; Anigstein, 1927; Zacharias, 1928; Glaser, 1930 ; Aschner, 1931 ; for more complete resumes, see Buch- ner, 1930, pp. 609-614; J. Bequaert, 1942a, pp. 196-197). The intracellular symbionts of the sheep-ked are of two types. All keds of both sexes examined in Europe and North America contain spheroidal or short rod-like organisms, which fill most of the proto- plasm of enlarged epithelial cells. The infected cells form a large, ring-like mycetome at the bend of the right-side coil of the mid-gut (Fig. 15, my), along one-eighth to one-tenth of the length of the abdominal mid-gut. The symbionts are not “inherited” through the egg, but are transmitted with the secretion of the milk-glands to the larva, where they are stored until pupation in enlarged cells (mycetocytes) forming the epithelium of the anterior tubular por- tion (proventriculus) of the mid-gut. During pupation the larval mycetocytes are freed from the epithelium and maintain their indi- viduality at first in the anterior part of the mid-gut. Zacharias (1928, pp. 695 and 703) states that about this time some of the mycetocytes break down, releasing symbionts which penetrate through the intestinal epithelium into the body cavity, where some were observed. He suggests that these symbionts might migrate then to the milk-glands, presumably passing through the secretory cells of the glands, although none were ever found there. In the mid-gut of the pupa the larval mycetocytes migrate backward and eventually break down, releasing the symbionts which now enter some of the epithelial cells of the newly-formed adult mid-gut. Zacharias (1928, pp. 705-711) described from some only of the European sheep-keds a second, more slender, thread-like symbiont. This fills ordinary, flat, epithelial cells, usually of the mid-gut, more 102 Volume XXXII rarely of the Malpighian tubules, and is said to be transmitted in the same way as the other type. In Lipoptena cervi, symbionts were noticed first by Roubaud (1919, p. 532) and later by Zacharias (1928, pp. 711-717), in a mycetome located much as in Melophagus. They are also of two types and transmitted in the same manner. According to Aschner (1931, pp. 376-380), the symbionts of Lipoptena capreoli (= L. caprina) are in every respect like those of L. cervi. In Lipoptena, the mycetome extends over one-eighth to one-tenth of the length of the abdominal mid-gut. The mycetome of Ornithomyia biloba, studied by Zacharias (1928, pp. 718-725; as 0. avicularia9) , differs markedly from that of the Melophaginae. The freshly dissected mid-gut of newly- emerged, unfed flies shows no distinct mycetome, but is swollen at about mid-length over a short stretch. When the gut is stained with borax carmine, the swelling discloses two lengthened, darker- stained areas separated by two narrow, paler bands. In cross- section, the epithelial cells of the dark areas are normal and contain no symbionts • but the layer beneath the epithelium is a mycetome, the several cells of which are fused into a syncytium with many nuclei and are filled with long, thread-like, twisted organisms, interwoven with the fibrillae of the muscle coat. According to Mr. I. B. Tarshis (in litt ., 1951), the mycetome of Stilbometopa impressa is located as in 0. biloba and has a somewhat similar structure. It shows, however, certain interesting peculiari- ties which he will describe in a future paper. Aschner (1931, pp. 378-380) found that the mycetome of Pseudo- lynchia canariensis (= Lynchia maura) extends over a relatively short stretch of the mid-gut (about one-twentieth of the length of the abdominal part), where the epithelial cells are greatly enlarged. However, the symbionts in this case fill only the apical portion of the cells, nearest the gut cavity, and are disk-like or vesicular and diffi- cult to stain. Similar organisms were found in the secretion inside the lumen of the milk-glands. In Hippobosca equina , H. longipennis (=H. capensis), and H. camelina also, Aschner (1931, pp. 380-390) found symbionts in 9 Zacharias’ flies were doubtless Ornithomyia biloba Dufour, since they were bred from puparia found in abandoned swallows’ nests in Germany. Although often recorded by error as a parasite of the European barn swallow ( Hirundo rustica), 0. avicularia does; not occur normally on this host or in its nests. 103 ENTOMOLOGICA AMERICANA enlarged epithelial cells; the mycetome is very extensive, being continuous over six-tenths, five-sixths or six-sevenths of the length of the abdominal mid-gnt. The organisms of H. equina are mostly rod-like and congregate in the basal portion of the cells, not apically as in Pseudolynchia. They are variously shaped in H. longipennis and H. camelina. In Hippobosca also the lumen of the milk-glands is infected with micro-organisms. The symbionts of H. equina were first seen by Roubaud (1919, p. 532), who regarded them as intracellular yeasts, and later also by Zacharias (1928, p. 717) . Aschner (1931, pp. 412-416) attempted to grow the intracellular symbionts of Melophagus, Lipoptena, and Hippobosca outside the insects, but without success. The extracellular Rickettsia- like organisms of the gut of Melophagus ovinus have been cultivated on various media by several investigators; Kligler and Aschner (1931) have grown successfully, not only those of M. ovinus , but the similar organisms in the gut of Lipoptena capreoli (= L. caprina), Hippo- bosca longipennis (= H. capensis), and H. equina as well. The following peculiarities of the intracellular symbionts sug- gest a possible connection with digestive or other metabolic processes of the flies: (1) the regular occurrence in all strictly blood-sucking species, including besides the Hippoboscidae all the other pupipar- ous blood-sucking Diptera, and their absence in blood-sucking flies with a different larval diet; (2) the elaborate mechanism ensuring their transmission to the offspring of the fly; (3) the location of the mycetomes in the epithelium or the muscle coat of the mid-gut. Although such a combination of peculiarities in the same insect is unusual, it should be noted that, taken individually, some of these features occur also in other types of arthropods and for other types of micro-organisms. They may therefore be incidental and without any particular significance to the insect. Possibly the ^symbionts merely take advantage of the unique digestive set-up of the pupipar- ous flies, without giving much if anything in return. Intracellular symbionts are fairly widespread among the Arthropoda, even among the non-hematophagous types. Probably they originated from some of the extracellular, saprophytic or mildly parasitic micro- organisms, found frequently in the mid-gut of arthropods. Even- tually some of the organisms invaded the epithelium of the wall of the gut, causing the tumor-like enlargement of certain cells or the formation of a syncytium, features which may have become heredi- tary in certain cases. The location and extent of the mycetomes vary with each species, possibly indicating that in certain areas of the gut the epithelial cells were originally more vulnerable, more ac- 104 Volume XXXII cessible to invasion, or more suitable to the growth of the micro- organisms. As for the mechanism of transmission, this is actually no more intricate than that of many other parasitic micro-organisms of vertebrates transmitted by arthropods. Moreover, it is often nearly impossible to draw the line, either on morphological or on biological grounds, between the so-called “symbionts” and other innocuous or pathogenic micro-organisms occurring sometimes in the same species of arthropod. On the other hand, there is ample proof that certain free-living, saprophytic or parasitic micro-organisms, morphologically similar to the symbionts of the Arthropoda, produce powerful biotic sub- stances, such as hormones, vitamins and enzymes, or are able to fix di- rectly free atmospheric nitrogen into organic products and to break down nitrogenous organic substances at least to the ammonia stage. This makes it most probable that the endosymbionts of the blood- sucking insects influence somehow the metabolism of their hosts. The possible physiological role of the symbionts, particularly those of arthropods with a specialized food diet, is now much in the lime- light. Comprehensive accounts have been published by Buchner (1930), Aschner (1931, pp. 416-439), Steinhaus (1946, pp. 188-255; 1949, pp. 123-165), and Wigglesworth (1950, pp. 345-349). Latterly the whole subject was discussed from many angles by several specialists at a colloquium on the occasion of the IXth International Congress of Entomology (August, 1951) at Amsterdam (1952, Tijdschr. v. Entom., 95, pts. 1-2, pp. 23-196). The reader must be referred to these sources for the details, which seem irrelevant in the present work, since none of the observations and experiments refer directly to the symbionts of the Hippo- boscidae. Discussing the possible function of the symbionts in strictly blood-sucking insects, Wigglesworth (1952) concludes, from obser- vations in the tsetse-fly ( Glossina ) and blood-sucking Reduviidae ( Triatoma ; Rhodnius ), that the organisms are most probably not concerned directly in the digestion of the blood. There is evidence, however, in the sucking lice ( Pediculus ) and Reduviidae ( Triatoma ; Eutriatoma ; Rhodnius) , that they influence growth, metamorphosis and reproduction. It seems likely that they supply certain growth factors, presumably vitamins, in which sterile blood is deficient. Differential Feeding and Host Specificity. Blood is an ex- tremely complex substance. It also shows physical and chemical peculiarities among the several vertebrates, aside from the more obvious differences in the size, number and structure of the corpus- 105 ENTOMOLOGICA AMERICANA cles. The proportion and composition of the mineral and organic components of the plasma are decidedly specific and fairly constant for each species. For instance, in the case of the blood proteins, particularly important elements in the diet of blood-sucking insects, not only does their concentration vary with the species, but they show also a more subtle serological specificity brought out by their antisera-antigen reactions. The differential properties of the blood of the several available hosts no doubt influence to some extent the ectoparasite’s metabolism. In the Pupipara, which feed on blood only, such differences may have an important survival value, certain types of blood being better suited for continued, normal reproduc- tion and others perhaps wholly unsuitable. Conceivably the effect of the type of blood diet upon the rate of reproduction has been the primary factor inducing by natural selection whatever host specificity is now observed in the Hippoboscidae. Other factors, such as the opportunities for shelter and the peculiarities of the ecology and behavior of the vertebrates, certainly contributed also toward determining the true breeding hosts of the flies ; but I am inclined to regard them as of secondary importance. It is unfortunate that so little attention has been paid to the possible role of differential feeding in the normal life of blood-suck- ing arthropods. A few, rather unsatisfactory experiments are available for the tsetse-flies. Lloyd (1914), in Northern Rhodesia, fed in captivity series of freshly caught, wild female Glossina mor si- tans separately on goats, monkeys, fowls, ducks and chameleons, recording the length of life of each fly, and the number of puparia and aborted larvae deposited, each puparium also being measured. Flies fed on chameleons did not produce larvae, perhaps because they failed to obtain a satisfactory blood meal. It was concluded that mammalian blood has a slight advantage over that of birds, as it seemed to be more easily digested. Puparia obtained on mam- malian blood averaged larger than those produced by flies feeding on birds.10 I can find no experimental data bearing on the influence of various types of blood upon the reproductive rate of the Hippo- boscidae. Coatney (1931) fed the pigeon-fly, Pseudolynchia can- i° rppe resuits of Lloyd’s experiments with G. morsitans have been criticized adversely on the basis of observations (not experi- ments) with G. palpalis. However, if the effects of differential feeding are specific, as may be suspected, they should not necessarily be the same for all species of Glossina. 106 Volume XXXII ariensis, for about 2 weeks on domestic fowl in the laboratory, obtaining normal pnparia meanwhile ; but he does not compare his results with flies fed under similar conditions on the normal host. Kemper’s (1951, p. 239) experiments with Crataerina pallida fed on abnormal hosts, particularly the domestic pigeon, furnish no information on the present topic. Mechanism of Feeding. Two hundred years ago Reaumur (1742, p. 602), in France, described as follows how Hippo~bosca equina bites and sucks blood : “If I had been uncertain how these flies fed, if I had doubted that the very thin thread sometimes ap- pearing in front of the head was actually a proboscis, as I have called it, my doubts would have been removed by an experiment performed without my trying. In spite of my efforts several spider- flies escaped from a pounce box where I kept them and from which I intended to remove one only. One of those that escaped did not go very far ; after a fairly short flight it alighted on my arm. I was careful not to chase it away; I wanted to know whether it might not like to pierce my skin the same as that of a horse or an ox : it did not pause long to teach me that my blood was to its taste ; it pro- truded the thin thread which I have called the proboscis and almost at once thrust the tip in my skin. The bite was felt no more than that of a flea. Horses must bear the bites of these flies with com- posure, if they are not more painful to them than this one was to me. The fly sucked my blood without stopping for nearly a quarter of an hour and during all this time I felt only a strong itching. The wound left open after the fly departed was marked only by a small red spot, which disappeared in less than half an hour and which produced no raised area : from which it follows that these flies are not as dreadful as mosquitoes, which do not fail to envenom the wounds they make. The fly was so engrossed with imbibing my blood, that it let me take my hand to a spot lighted by the sun’s rays, where I could observe it at leisure under a lens with a 6 to 7 lines focus. At first the fly pushed in more and more of the pro- boscis ; but once this had penetrated as far as wanted and apparently as far as possible, it was pulled out a little and then pushed back again as far as before. This process was repeated several times, but at unequal intervals. I saw then that the proboscis started from a sort of membranous pouch, oblong in shape and thicker close to the head than elsewhere. As long as the fly kept the pro- boscis inserted in my flesh, the two pallets [palpi] that enclose it in a sheath were spread apart in a rather broad angle, allowing the por- tion of the proboscis kept between them to enter the flesh. The fly 107 ENTOMOLOGICA AMERICANA left only after being fully engorged, when its gaster was distended and tight with the blood that filled it.” It may be interesting to compare the French naturalist’s account with the following, more recent observations made on the same species in Spain by Hase (1927, pp. 218-233). When the proboscis is not in use, most of the haustellum is retracted within the rostrum membrane, only the apical portion with the labella being hidden within the slightly hollow sheath of the palpi. A fly about to feed takes a firm hold of the skin by anchoring the deeply divided, strong claws to the hairs or sometimes to the minute folds of the epidermis. Next it explores the surface, at first with the setiferous tips of the palpi, later also with the labella as soon as the tip of the haustellum protrudes from the slightly spreading palpi. During the preliminary probings the labella are often covered with a clear liquid, presumably saliva oozing from the hypopharynx. Differences in the temperature or toughness of the skin may possibly guide these explorations. A further eversion of the rostrum membrane frees the entire haustel- lum from the palpi, which do not guide its course nor take an active part in feeding. The prestomal teeth of the labella do the actual cutting of the skin, after which the haustellum bores in rapidly under the pressure of head and thorax, meanwhile bending forward and sideways, as well as twisting. The whole or part of the needle- like portion of the haustellum enters the skin, until the labella reach a capillary of appropriate size in the stratum corneum. If a suit- able capillary is not found at a first try, the fly removes the haustel- lum and bores at another spot. The protruding portion of the ros- trum membrane shows active sucking motions during the actual imbibing of blood. Near the end of the meal, the fly voids excreta from a previous digestion and eventually also some of the freshly absorbed blood. Cleaning motions with the legs, starting over wings and eyes, indicate near-satiation, the fly preparing to leave the spot of the bite. As a rule the haustellum is then easily and swiftly removed; but sometimes it can only be pulled out with an effort. Once feeding is well under way, the fly is not easily dis- turbed and ignores most of the stimuli to which it responds quickly at other times. Hase ’s drawings of six successive stages of the bit- ing process are reproduced in my Fig. 6H. The duration of a complete blood meal varies greatly: in Hase’s experiments several flies engorged respectively within 3, 7-J, 10, 14, 20 and 24J minutes (15 minutes on the average) ; one female fed without interruption for 43 minutes. Jobling (1926, p. 345) describes the feeding of Melophagus 108 Volume XXXII ovinus on Man in the laboratory : 1 ‘ Before the ked starts feeding it must attach itself to the host. As the human skin is very poor in hair much time is required before the ked becomes fixed. The ked then protrudes the haustellum between the maxillary palps, apply- ing the end of the labellum to different points of the skin, apparently in search of a suitable spot for piercing. When such a spot is se- lected the insect pushes its proboscis against the skin with such force that the middle part of the haustellum bends into a bow. Some- times in piercing the ked twists the haustellum as one twists an awl. Immediately afterward there can be observed forward and back- ward movements in the labellum, the distal part of the proboscis rapidly penetrating into the skin. The haustellum penetrates into the skin as far as two-thirds its length. The appearance of a red spot under the vertex indicates the blood being sucked into the •oesophagus. All the keds fed under observation were gorged in about 10 minutes after the appearance of the red spot in the region of the oesophagus. It has also been observed that forcible removal of the feeding ked involves the risk of tearing off the haustellum, as it is fixed by the everted prestomal teeth to the bottom of the hole formed in the skin. I felt no irritation during the feeding of the sheep-keds, but four or five days later small yellowish papules made their appearance. Bach had a very minute opening through which serum exuded. The irritation caused by these papules is not very severe, but they heal very slowly, disappearing three or four weeks after the feeding of the insects. ” Brumpt (1913, p. 638) states that Lipoptena cervi engorges on Man in 12 to 20 minutes. Hase (1939, p. 414), in Germany, also made hungry, volant L. cervi bite Man, using flies caught in the open in November and brought to the laboratory. They engorge within 15 to 25 minutes. Once a suitable spot is selected by feeling with the palpi and probing with the haustellum, the latter is inserted in the skin, either obliquely or at a right angle, by pushing the head downward or sometimes also by forward thrusts of the whole body. While feeding, the fly remains as if “ frozen ’ ’ on the spot and is not disturbed by either touch or pressure or by moving its legs. When placed on mice, L. cervi attaches behind the ears and to the neck, moving to the darker belly when engorged without attempting to leave the animal. The mice seem to be annoyed not so much by the bite as by the crawling of the flies. One mouse succeeded in catch- ing and chewing two of the flies but did not actually eat them. The deer-ked likewise starts defecating while feeding and eventually the feces contain some undigested or partly digested blood. 109 ENTOMOLOGICA AMERICANA It is rather unfortunate that the foregoing observations on mam- mal-infesting flies were made on abnormal hosts (Man and mice) and usually performed in the laboratory. There is no certainty that they conform in every detail with normal feeding under natural conditions on the regular hosts. In particular, differences in the texture and thickness of the skin, density and nature of the pelt, and the individual reaction to the bites may influence the method and speed of feeding. There are as yet few comparable observations of bird-flies. Kemper (1951, p. 236) states that if Crataerina pallida, kept with- out food 3 or more days in the laboratory, are put on the plumage of a swift, Apus apus, they usually slip at once under the feathers and presumably start feeding immediately. They select to some extent the belly and the neck, and have not been observed biting the head. Within 7 to 15 minutes (about 10 on the average) the flies reappear, mostly engorged, run for a short while over the body and eventually hide in the plumage, as a rule beneath the wings. Kemper induced the swift fly to feed on a tree sparrow, Passer montanus, a great tit, Pams major, and domestic pigeons. He observed that the fly did not enter the plumage as quickly on these strange birds as on the swifts. The duration and size of the meal were the same as on the normal host and the longevity of the flies seemingly was not impaired. No information is given as to the production of puparia. Attempts to feed the swift fly on guinea pigs and white mice failed. Noller (1920, p. 166) was unable to make Ornithomyia biloba (cited as 0. fringillina) bite a crossbill, Loxia c. curvirostra, using flies taken from barn swallows, Hirundo rustica, or bred from puparia found in swallows’ nests. Even when starved for 3 to 7 days, the flies refused to bite. This would seem to point to a strong selectivity in the choice of a blood supply by this species. Schuurmans Stekhoven and his co-workers, in Argentina, re- cently observed how Pseudolynchia canariensis feeds on the domestic pigeon (unpublished observations). Newly-emerged flies fast one or more days before attempting to bite, while older flies taken from birds sometimes feed within a few seconds after being placed in a vial on the skin. After probing the skin with the palpi, they pro- trude and push in the haustellum. Meanwhile the fly rises on its hind legs, tilts the body forward, and rests on the fore tarsi and the strongly bent fore femora and tibiae. The palpi are spread out on the skin. The fly engorges within 5 to 10 minutes, during which time it is difficult to disturb. If the meal is interrupted, the fly usually starts feeding again at another spot. Defecation by feeding 110 Volume XXXII flies was not observed. Huff (1932) kept P. canariensis for 2 to 8 days on the American mourning dove, Zenaidura macroura carolin- ensis, the fly feeding meanwhile. Kartman (1949), in Hawaii, also fed pigeon-flies on the Chinese dove, Streptopelia chinensis, and the ring-necked dove, S. decaocto; some were kept alive from February 29 to March 29 on the Chinese dove and from May 12 to July 4 on the ring-necked dove. Olfersia aenescens is a specific fly of certain Oceanic birds. Kartman (1949), in Hawaii, placed 20 flies taken from a red-footed booby, Sula sula rubripes, in a cage with a domestic pigeon on February 29. Most of the flies showed no interest in the pigeon and died within a few days. On March 10, however, one surviving 0. aenescens was recovered from the pigeon and showed fresh blood in the mid-gut ; two other flies, bred from puparia, remained alive on a pigeon from April 24 to May 4. Size of Blood Meal. Until now it has been generally believed that the diverticulum or storage crop of the fore-gut is atrophied and non-functional in all Hippoboscidae, and that the amount of blood they absorb at one meal is therefore limited by the capacity of the fore-gut and of the anterior, widened portion (duodenum) of the mid-gut. While this may be true for the sheep-ked, Melopha- gus ovinus , and possibly for other mammal-infesting flies, recent, unpublished observations by Mr, I. B. Tarshis show that in some of the bird-flies, particularly in Stilbometopa impressa, a certain amount of blood enters the diverticulum toward the end of the meal. Even in this case, however, the quantity of blood stored in the crop is relatively small and the fore and mid-gut remain the chief food reservoirs. Nevertheless, the total amount of blood imbibed at one meal may be very large, but varies greatly even under the best feeding conditions. In Hase’s (1927) experiments with starved Hippobosca equina fed on Man, one female weighed 31 mg. before and 35 mg. after a full meal, having gained 4 mg. or about 1/8 of its original weight. These figures were 31 mg. before and 43 mg. after feeding for an- other female, which gained 12 mg. or 1/3 of its weight. Freund and Stolz (1928) found that Melophagus ovinus imbibes from 8 to 14.2 mg. of blood at each meal. A newly emerged ked nearly doubled its original weight, of 15.1 mg., to 29.3 mg. after feeding 10 minutes. In Kemper’s (1951) experiments with M. ovinus fed on Man, 10 females imbibed at one meal slightly more than 10 males, the females gaining 3 to 15 mg. (8.1 mg. on the aver- age) and the males, 6 to 13 mg. (7.4 mg. on the average). Ill ENTOMOLOGICA AMERICANA Kemper (1951, p. 237) determined the size of the blood meal of the swift fly, Crataerina pallida. In this fly the length of the com- bined head and thorax is about the same in both sexes, but the abdomen is much wider and also longer in the female than in the male, whether unfed or fed. The female averages 7.41 mm. (6.9 to 8.1) in length and 3.15 mm. (2.9 to 3.5) in width when starved, 9.15 mm. (8.0 to 9.9) in length and 6.89 mm. (5.5 to 8.4) in width when engorged. The corresponding measurements of the male are 7.19 mm. (6.3 to 7.7) in length and 3.12 (2.9 to 3.5) in width, unfed, and 8.03 mm. (7.4 to 9.1) in length and 4.69 mm. (4.3 to 5.3) in width, engorged. Flies kept starved for 5 days increased their weight with one blood meal from the original 22.7 mg. to 59.0 mg. in the female, and from 18.4 mg. to 31.4 mg. in the male, in an experiment using 10 flies of each sex. Occasionally even more blood may be taken np in one meal : a female which had engorged more fully than usual weighed 91 mg. ; after fasting 5 days its weight was down to 37.5 mg. ; on the 8th day it voided a premature larva of 16 mg. ; when it died on the 10th day it weighed only 12 mg. f it therefore had taken up at least 60 mg. of blood at one feeding, nearly 5 times the original unfed weight. The amount of blood ingested by the hippoboscids compares favorably with the blood meals of the tsetse-flies, although the latter are provided with a large and fully functional storage crop. Macfie (1912) concluded, from experiments with Glossina palpalis, that the male of this fly imbibed on the average 27 mg. of blood at one meal and the female slightly more, or nearly If times the unfed weight for the male and If times for the female. Frequency of Meals. Since permanent ectoparasites have the food supply within easy reach at all times, there seems to be no reason why they should not feed at fairly regular intervals, regard- less of day or night, the changing seasons or the periodical activities of the host. However, little is known as yet of the time required to digest a full meal. Due to the feeding needs of the developing larva, digestion is probably more rapid in the female hippoboscids than in the males. The undigested material of the blood and the products of excre- tion are eventually evacuated as feces. In many cases defecation starts toward the end of a feeding. Fecal matter first appears at the anus as a droplet of soft or semi-liquid material which hardens in the air, presumably by drying. A new droplet pushes the first farther out and the successive droplets usually adhere in bead-like strands, which eventually break up in shorter strands or in individ- 112 Volume XXXII ual pellets or feculae. When the fecnlae are dry, they have a char- acteristic shape. Hase (1927, pp. 226-228, fig. 15) describes them for Hippobosca equina , and in a later paper (1939, p. 415, fig. 2) for Lipoptena cervi; Kemper (1951, pp. 245-246, fig. 7) for Crataerina pallida and Melophagus ovinus. Kemper observed that the feces of M. ovinus are softer than those of C. pallida and consequently build up only very short strands (about 3 mm. long), or none at all. On the other hand, the fecal strands of C. pallida may reach 15 mm. in length before breaking up. The feculae of Pseudolynchia canarien- sis will be described in a forthcoming paper by Schuurmans Stek- hoven and his co-workers, in Argentina. Hase (1927, pp. 230 and 232) found that one of his TLippobosca equina, fed on Man, refused to feed the first, second and third days after repletion, but fed readily on the fourth; another fly fed eagerly the third day after a full meal. Hase (1939, p. 411) also kept winged Lipoptena cervi, captured in the open, alive from 6 to 8 days in a humid atmosphere at 27° C., when they would suck blood every day on Man or on mice. At room temperature the flies died within 3 days. At 5° C. they became stiff and digestion was arrested, to be resumed when the temperature was raised. Some flies were fed on Man 6 or even 8 times on successive days and seemed to stay healthy. Four flies that had engorged the same day on Man, fed once more on a mouse the next day and again on Man the third day. Kemper (1951, p. 237) observed that in Crataerina pallida four to five days elapse between consecutive feedings. In this connection, Mr. I. B. Tarshis {in litt., 1951) made the interesting discovery that in Stilbometopa impressa the feeding re- quirements are not the same for both sexes. In the male the gut is at all times generally distributed in the abdomen, so that feeding may be resumed as soon as digestion is completed. In the female, however, while a larva is developing in the uterus, the gut is pushed far forward toward the fly’s waist, thus making room for the devel- opment of the very large offspring. Throughout the period of gestation no blood is taken in by the female. She feeds once after voiding a larva and this blood meal lasts her until her next feeding after the deposition of the next larva, and so on. Just after feeding the female ’s gut is well distributed throughout the abdomen, as it is at all times in the male. The information at present available on the size and frequency of the meals is inadequate to calculate the total amount of blood re- quired for the full development of one larva, from the hatching of the egg until parturition, as well as the amount imbibed during the 113 ENTOMOLOGICA AMERICANA average life of a fly. Such data might be useful to determine more accurately the damage done by feeding alone to the well-being of the host by an average hippoboscid population. Resistance to Starvation. The ability to survive without food for some time must often be important to ectoparasites and partic- ularly to forms emerging away from the host, as is the case for nearly all Hippoboscidae (except Melophagus and perhaps a few others). It would seem that as a rule newly-emerged, unfed flies withstand starvation better than those that have previously fed on a suitable host. Curtice (1890) and Swingle (1913), in North America, con- cluded that Melophagus ovinus, removed from sheep and exposed to normal atmospheric conditions, dies within 3 or 4 days. In care- fully controlled experiments, in Australia, Sweet and Seddon (1917) found that keds live from 2 to 11 days away from the host. They appear to stand dryness and lack of food best when the tem- perature is moderately low and uniform. Moisture is needed if extremes of temperature are to be withstood. In shorn or shed wool, life is always short and seems to be little influenced by the fed or unfed condition. Hill (1918), in Australia, experimented with large numbers of adults, unfed and fed, as well as of different ages, and under a variety of conditions. He states that they may survive away from the host longer than had been generally believed. One ked, kept in a cellar, lived for 18 days ; two for 14 days under simi- lar conditions ; a few others, in a more natural environment, for 11 to 13 days. These were, however, exceptional individuals, for his tables show that 75 to 80 per cent of the starved keds died within 1 to 6 days. Graham and Taylor ( 1941 ) , also in Australia, obtained similar results : newly-emerged keds survived in unfed condition for 4 to 5 days at 16° to 30° C. At lower temperatures life was pro- longed to 14 to 24 days ; while higher temperatures reduced it to as little as one day. Kemper (1951, p. 247), in Germany, was unable to keep fully-engorged sheep-keds alive for more than 7 days in a thermostat at 20° to 27° C., the majority dying within 5 days; but he may have kept the keds under too dry conditions. Cowan (1943, p. 186) experimented with Lipoptena depressa in British Columbia. Survival tests were conducted upon newly- emerged flies and also upon sexually mature individuals removed from the host. Seven newly-emerged flies kept in an incubator at room temperature over damp sand, survived for 72, 48, 52, 60, 62, and 52 hours, respectively, the average for this sample being 57 hours. Individuals taken from a deer started dying approximately 114 Volume XXXII 24 hours after removal, the last surviving 6 days. Of 201 flies re- moved from the head and neck of a buck shot October 8, 90 per cent were dead 48 hours later. Flies seem to survive somewhat longer when left upon the dead host, a single living specimen being observed after 11 days, and several taken alive after 6 days and 8 days under such conditions. According to Cowan, the short survival time of the winged individuals must be a potent factor in limiting the popu- lation in areas where deer are scarce. Hare (1945a, p. 56), in California, also working with L. depressa , determined that newly- emerged adults may survive away from a host much longer than Cowan thought, if kept under proper conditions. Volants reared from normal puparia collected on a captive deer survived from 4^ to 8 days without food. Those caught in the open lived only up to 3| days if kept under daylight conditions, but up to 6 days if kept in the dark. In his experiments, 50 per cent of the reared flies settled successfully on a host 4 days after emergence. He con- cluded that under natural conditions volants might remain active for from 1 to 4 days. Theodor (1928), in Palestine, could keep newly-emerged Lipoptena capreoli '( = L. caprina) alive for at most 2 days without feeding. Beebe (cited by Bequaert, 1942a, p. 130), in British Guiana, kept Lipoptena mazamae alive for 3 days after removal from the host. Hase (1927, p. 200), in Spain, kept normal, hungry Hippobosca equina, removed from the host, alive usually for 3, exceptionally for 4 days in a dry atmosphere, and for up to 6 days under moister con- ditions, at 22° to 25° C. He believes that this fly withstands starva- tion better when the air is very humid. In experiments with Hippo- bosca variegata (=H. maculata), in Indonesia, Schuurmans Stek- hoven found that most flies, kept away from the host without feed- ing, died within 1 or 2 days, the males usually somewhat before the females ; a few females survived up to 3 days. In Prouty and Coatney’s (1934) experiments, in Nebraska, newly-bred Pseudolynchia canariensis ( =P. maura ) survived unfed for approximately 109 hours if left undisturbed; if disturbed by shaking, they lived for only 75 to 92 hours. Ed. and Et. Sergent (1909), in North Africa, had previously determined that this fly survives only 2 or 3 days when kept starved. A forthcoming paper by Schuurmans Stekhoven and his co-workers, in Argentina, de- scribes further experiments on the resistance to starvation of pigeon- flies, most of them surviving 4 or 5 days. Falcoz (1922, p. 228) kept 4 Ornithomyia biloba (cited as O. 115 ENTOMOLOGICA AMERICANA avicularia), taken from a swallow’s nest in France, alive without food for 6 days. Galli-Valerio (1930, p. 214; 1932, p. 132), in Switzerland, noted that Cmtaerina pallida could live without food for from 3 to 5 days after being removed from swifts, Apus apus. Kemper (1951, p. 247 ) was able to keep this fly alive much longer in starvation : of 30 specimens kept at from 21° to 24° C. indoors, 23 lived 5 days and one survived 9 days ; one exceptional overfed female lived for 13 days. Cannibalism. The only information available on this point is a rather vague statement by Eysell (1924, p. 387) : “In 1904 I made the interesting observation that hungry louse-flies bore into the blood-filled gut of members of their own species and imbibe the contents. The wound healed normally and the insects survived for weeks after.” On p. 235 there is another brief reference to this case, from which it appears that the observation was made on Melo- phagus ovinus and that flies that had been attacked were fed again on normal blood, which explains their survival for some time after. II. Shelter Adequate protection against adverse factors in the environment is essential for the welfare, of the individual and hence for the survival of the species. The problem of how to secure this is solved neatly by the Hippoboscidae and other obligate ectoparasites, which use the source of food also as a permanent abode, thus eliminating many of the vicissitudes of a free life. To be successful in this, how- ever, the ancestors of the louse-flies needed far-reaching structural and physiological adjustments or adaptations, which eventually led to such extreme specialization that their fate is now tied up ir- revocably with that of the hosts. It may be pertinent to consider the peculiar environmental conditions offered by the body cover of birds and mammals and the manner in which they affect the bionom- ics of the flies. Physical Characteristics of Feathers and Hair. From various considerations to be presented in Part II of this paper, it is believed that the earliest hosts of the ancestral Hippoboscidae were most probably birds, so that the peculiarities of plumage should be con- sidered first. Feathers are unique horny products of the epidermal cells of the avian skin, not duplicated elsewhere in the animal king- dom, and the most outstanding characteristic of the birds. Typi- cally a feather is a thickened and usually stiff main stem or quill, bearing an inner and an outer web or vane of softer barbs, which 116 Volume XXXII are divided into barbules and these in tnrn into barbicels. The several parts, as well as the whole, vary greatly in size and shape. The outer or surface feathers are the relatively strong and elastic contour feathers, some of which become very large on the wings and tail, being then used for flight. Elsewhere on the body the con- tour feathers overlie the inner or under down feathers, which are smaller, more fluffy and often more numerous. In most birds the feather coat or plumage is a thick and compact, though light and elastic mass of regularly imbricated contour feathers, slanting back- ward or downward, thus forming a smooth outward surface and a nearly waterproof cover for the under down. From our present viewpoint, the plumage encloses most of the body in a compara- tively thick layer of light, aerated and fairly waterproof material. Its elastic resilience guards the body against mechanical injury, while the blanket of air held in the interstices of the feathers insu- lates the skin, protecting it against rain and dampness and reducing the loss of heat by convection and radiation, particularly during flight. A microclimate of unusual uniformity for temperature and humidity prevails in the rather loose layers of feathers, as com- pared with that of the outside environment. In this respect bird plumage is clearly one of the most favorable biotopes, for insect life. It should be noted that, owing to the lack of sweat glands in the avian skin, the air enclosed within the plumage never becomes over- saturated with moisture by perspiration when the temperature in- creases unduly. Excess body heat is released mainly through many air-sacs distributed throughout the body and connected with the lungs (Wetmore, 1921, p. 23). In addition, the insulating blanket of air in the plumage can also be regulated; pulling the feathers closer removes some of the air and increases the loss of body heat, while puffing them out reduces the loss by admitting more air, a method of heat regulation affecting the ectoparasites directly. The inner body temperature of birds is known to average higher than that of mammals and to vary more among the several species as well as individually. It fluctuates also following changes in the bird’s activity and for other causes; there seems to be no marked seasonal variation. According to Wetmore (1921), resident birds of cold regions become more heavily feathered before winter sets in and more thinly feathered for summer, thus decreasing or increas- ing radiation of heat from the skin. The inner temperatures of birds were investigated and summarized by Wetmore (1921) and by Baldwin and Kendeigh (1932). A short paper by Becker and Stack (1944) gives valuable data taken from live birds of 37 corn- 117 ENTOMOLOGICA AMERICANA mon species from the central United States (30 passerines, 1 wild dove, 1 quail and 5 Picidae), some of which are among the most frequent hosts of two common American flies, Ornithomyia fringil- lina and Ornithoica vicina. The average inner body temperature of these birds varied from 108.1° F. to 110.2° F., the maxima from 108.3° F. to 115.8° F., and the minima from 104.3° F. to 109° F. In T urdus migratorius, which showed the greatest variation, the temperature ranged in 48 birds between 105.2° F. and 115.8° F., the average being 109.5° F. Wetmore (1921, pp. 18-21) also found that there is a marked difference in the temperature of nestling or young birds between the so-called precocial and altricial species. In the precocial birds, the young are fairly well feathered, able to run about and feed themselves as soon as they are hatched ; their body temperature ap- proximates closely that of the adult of the same species. The al- tricial birds, however, hatch in a weak and usually naked condition and require to be fed and protected in the nest for some time by the parents. Their nestlings show a lower average temperature than the adults. This was found to be the case for certain Columbidae ( Zenaidura ), Tyrannidae ( Myiarchus ), Hirundinidae ( Petrocheli - don), Muscicapidae (Geothlypis, Dumetella, and Sialia), and Pari- dae (Pentkestes) . The young of these birds are evidently depen- dent upon brooding by a parent to maintain their bodily heat. For instance, in the catbird, Dumetella carolinensis, the body tempera- ture of 6 nestlings averaged 101.7° F., while it was 108.6° F. for the male and 108.7° F. for the female. Since some of the common bird-flies often parasitize altricial birds and show a decided pref- erence for nestlings, the lower temperature of the young may be of some importance to them. The inner body temperature regulates that of the blood on which the flies feed ; but the surface body temperature controls the biotope of the insects. The skin temperature, derived from that of the body, is generally lower. It is less known than that of the body. Kallir (1931) determined the skin temperature of some 10 Euro- pean species, as well as the variations for several areas of the body. Baldwin and Kendeigh (1932) found that the skin temperature of the American house wren, Troglodytes aedon, is 0.8° F. to 3.9° F. lower than that of the body and differs in both sexes as well as in several parts of the body ; it is not affected by moderate changes in the temperature of the air, being on the whole less variable than the body heat. In 9 males, the inner body temperature averaged 108.5° F., while the skin temperature was 107.2° F. on the belly, 118 Volume XXXII 105.5° F. on the breast, 106.7° F. on the side, and 106.6° F. on the back. In 15 females, the body temperature also averaged 108.5° F. (as in the males); the averages of the skin temperatures were: 107.2° F. on the belly (as in the males), 107.2° F. on the breast (higher than in the males), 106.7° F. on the side (as in the males), and 105.6° F. on the back (lower than in the males). It seems rather donbtful that such slight variations conld seriously affect the presence, distribution over the body, or seasonal occurrence of hippoboscids on a bird host. Although pelt, the hair cover of mammals, differs greatly from plumage, it is equally effective as a protective device. In most mammals the skin is yuore or less clothed with hair, a modified form of epidermis exclusively found in this class of vertebrates. Hairs are of different length, thickness, rigidity or softness, density, direc- tion and color. Three main types, connected by transitional forms, may be distinguished (Noback, 1951). Guard hairs (“Leithaare” or “Borstenhaare” of Toldt, 1935) are fairly straight, evenly thick throughout or gradually thinner at the tip, and are sometimes modi- fied into bristles or spines. Pile hairs (“Grannenhaare” of Toldt) or awns are also straight, but relatively thin basally and thickened over the apical portion. These two types form the over- hair or upper fur, the major and coarser part of the' pelt of many mammals, particularly among the Ungulates. The third type, or fur hairs (“Wollhaare” or “Flaumhaare” of Toldt), are much softer and finer, often showing a tendency to curl. They form the underhair or under fur, beneath the cover of guard and pile hairs, and give the woolly feel of most fur animals. Variations in the relative proportions of the three types of hairs impart to each species of mammal its characteristic pelt. These differences may be of some importance to ectoparasites in selecting or avoiding avail- able hosts. The pelt guards the body and especially the skin against me- chanical injury, owing to the thickness, durability and resilience of the upper fur. It also protects against excessive atmospheric humidity, particularly fog, rain and snow, as well as against loss of body heat in a cold or windy environment. Insulation is achieved mainly by a series of thin layers of motionless air within the pelt (Herrington, 1951). The woolly under fur is a particularly effec- tive insulator and is therefore as a rule unusually developed in ani- mals apt to be exposed to extremely low temperatures. On the other hand, hair also helps cool off the body when it is exposed to excessive heat, inasmuch as the animal can erect the upper fur. 119 ENTOMOLOGICA AMERICANA This releases some of the enclosed air layers and increases the sur- face exposed to the evaporation of moisture secreted by the sweat glands, which are present in the skin of most mammals (except the Carnivora and Rodentia). On the whole, the texture and microclimate of the mammalian pelt seem less advantageous to ectoparasitic insect life than those of the avian plumage. Regardless of how they were acquired in the course of evolution, nearly all the peculiar external features of the majority of Hippoboscidae now function as near-perfect adap- tations to life within plumage rather than within pelt. The general dorso-ventral flattening and correlated modifications of the body, notably the greatly widened sternum, the laterally attached and usually long legs, the slender and often deeply split claws, the emarginate notum, the wedge-shaped head, and the small, concealed third, antennal segment, all combine to allow the fly to glide without hindrance or harm beneath the contour feathers, to move in any direction amid the down, and to maintain a firm hold while feeding or when the bird is moving. On the other hand, within the pelt of most mammals these structural peculiarities lose certain of their advantages. It is noteworthy that some of these features may be lacking in the mammalian louse-flies, possibly because they were never acquired by their ancestors ; others may have been merely retained from the ancestral types of bird-flies from which the louse- flies of mammals were derived, without being now of any particular advantage. Certain mammal-infesting flies are noticeably less flat- tened than those of birds and all of them are decidedly less stream- lined. However, the legs of the mammalian flies are shorter and often thicker, with a pair of heavier claws (sometimes unequal in length) than those of the bird-flies, enabling them to cling more securely to the coarse hairs of the rather loose pelt. It should also be pointed out that most mammalian pelts compare unfavorably with the average avian plumage as a protective cover against ad- verse climatic factors and potential enemies. The less effective hair cover may account to some extent for the paucity of louse-flies on mammals (34 species of 8 genera on mammals, as compared with 90 to 100 species of 13 genera on birds) and for the fact that their flies occur on fewer types of hosts than those of birds. Effect of Macroclimate. The foregoing considerations might convey the impression that the behavior and distribution of the bird-flies, at any rate, must be relatively little influenced by the usual climatic factors, or macroclimate, of the host’s environment. There is nonetheless evidence that these insects are by no means 120 Volume XXXII completely immune to them. Many bird-flies show very little host specificity and therefore tend to occur over wide areas, some species being nearly cosmopolitan (such as Lynchia albipennis and Orni- thoica confluent a of wading birds) or circumpolar or Holarctic (such as Ornithomyia fringillina and Olfersia fumipennis) . This feature may obscure the effect of climatic conditions. Yet it is noteworthy that Ornithomyia fringillina and Lynchia americana, two of the most common bird-flies of temperate North America on a variety of hosts, are practically unknown in the Neotropics. Since they must be carried occasionally to tropical America by some of their migratory hosts, failure to become permanently established there can only be due to some adverse macroclimatic factor. On the other hand, like most of the Hippoboscidae, the majority of the American bird-flies are essentially tropical and subtropical insects, only some of them spreading normally or accidentally to more tem- perate or colder areas, particularly along the eastern Atlantic sea^ board. Such species as Ornithoica vicina , Ornithoctona erythro- cephala, and Ornithoctona fusciventris, which reach southeastern Canada, are scarcely permanent members of the local fauna in the northernmost part of their range. They should be regarded there rather as seasonal migrants, the adults and possibly even the pu- paria being normally absent during the winter. In these northern areas, these flies are most probably introduced anew in the spring by the migratory hosts returning from the south, at least after severe winters. A low average yearly temperature and extreme cold during the protracted winter are the two main factors limiting the northward extension of the Hippoboscidae. The normal limit for the family in the New World is at about the 50th parallel, north of which the bird-flies at any rate are scarcely part of the autochthonous fauna. Among many hundreds of American bird-flies examined in recent years, only half a dozen Ornithomyia fringillina were from Alaska at the following localities: Crater Mt., on a ‘‘Columbia falcon”; Nelchina River, north of Mt. Witherspoon, N. W. of Valdez; Ta- kotna, 63° N., 156° W., on Canachites c. canadensis. Old Crow River, Timber Creek, Yukon, about 68° N., 140° W., at present the northernmost New World locality for an avian hippoboscid, is near the annual isotherm of 20° F. In the eastern part of the continent, where the flies stay much farther south, the northernmost record is for an Ornithomyia fringillina (recorded as avicularia) taken by Eidmann (1935, p. 99) from Junco h. hyemalis on the Matamek River, in the southern Labrador Peninsula (15° 17' N.) ; this local- 121 ENTOMOLOGICA AMERICANA ity is also near the annual isotherm of 30° F. As some of the pas- serine birds known in North America as hosts of 0. f ring illina migrate in the summer to the Arctic Circle or even beyond the scar- city of hippoboscids in the Far North must be due to the climate being unsuitable even in summer. No louse-fly has ever been taken in Greenland. The Hippoboscidae of birds extend farther north in Europe than in America, which agrees with the more northern course of the annual isotherm of 30° F. in Scandinavia, where it approaches or even exceeds the 72nd parallel. The Holarctic Ornithomyia fringillina is known from Iceland, Scotland, the Shetlands, northern Scandinavia and northern Finland. I have seen this fly from Levajak on the Tana River, Finland, between 69° 45' and 70° N., the northernmost locality known for any hippoboscid. The other species of northern Europe, Ornithomyia avicularia , Stenepteryx hirundinis, Crataerina pallida , and Olfersia fumipennis, do not occur north of the Shetlands, southern Scandinavia and southern Finland, where the climate is about that of southern Canada and the northeastern United States. The effects of climatic variations in the relative humidity of the air are more subtle than those of the temperature and therefore more difficult to discover. They should not be discounted, however, since it has been shown that certain Hippoboscidae are averse to excessive moisture, as I pointed out in discussing various tropisms of the flies. Mr. H. Hoogstraal, who has had much experience with the several species of Hippobosca infesting domestic animals in Egypt, informed me recently that these parasites are relatively scarce in the Nile Valley proper and the Delta, but increase in num- bers as one travels toward the drier or desertic areas. Macroclimatic factors also might possibly account to some ex- tent for the periodic fluctuations in successive years of the hippo- boscid population in the same locality. Herman (1937, p. 164) noted that at North Eastham on Cape Cod, Massachusetts, Orni- thomyia fringillina and Ornithoica vicina were much more numer- ous on trapped passerine birds during August, 1937, than during August, 1936. According to information he received from Mr. C. L. Whittle, the reverse was true for these two years at Peterboro, New Hampshire. In 1937 there was practically no rain during July on Cape Cod, while in New Hampshire there was more rain in July, 1937, than the previous year. Herman suggests that per- haps hippoboscid flies thrive better in a drier atmosphere ; but it might be difficult in this case to dissociate the temperature factor 122 Volume XXXII from the atmospheric humidity, since in the northeastern United States summers with a heavier rainfall are . generally cooler than those of drier years. It is known that the temperature influences the rate of reproduction of the louse-flies, the puparia requiring more time to produce flies in cool weather. Pseudolynchia canariensis is thus far the only bird-fly whose range has been affected appreciably by man. At first mainly a parasite of wild Columbidae and diurnal birds of prey in the Old World tropics, whence it occasionally strays to Central Europe, it has become a regular parasite of domestic pigeons, on which it was carried by man at one time or other no doubt to nearly every part of the Earth. Yet it is far from being as cosmopolitan as its do- mesticated host. It failed to become established in Europe north of the 45th parallel, while in America it is rarely seen north of the 40th parallel. The few sporadic captures in pigeon lofts or on feral pigeons in New York and Boston are exceptional and prob- ably seasonal. There is no evidence that either the adult or the puparium could survive a normal winter in the northeastern United States (J. Bequaert, 19435) . No doubt owing to the poorer protection provided by hair pelt, the louse-flies of wild mammals are more sensitive to the macro- climate than those of birds. Their range is always more restricted, although this may be due in part to the more sedentary habits of the hosts. Most species are strictly tropical and the few that ven- ture into colder temperate areas seldom extend as far north as their potential breeding hosts ( Lipoptena cervi in Europe and Asia to 60° N. ; L. depressa in northwestern America to 50° N. ; L. mazamae, a tropical species, only to about 34° N. in the eastern United States). Seven species of hippoboscids, Melopliagus ovinus, Lipoptena capreoli , Hippobosca equina, H. longipennis, H. rufipes, H. came- lina, and H. variegata, now use domesticated mammals as normal breeding hosts. Although some of these flies have been spread un- intentionally by Man over wide areas, none has been able to be- come established over the same territory as its host and none is ac- tually cosmopolitan. The sheep-ked, M. ovinus, the most success- ful of them, nowadays has only the domestic sheep as a normal breeding host; but the common assumption, often repeated in text- books, that sheep carry keds wherever they are kept, is erroneous, as shown by an analysis of the known distribution (J. Bequaert, 1942a, pp. 183-186). Keds are either entirely lacking or occur only as temporary introductions in many areas where sheep are regularly kept. They avoid particularly the hot tropical and sub- 123 ENTOMOLOGICA AMERICANA tropical lowlands, as well as many oceanic islands. The climatic factor which eliminates the ked after its introduction in unsuitable territory appears to be the high air temperature. It has been ob- served that at higher ambient temperature sheep-keds move away from the skin to the outer wool (Swingle, 1913; Hoffmann, Roth and Lindquist, 1950; Evans, 1950, p. 467). M. ovinus is the hip- poboscid which reaches farthest north, as it extends close to the Arctic Circle in Alaska, Iceland, the Faroes and Lapland. In the Southern Hemisphere it occurs on the Falkland Islands (52° S.). Lipoptena capreoli, a specific parasite of the domestic goat, is at present restricted to the eastern Mediterranean countries and north- western India, where snbarid conditions prevail. It shows no tendency to spread outside this area with its host. Hippobosca equina, the cattle and horse louse-fly, not known at present from a wild host, is a common parasite of equines and cattle in many trop- ical and subtropical parts of the Old World, extending northward to a few disconnected areas as far as southern England and southern Sweden. It is, however, by no means uniformly distributed over this territory, since it occurs only as exceptional and temporary introductions in Africa south of the Tropic of Cancer ( J. Bequaert, 1931, pp. 310-311)/ Even more remarkable is its failure to become established in the New World, although it must have been intro- duced there repeatedly with horses. Hippobosca longipennis (= H. capensis), the dog-fly, is fairly common on a variety of wild Car- nivora in East Africa ; it is chiefly an abundant parasite of domes- tic dogs in many Mediterranean countries and southern Asia, ex- tending northward as far as northern China. In Africa it avoids the moister tropical areas, being unknown from most of West Africa and the Congo Basin. It is also not found in Indonesia, Australia and the New World. It could scarcely have failed to reach America from time to time, yet has never become established there. Hippo- bosca variegata (= H. 'maculat'd ), another horse and cattle fly, with- out known true wild host, is very abundant over the northern half of Africa, India and Indonesia ; but again, it has never been able to gain a foothold in the New World. The climatic or possibly other factors which restrict the spread of these abundant parasites of cos- mopolitan domestic hosts should be investigated more carefully. Preferential Distribution on the Host. Many ectoparasites tend to select certain areas of the host ’s body in preference to others. This is particularly true of the biting and sucking lice, where two or more species living on the same individual host may each be re- stricted to its own area of the plumage or pelt, thus avoiding corn- 124 Volume XXXII petition. Little such topical selectivity is exhibited by the Hippo- boscidae, presumably due to their great mobility, the generally light individual infestations and the scarcity of infestations with more than one species of fly. On birds, the flies are, moreover, difficult to locate, being usually concealed on undisturbed hosts, while they either leave or scatter over the body if the bird is handled or other- wise disturbed. There should be, however, a certain amount of topical prefer- ence, conditioned by such factors as the feeding facilities, differences in the temperature and moisture of the skin, and variations in the possibility of escaping the destructive preening and cleaning efforts of the host. As pointed out before in discussing the sense organs, some louse-flies are sensitive even to relatively slight changes, par- ticularly in the temperature. Unfortunately, little information is now available on the surface temperature of mammals and birds in different parts of the body. That such topical differences may be substantial is shown by Kallir’s (1931) study of the European turtle dove, Streptopelia turtur, in which variations up to 3° C. were observed. The lowest temperatures (38° to 39° C.) were in the plumage of head and croup ; the highest on the sides of the rump and near the anus (over 41° C.) as well as on the back and beneath the wings (nearly 41° C.). The topical variations of the skin tem- perature of Troglodytes aedon were mentioned in the general dis- cussion of the body temperature of birds. The uneven distribution of Melophagus ovinus over the sheep’s body has often been noted. It was most carefully studied by MacLeod (1948), in England. He found that, in winter and early spring, the upper side wool is equally if not more favored by the keds than the lower side wool, and that an appreciable concentration occurs on the back. With the advancing warm season, the keds tend to forsake the upper surfaces. The lower wool, including the abdomen, a site little favored in winter, now becomes the principal territory. The chest and throat are favored throughout the year, but particularly in winter and spring. Evans (1950, pp. 470-472), also in England, offers further statistical evidence in support of these conclusions and emphasizes the periodic migrations of the keds on the sheep ’s body. Cowan (1943, pp. 183 and 185) studied the distribution of Lipoptena depressa over the body of coast deer, Odocoileus Jiemi- onus columbianus, in British Columbia. He states that there is a strong tendency for the gravid keds and their clasping mates to congregate on the lower surface of the host from the base of the 125 ENTOMOLOGICA AMERICANA neck to the abdomen. On hot summer days, large numbers of flies frequently move about over the outer pelt. One deer in mid- August showed many crawling over the face and legs in nature. Hippobosca equina and H. variegata, of cattle and equines, often cluster in numbers under the tail and between the thighs. Gachet (1915) reports that the dog-fly, Hippobosca longipennis (=H. canina) , in Iran, frequently was observed biting sleeping dogs on the nostrils and at the edges of sores. Speiser (1907, p. 5) stated that East African specimens of H. struthionis were said by the col- lector to occur mostly beneath the ostriches ’ wings ; it may be that the unusually heavy plumage made the detection of the flies difficult elsewhere. There are as yet few observations on this topic for the bird-flies. They suggest that the flies seek out areas of the body where there is a better chance of escaping the destructive pecking endeavors of the host. For instance, Olfersia fossulata, the cQmmon parasite of the guanay, Phalacrocorax bougainvill'ei, on the coast of Peru, often selects the bird’s head and neck, as shown on a photograph by Dr. R. C. Murphy (1924, National Geographic Magazine , 46, p. 294). The photograph by Dr. J. P. Chapin (Fig. 21) of Olfersia aenescens infesting the red-tailed tropic bird, Phaeton rubricauda, on Ducie Island, shows a similar preference of the flies for the neck and upper breast. It is also interesting to note that several specimens of Or- nithoica vicina, which had strayed onto young domestic fowl in Virginia, were all on the head. Biittiker (1944), in Switzerland, observed that Crataerina pallida stays mainly on the neck and head of the swift, Apus apus. Protective Behavior. Even the fully-winged species of Hip- poboscidae rely mainly on running and hiding in order to escape danger while on the host. The wings seem to be used more to reach a host after emergence, in some cases at mating time, and when the flies are forced to leave a dead host and seek a new one. While winged louse-flies are not among the swiftest or most skillful fliers, they are nevertheless not as slow and clumsy on the wing as might be supposed. They take off and alight rather quickly, but their flights are always jerky and short. Hippobosca equina, for in- stance, which has rather powerful wings, flies when disturbed a distance of \ to 1 meter or less and then alights suddenly. Hase (1927, p. 193) observed also that it rises sometimes from the host for a brief, sharp turn in the air, to alight again close to the spot it had left on the skin. Roberts (1925) observed an unusual behavior of Hippobosca 126 Volume XXXII equina in North Wales, England, where it is very common and well established in the open in certain areas. In the summer the fly is abundant on both horses and cattle on bright, sunny days, but much rarer on cold and rainy days. Its distribution is closely correlated with the dense covering of bracken fern ( Pteridium aquilinum) on certain hill slopes. During the day flies are seen occasionally sun- ning on the fern fronds ; at sunset or when a spell of cold weather or rain is imminent, they generally leave the host and seek shelter on the under side of the fronds where they spend the night. Whether fully-winged or subapterous, most Hippoboscidae run swiftly and skilfully through the pelt or plumage of the host. The long legs inserted at the extreme sides of the flattened thorax, and the unusually developed mid and hind trochanters allow them to move without having to raise the body, with equal ease and rapidity in every conceivable direction, forward and backward as well as sideways, as Dufour (1845) first pointed out for Hippobosca and Ornithomyia. The running motions of Hippobosca equina were studied by Hase (1927, pp. 194-196), who states that it may cover 3 to 7 cm. per second, its movements amid the hairs being greatly helped by the flattened body and particularly by the wedge-shaped head. Many of the bird-flies run even faster than Hippo~bosca, especially the subapterous species, which must rely on running alone for their safety and for reaching a host. The swift-fly, Cra- taerina pallida, studied by Kemper (1951, pp. 249-251), is ex- tremely agile; it does not travel far in one stretch, usually a dis- tance of 5 to 10 cm., sometimes only 2 to 5 cm., exceptionally 31 cm. In unfed condition, it covers about 5 cm. per second (calculated from 6 flies) and slightly over 4 cm. per second when fully engorged (calculated from 4, flies). Eichler (1939, p. 215) claimed that C. pallida could in addition perform small jumps ; but Kemper was unable to confirm this and thinks that the jerky motions of the fly may have simulated jumps. Biittiker (1944, p. 28) observed on one occasion that C. pallida used the reduced wings to glide a short dis- tance in the air. Schuurmans Stekhoven and his co-workers, at Tucuman, Argentina, determined the speed of Pseudolynchia canariensis by allowing flies to run in a glass tube 75 cm. long and 11 mm. wide (unpublished observations). As a rule this fly does not run very far before stopping for a few seconds : one fly covered the 75 cm. in four stretches or “heats,” at speeds of 108, 300, 330 and 166 cm. per minute for each successive heat. The highest speed observed was 540 cm. per minute. This fly often moves the wings also while on the run. 127 ENTOMOLOGICA AMERICANA The Melophaginae, or ked-flies of certain mammals, are excep- tional in the family for their rather slow movements. Hase (1939, p. 412), in Germany, noted that winged, unfed Lipoptena cervi, of European deer, are rather slow runners, usually covering 0.5 to 1 cm. per second, rarely 2 cm., on an open surface, away from a host and unhampered by hairs. The apterous Melophagus ovinus man- ages to move about a great deal on the host, in spite of the short and thick legs ; but its gait is very slow and clumsy. According to Freund and Stolz (1928), it covers from 6 to 12 cm. per minute, with an average of 8.5 cm., as compared with the 240 to 300 cm. per minute of Crataerina pallida. Kemper (1951, p. 251) noted that a crawling M. ovinus makes rapid grasping motions in the air with the fore legs before putting them down, no doubt in attempts to grab a host. In Allobosca crassipes of the lemurs, in Madagascar, the legs are even shorter and much more swollen than in Melophagus. Mr. H. Hoogstraal, who observed this insect alive on the hosts, informs me that it is slow moving and secretive, keeping in the pelt near the roots of the hairs. Most Hippoboscidae exhibit a strong positive thigmotropism (or stereotropism) and flatten the depressed thorax and abdomen against the host’s body or any other surface. Even a decapitated Hippobosca equina, placed on its back, turns over at once so that the sternum and venter touch the support, after which it remains motionless on the spot, the fly being evidently warned of its normal or abnormal position by the many tactile setae (Hase, 1927, p. 202). H. A. Macpherson (quoted by Austen, 1926, p. 254) noted that, if a Crataerina pallida is surprised in a swift ’s nest away from a bird, it often flattens the body against the background and remains mo- tionless. Cowan (1943, p. 185), in British Columbia, found that the tropistic responses of Lipoptena depressa change with age. Newly- emerged, volant or recently dealated, unfed flies of both sexes show a strong negative geotropism, always moving upward, either in a tube or on any object within reach. They are also positively photo- tropic, moving toward the light, but this tropism is secondary to the negative geotropism. Winged individuals seem to prefer light to dark clothing when alighting on a person. On the other hand, dealated flies that have fed for some time tend particularly to cling to any small object or to each other when removed from the host, a positive thigmotropism not noticed in the newly-emerged indi- viduals. If several L. depressa of different ages are placed in a vial, the older individuals cling together in a mass at the bottom of '128 Volume XXXII the vial, while the younger keds climb actively, each by itself. Prouty and Coatney (1934, p. 252) found Pseudolynchia canari- evisis (= P. maura) positively thigmotropic, seeking to push its way into small cracks or under the edge of pads or sheets of paper ; this may explain why the female prefers to larviposit in rather tight cracks. According to Kemper (1951, p. 252), Crataerina pallida also turns over immediately to a normal position when placed on its back. Melophagus ovinus, however, is unable to turn over to the right position, unless its legs can grasp a near-by object (Freund and Stolz, 1928; Kemper, 1951). The peculiar tropistic responses of the louse-flies no doubt help them in escaping their enemies. They certainly make their capture unusually difficult. Predacious Enemies. It is generally recognized that the hosts themselves are by far the most dangerous enemies of all ectopara- sites. This important fact should not be overlooked when studying the effects of external factors or changes in diet upon populations of such arthropods. Any circumstance that either enhances or impairs the host’s vitality and efficiency in combating its ectopara- sites, tends to decrease or increase the parasitic population. Irri- tated by the bites or by the movements of the parasites, the host attempts to destroy them by every possible means, such as preening, rubbing, scratching or catching in mouth or bill, or even devouring them. It is not unthinkable that certain peculiar activities of birds, such as bathing in water or dust, smoke-flights and 1 ‘ anting, ’ ’ are more or less deliberate attempts to get rid of annoying parasites (Rothschild and Clay, 1952, pp. 125-128). It is improbable, how- ever, that such methods are very effective against the swift Hippo- boscidae. Ed. and Et. Sergent (1906), Drake and Jones (1930), and Coat- ney (1930) noted that domestic pigeons fight Pseudolynchia canari- ensis by pecking and thrusting the bill in the plumage, thus destroy- ing many flies. Austen (1921, p. 122) observed: “It was found in Palestine that an infested pigeon on returning from a flight, in- stead of at once seeking its resting place, as these birds usually do, would alight on the floor of the loft and proceed to stamp and peck itself. ’ ’ Grimshaw (1911, 2, p. 86), while examining the contents of the crops and gizzards of red grouse, Lagopus scoticus, in England, found grouse-flies, Ornithomyia fringillina (= 0. lagopodis) , in 3 young birds, 28 to 40 days old: one fly in the gizzard of one bird and 2 flies in that of another ; and one fly in the crop of the third bird. 129 ENTOMOLOGICA AMERICANA Herman (1944, p. 114), working with Stilbometopa impressa of California qnail, Lophortyx calif ornica vallicola, observed that sev- eral flies placed in a cage with one or 2 quail yielded many puparia, while an equal number placed with 7 quail produced very few or no offspring. The birds were seen eating flies that leave the host temporarily when it becomes excited, as flies exposed in the open have little chance of escaping capture when many birds are con- fined in a small space. When 25 flies were placed in an escape- proof cage with 7 quail, 24 hours later only one fly could be found after extensive examination; but 17 flies were recovered 3 days after 18 flies had been placed in a cage with 2 quail only. Mr. I. B. Tarshis also noted that California quail eat their flies and found in addition that the birds catch the smaller Lynchia hirsuta more easily than the larger Stilbometopa impressa (unpublished obser- vations). Some years ago, Dr. C. M. Herman made the following interest- ing unpublished observation on a chipping sparrow, Spizella pusilla passerina, on Cape Cod, Massachusetts : ‘ ‘ This bird, originally with 3 specimens of Ornithoica vicina and heavily infected with Haemo- proteus, was placed in a gauze-covered cage about one foot on each side. During a period of one week 117 specimens of 0. vicina, taken from various hosts, were placed in the cage with this sparrow. At the end of this period the bird was removed and examined. Only 2 flies were found. Further flies were then planted in the cage with the bird and kept under close observation. The flies would periodically leave the bird to rest on the gauze wall and then re- turn to the host. Every so often, en route, they would become a cherished part of the bird ’s diet. ’ ’ Sheep eat Melophagus ovinus regularly and in large numbers, this being the only known way in which they become infected with a blood protozoan, Trypanosoma melophagium, part of whose life cycle necessarily occurs in the ked (Hoare, 1923; Turner and Murnane, 1930). Newman and Cordeaux (1871), in England, noted that sheep “ pluck themselves of keds.” Evans (1950, p. 467), also in England, saw on several occasions “sheep biting keds from the surface of the fleece and then devouring them. This procedure was very marked during warm weather when some of them were on the surface of the fleece, and after shearing when the sheep were able to pick off keds without much difficulty. During colder weather, when the insects were in the depth of the fleece, the sheep had to be content with biting the fleece in an attempt to rid itself of the irritation.” 130 Volume XXXII Theodor (1928, p. 316), in Palestine, observed goats removing goat-keds, Lipoptena capreoli (=L. caprina), from the hairs with the lips and eating them. As in the case of Trypanosoma melo- phagium, the goat thus becomes infected with a similar protozoan, Trypanosoma theodori, which goes through part of its life cycle in the Lipoptena. Modeer (1785), Girard, (1885), Neumann (1888), Evans (1950, p. 467), and others report that various birds, such as crows, star- lings, wagtails and magpies, will catch and eat keds on sheep. Un- fortunately there is no confirmation of this habit by actually find- ing keds or their puparia in the stomach of any of the birds. Dixon (1944) mentions that the California jay, Aphelocoma calif ornica, was seen picking “ ticks and deer tick flies” from the back of mule deer in Sequoia National Park, California. This state- ment again needs confirmation by examination of stomach contents. Melin (1923, p. 21) saw an asilid fly, Dioctria hyalipennis (Fabricius), which had captured and was sucking an Ornithomyia sp. in Sweden. Freund and Stolz (1928, p. 105) mention incidentally that, in Timor, Indonesia, they were told of a large hornet, Vespa auraria nigrithorax , eating the louse-flies of horses. Schuurmans Stek- hoven himself did not observe any insect predator of Hippobosca in Indonesia; he was informed (1926, p. 47) that another person on the island of Soemba witnessed a fossorial wasp capturing Hippo- bosca variegata. These two statements should be confirmed by more careful observations, since they may be due to some confusion. The wasps may have been catching horse-flies (Tabanidae), which are the usual prey of certain species of fossores and more easily caught than Hippobosca. According to Dr. W. W. Wirth (in lift., 1946), on Rabbit Island, off the coast of Oahu, Hawaiian Islands, spiders were seen catching Olfersia aenescens resting on rocks in the rookeries of noddy terns and shearwaters. Parasites. Compared with the free-living arthropods, most permanent ectoparasites of vertebrates are unusually free of para- sites of their own, presumably owing to the more effective protec- tion afforded by the host’s body cover. In the Hippoboscidae, in addition, the more vulnerable puparia are few, widely scattered, and usually hidden either in the pelt or plumage or away from the host. It is doubtful that any of the parasitic organisms discussed below reduce appreciably the total population of a louse-fly species. In view of the very low rate of reproduction, few losses are needed 131 ENTOMOLOGICA AMERICANA to maintain the population in a state of equilibrium and these are mostly caused by accidental injuries and by the predacious activity of the hosts. 1. Bacteria and Bacteria-Like Organisms. Under this head- ing will be grouped a variety of micro-organisms, including true bacteria, so-called bacteroids, true Rickettsiae, and Rickettsia- like organisms. They are conveniently studied together, particularly since it is often difficult to draw the line among them. Moreover, certain authorities regard even the Rickettsiae as merely a pecu- liar type of bacteria (Family Rickettsiaceae) . At first sight, it might seem important or at least useful to distinguish between extracellular and intracellular organisms. This again does not always prove feasible in practice. No doubt some of the micro- organisms observed in insects never enter tissue cells, while others live normally inside cells ; but some seem to have both an intra- and an extracellular stage. I am satisfied that the forms men- tioned in the sequel are true organisms and not waste or other metabolic, inert products of the tissues. Several of them grow and multiply on appropriate media or in tissue cultures outside the insect’s body, and in cases where cultivation has failed thus far, the organisms are similar to those that can be maintained in culture. A previous discussion of the blood diet of the Hippoboscidae considered at some length many of the micro-organisms reported from these insects, so that only a summary, with more complete bibliographic references, need be given here. I commented at that time upon the scarcity of such organisms and particularly of the extracellular types in these flies. Cory neb act erium lipoptenae Steinhaus (1943; 1946, p. 44, fig. 15, and p. 245), described as a true bacterium of the Family Mycobacteriaceae, was found in Montana in the deer-ked, Lipop- tena depressa, chiefly in the intestinal tract. It was successfully cultivated first on egg fluids and later also on other media. C. lipoptenae should be compared with various micro-organisms reported from Lipoptena cervi and L. capreoli in the Old World; perhaps also with Cory nek act erium ovis , known from domestic sheep and reported also from mule7 deer in British Columbia (Humphreys and Gibbons, 1942). Bacillus anthracis Cohn, the bacterium causing anthrax, is stated by Zumpt (1939, p. 704) to have been found in the gut of sheep-keds, Melophagus ovinus, taken on sheep which had died from anthrax. He also cites presumably unpublished experiments 132 Volume XXXII by Bongert, in which anthrax is claimed to have been transmitted from infected to healthy sheep by the bite of keds. It is partic- ularly in South Africa, where Hippobosca is sometimes unusually abundant on domestic animals, that* investigators have been inter- ested in the role such flies might play in anthrax epidemics. This possibility was investigated carefully by Viljoen, Curson and Fourie (1928). They state that Hippobosca rufipes is very common in South Africa, often appearing in the thousands on horses and mules, while cattle run on the same farm are relatively free from it. They note that anthrax is generally confined to equines, even where susceptible cattle and sheep freely mingle with them. The anthrax swellings were observed to start on the places where the flies cluster and to extend from there to adjoining parts. Bacillus anthracis was actually isolated from H. rufipes taken on horses infected with the disease. The epizootic form of anthrax occurs only during the summer, when louse-flies are particularly numerous on horses. It would seem therefore that H. rufipes is an important factor in the mechanical transmission of the disease in South Africa. Of course, there are several other ways in which anthrax is spread from diseased to healthy animals. True Rickettsiae and similar organisms are common in the sheep-ked, Melophagus ovinus, in the lumen as well as in the epithelial cells of the mid-gut, and also in the milk-glands of the fe- male. They were briefly mentioned in the discussion of the blood diet of the flies. Rickettsia melophagi Noller (1917 b, p. 70) was described and named from extracellular forms abundant nearly everywhere in all keds of both sexes* forming a layer over the cuticula of the epithelium in the lumen of most of the mid-gut. Notwithstanding unsubstantiated claims by Noller and Kuchling (1923), Gordon (1933), and Katie (1940), it seems fairly certain that the organism is not inoculated to sheep by the ked and that it is not pathogenic either to the insect or to vertebrates. Although the transmission from ked to ked is not completely elucidated, it is most probably acquired by the larva feeding on the infected secre- tion of the female’s milk-glands.11 It has been cultivated outside keds on certain media. The intracellular Rickettsia- like micro- 11 Jungmann (1918) claimed that he found the “ova” of M. ovinus infected with R. melophagi. His “ova” were not the true eggs (in the ovaries), but larvae developing in the fly’s uterus, a confusion commonly made by non-entomologists unaware of the peculiar reproduction of the Hippoboscidae. 133 ENTOMOLOGICA AMERICANA organisms of M. ovinus have not been given a specific name and it is as yet a moot question whether or not they are more than devel- opmental forms of R. melophagi. They are usually regarded as mutualistic symbionts and are -of two types. One form, apparently of general occurrence, fills the protoplasm of enlarged, specialized cells of a ring-like mycetome in the posterior portion of the mid-gut. A second form, different in shape, was found by Zacharias (1928) in some keds only, filling ordinary, flat epithelial cells usually of the mid-gut, rarely of the hind-gut or of the Malpighian tubules. Both types are harmless to the insect and are not transmitted to the sheep. According to Zacharias, some of these organisms occur freely in the lumen of the milk-glands of the female, whence they pass with the secretion of the glands to the alimentary tract of the intra-uterine larva. Here they enter specialized epithelial cells in the anterior portion of the mid-gut, close to the oesophagus. Zacharias describes also the complicated mechanism by means of which the symbionts reach the final mycetome of the adult during nymphosis. The micro-organisms of the sheep-ked are favorite study objects, so that the literature dealing with them is volumin- ous, as shown by the following references: Noller (19175), Jung- mann (1918), Sikora (1918), Hindle (1921), Arkwright and Bacot (1922), Arkwright (1923), Noller and Kuchling (1923), Kuchling (1924), Hertig and Wolbach (1924), van Thiel (1925; 1926), Anigstein (1927), Zacharias (1928), Glaser (1930), Buch- ner (1930), Aschner (1931), Kligler and Aschner (1931), Katie (1940), and Steinhaus (1946, pp. 244—245 and 312-315; see also pp. 205-206 for Kligler and Aschner’s method of cultivating the micro- organisms ) . The extracellular and possibly symbiotic intracellular bacter- oids of Lipoptena, Hippobosca, Ornithomyia, Stilbometopa, and Pseudolynchia were discussed before in the section dealing with their possible role in digestion. It was stated that intracellular symbionts, located in mycetomes of the wall of the mid-gut, probably occur in all Hippoboscidae. Whether this is true of the extracel- lular forms also is more doubtful ; and there is a possibility that the extra- and the intracellular forms in the same fly may be stages of one and the same organism. Both the bacteria-like and the Rickettsia- like forms vary in size and shape : R. melophagi may be spheroidal or coccoid, 0.4 to 0.6 /* in diameter on the average ; or more elongate and rod-like, 0.5 to 0.6 fi in length and 0.3 to 0.4 y in width. The intracellular symbionts are often very long and thin, thread-like, even simulating spirochetes. Spirochaeta melophagi 134 Volume XXXII Porter (1910), described from the gut, ovaries and puparia of Melophagus ovinus, in England, was perhaps based on bacteroids escaped from ruptured mycetocytes. No spirochetes have been reported since by the several investigators who studied the micro- organisms of the sheep-ked. 2. Protozoa. The relatively few types of Protozoa known at present from the Hippoboscidae belong to the two classes Mastig- ophora (or flagellates) and Sporozoa. Flagellates of the Family Trypanosomidae are very common in insects, some species being restricted to invertebrates, while others have both a vertebrate and an invertebrate host. In the latter case, the organism has at least a stage in the blood of the verte- brate, being often called a hemoflagellate ; while the invertebrate host is invariably blood-sucking and acquires the flagellate by feeding on the infected vertebrate. At least part of the life cycle of the flagellate occurs in the invertebrate’s intestinal tract. In most cases the vertebrate must acquire the flagellate from the in- vertebrate host, which usually, but not always, inoculates the protozoan with the mouth-parts while feeding. Both the verte- brate and the invertebrate are therefore essential links in the per- petuation of the hemoflagellate. In the Hippoboscidae, hemoflagellates are known at present only from the keds, or mammalian flies of the subfamily Melopha- ginae. The best-known is Trypanosoma melophagium (Flu) (= Crithidia melophagia Flu, 1908), originally regarded as a specific flagellate of the ked, where it was first seen and described by E. Pfeiffer (1905) in the lumen of the mid-gut, in Germany. It is now recognized that the forms in the ked are developmental stages of a trypanosome found in the blood of most domestic sheep, for which it is not pathogenic. Laboratory-bred, clean keds have been infected experimentally by feeding them on the blood of sheep harboring trypanosomes. The flagellate upon reaching the ali- mentary tract undergoes a definite cycle of development : a so-called crithidial stage is produced in the mid-gut, followed by forms in- fective for sheep (metacyclic trypanosomes), which line the walls of the hind-gut. According to experiments by Hoare (1921; 1922; 1923a; 19235) and Turner and Murnane (1930), infected keds do not inoculate the trypanosome by the bite (proboscis), but by the contaminative method, by way of the sheep ’s mouth. Sheep become infected by biting keds off the fleece and crushing them in the mouth, the metacyclic trypanosomes presumably penetrating through the buccal mucous membrane. In all experiments in 135 ENTOMOLOGICA AMERICANA which emulsions of infected keds were fed by month to clean sheep, the latter acquired the trypanosome; while control experiments to infect clean sheep either through the abraded skin or by the bite of infected keds, invariably failed. In addition to the authors cited above, T. melophagium was also studied by Swingle (1909; 1911a; 19115), Georgewitch (1910), Annie Porter (1910), C. F. Bishop (1911), Cauchemez (1912), Chatton and Delanoe (1912), Dunkerly (1913), Laveran and Franchini (1914a; 19145; 1919a; 19195), Noller (1917; 1919), Kleine (1919), Boning (1920) , Witzky (1922), Sprehn (1922), Buchner (1922), E. R. Becker (1923), Boese (1923), Noller and Kuchling (1923), Kuchling (1924), Bozhenko and Tzeiss (1928), Colas-Belcour (1931), and Wotton (1940). A well-illustrated account of the life cycle of this parasite is given by Wenyon (1926, 1, pp. 502-507). Bishop (1911) claimed that he saw crithidial forms of T. melophagium in a tick of sheep, Ixodes ricinus, in England ; this finding has not been confirmed. Bouvier et al. (1951, p. 276) mention briefly that H. Gaschen found Melo- phagus rupicaprinus of the chamois, Bupicapra rupicapra, in Switzerland, infected with “ Herpetomonas melophagae” ; these may have been developmental stages of a trypanosome of the chamois. A hemoflagellate, first seen by Adler (1926) in the digestive tract of the goat-ked, Lipoptena capreoli (= L. caprina ), in Palestine, and also studied by Theodor (1928), was later called Trypanosoma ikeodori by Hoare (1931). Theodor recognized that it is a stage in the invertebrate host of a non-pathogenic trypanosome found in the blood of domestic goats. The developmental cycle is similar to that of T. melophagium. Newly-emerged L. capreoli are never infected. If kept on an infected goat, the ked acquires the trypano- some in from 5 to 22 days. The scarcity of the trypanosomes in the blood of infected goats explains the slowness and the low rate of infection of the keds. In a herd of 15 goats, presumably all infected, on 2 animals all keds were negative, on 11, from 6 to 70 per cent of the keds were infected, and only on 2 were all keds infected. In the ked, the flagellate occurs in the mid- and hind-gut, never in the fore-gut, proboscis, salivary glands or other organs. Theodor established that reinfection of the goat is not by active inoculation through the bite or by the feces of the ked, but by infected keds being crushed by the goat. Even emulsions of in- fected keds injected subcutaneously failed to inoculate the trypano- some. Herman (1945, p. 21) reported an unidentified flagellate from 136 Volume XXXII the intestinal tract of Stilbometopa impressa from the valley quail, Lophortyx calif ornica vallicola, in California; and Mr. I. B. Tar- shis (in -lift., 1952) observed a similar organism in the same fly. The affinities of this flagellate have not been elucidated. It is not impossible that the organism may be a developmental stage of the: trypanosome which Herman (loc. cit.) observed in the blood of the valley quail. In the course of his experiments with pigeon malaria, Aragao (1927), in Brazil, found domestic pigeons sometimes in- fected with Trypanosoma hannae Pittaluga (1905, Rev. R. Acad. Cienc. Ex. Fis. Nat. Madrid , 2, No. 3, p. 356). He observed that 12 Pseudolynchia canariensis (= Lynchia maura), bred from puparia and fed on a pigeon infected with the trypanosome, showed 4 days later some crithidial flagellates in the gut and more of them on the 8th day. Two squabs bitten by presumably infected flies and 2 others inoculated with saline emulsions of 3 flies containing many flagellates, all failed to show trypanosomes in their blood for the next two months. In the Class Sporozoa, only the Order Haemosporidia is defi- nitely known to occur in the Hippoboscidae. The complete devel- opment of the Sporozoa usually comprises both an asexual and a sexual cycle. In the Haemosporidia, the two cycles alternate : the asexual cycle (schizogony) occurs in a vertebrate, eventually pro- ducing gametocytes in the blood, .and most of the sexual cycle (sporogony) in an invertebrate, starting with the formation of a zygote in the digestive tract and ending in a form infective to the vertebrate. The blood parasites (hematozoa) of this order are usually classified in two families. The family Plasmodiidae consists of one genus, Plasmodium , which comprises the true malarial parasites of Man and other vertebrates. Although such parasites are frequent in birds, there is no evidence that the sporogony of any of them is completed in the Hippoboscidae or any of the other pupiparous Diptera, or that these flies transmit them to new vertebrate hosts. The Haemoproteidae, with the two genera Haemoproteus and Leucocytozoon, are of much interest to the student of the Hippo- boscidae. At the asexual or schizogony stage, they are frequent parasites in the blood system of many birds and some reptiles, at first in the endothelial cells of the blood vessels and later (as mature gametocytes) inside the circulating cells or blood-corpuscles. The sexual or sporogony stage proceeds in a variety of invertebrate vectors. So far as known, no hippoboscid is involved in this sexual phase of any species of Leucocytozoon. There is, however, con- 137 ENTOMOLOGICA AMERICANA elusive evidence for two species of Haemoproteus of birds that the further development of the gametocytes into mating gametes and the resulting sporogony occur normally in the digestive tract of certain Hippoboscidae. Eventually an infective form, or sporo- fzoite, is produced in the fly and may be inoculated by the bite of the insect into a new avian host. In these species the bird-fly is therefore an essential link in the> transmission from infected to clean birds, hence in the perpetuation of the hematozoa. The fly is also a second true host of these organisms, which are as much para- sites of it as of the bird. Of the two cases, Haemoproteus eolumbae Kruse (1890, p. 370) is the one for which the evidence for a normal transmission, from bird to bird by a louse-fly in nature, is most satisfactory. This blood parasite occurs frequently in domestic pigeons, Columba livia, particularly in tropical and warm temperate regions, and is the causative agent of a serious malaria-like disease. There is fully reliable experimental proof that it is normally transmitted from pigeon to pigeon by the bite of the pigeon-fly, Pseudolynchia canariensis, after undergoing sporogony in the insect. The names Lynchia maura, L. lividicolor and L. capensis, sometimes cited also among the vectors of pigeon malaria, are all synonyms of P. can- ariensis. As for Lynchia brunea , mentioned by Aragao (1907 ; 1908) as one of the vectors, it was based on a misidentification of P. canariensis, the true Pseudolynchia brunnea (Latreille) having nothing to do with the transmission of the disease ; moreover, Aragao synonymized his L. brunea with L. maura in a later paper (1927, p. 827). At one time Aragao (1916, p. 353) included Micro- lynchia pusilla (Speiser) among the bird-flies transmitting H. eolumbae in Brazil; unfortunately he never described his experi- ments with this fly. Microlynchia pusilla is a strictly American bird-fly, known from a variety of hosts, among them 4 wild Ameri- can species of Columbidae ; it has also been taken a few times as a straggler on domestic pigeons in the New World. Its possible role in the transmission of H. eolumbae and other closely related hema- tozoa of native American doves, such as Haemoproteus sacharowi and H. maccallumi, deserves to be investigated more carefully. Haemoproteus lophortyx O’Roke (1929u) causes a grave malaria-like disease in California quail, Lophortyx calif ornica. The parasite goes through the sporogony stage of its life-cycle in the digestive tract of two species of flies, Lynchia hirsuta and Stil- bometopa impressa, both commonly found on quail in western North America, and which act as the vectors of the disease. 138 Volume XXXII Although the sexual stages of H. columbae and H. lophortyx should be regarded as true parasites of the bird-flies in which they develop, it is not known to what extent these hematozoa affect adversely the health or life expectancy of the insects. In the bird hosts they are decidedly pathogenic. For this reason their trans- mission by the flies will be considered more in detail in a later section of this work, while discussing the effect of the hippoboscids on their hosts, an important aspect of their host-parasite relation. Meanwhile it may be noted that the three Hippoboscidae definitely known at present as vectors of malaria-like diseases are among the most host-specific of the bird-flies. In domestication, Pseudolynchia canariensis is a regular parasite of pigeons, having been carried by man on this host to many parts of the world. In the Old World, its original home, it has a wider host range, as shown in the sequel. Both Lynchia hirsuta and Stilbometopa impressa are restricted to the New World, where their only breeding hosts are quail and related game birds of the family Phasianidae. In the Class Cnidosporidia, the Order Microsporidia contains many parasites of insects, usually more or less pathogenic to them. None have been reported to date as occurring normally in the Hip- poboscidae. However, Fantham and Porter (1913) claim to have produced the experimental infection of Melophagus ovinus, by way of the mouth-parts, with Nosema apis Zander, the causative agent of a disease of the honeybee. 3. Fungi. The supposed fungi which certain observers have reported from the internal organs of the Hippoboscidae were bac- teria or related micro-organisms, discussed in earlier sections. Of the several groups of true fungi that normally or occasionally para- sitize arthropods, only the Order Laboulbeniales, of the Class As- comycetes, is thus far known to occur on some of the louse-flies. The Laboulbeniales are peculiarly shaped fungi, of small size, all strictly ectoparasitic on the outer integument of a variety of Arthropoda. These fungi are by no means rare on other pupiparous flies of the families Nycteribiidae and Streblidae ; but thus far they were practically unknown from the Hippoboscidae. Previously they were only mentioned briefly by Speiser (1905a, p. 359) in the de- scription of his Pseudolfersia mycetifera, based on a male from an “eagle” on Senafir Island in the Red Sea. Recently I examined the type of this fly, now at the Vienna Museum, through the cour- tesy of Dr. F. Keiser, and recognized that it belongs to Olfersia fumipennis (Sahlberg). The specific name mycetifera alluded to 139 ENTOMOLOGICA AMERICANA the presence of two clusters of what the author correctly recognized as Laboulbeniales, as I was able to confirm. Speiser stated that these fungi were to be studied by Brunnthaler; but I could not ascertain that they were ever mentioned again in print. Actually the Laboulbeniales are not so very rare on hippo- boscids. They have been merely overlooked, owing to their very small size and to the fact that they shrivel beyond recognition on dry specimens or are destroyed in the process of mounting the flies on slides. My attention was first called to them on a specimen of Al- lobosca crassipes from Madagascar. Shortly afterward I recognized them on the type specimen of Walker’s Ornithomyia simplex, at the British Museum, my identification being confirmed at the time by Mrs. F. L. Balfour-Browne. Some time later I was able to en- list the competent interest of Mr. Richard K. Benjamin, while he was a Graduate Student in the Department of Biology of Harvard University. Mr. Benjamin, who makes a special study of the Laboulbeniales, agreed to go over the several hundreds of louse- flies preserved in spirit, which I had accumulated over the years. The result has been most gratifying, as shown by the subjoined list of the specimens found infested thus far. Subfamily Ornithoicinae 1. Ornithoica pusilla (Schiner). a. Female on Dacelo leachii intermedia, from Dobodura, New Guinea. b. Female on Dacelo gaudichaud, from Hollandia, Dutch New Guinea. c. Female on Tyto aurantia, from New Britain. d. Female on Ninox odiosa, from New Britain. e. Female on Accipiter virgatus gularis , from Loquilocon, Samar, Philippine Islands. 2. Ornithoica stipituri (Schiner). a. Male on Coracina sp., from Dobodura, New Guinea. Subfamily Ornithomyiinae 3. Ornitheza metallica (Schiner). a. Female on Halcyon chloris (subsp.), from Boang Island, Tanga Group, Bismarck Archipelago. 4. Lynchia pollicipes Ferris. a and b. Female and male on Megalaima a. asiatica, from Myitkyina, Burma. 5. Lynchia schoutedeni J. Bequaert. 140 Volume XXXII a. Female on Anhinga r. rufa, from Katwe, Toro, Uganda. b. Female on Anhinga r. rufa, from Kampala, Uganda. 6. Lynchia ( Ornithophila ) simplex (Walker). a. Female (type), without host, from Menado, Celebes. 7. Microlynchia pusilla (Speiser). a. Male on domestic pigeon, from Cali, Yalle del Cauca, Colom- bia. 8. Olfersia bisulcata Macquart. a. One of 6 flies taken on Sarcoramphus papa, at Cayari Island, Uassa Swamp, State of Para, Brazil. 9. Olfersia aenescens C. G. Thomson. a. Female on Sula sula rubripes, from Kaneohe Bay, Oahu, Ha- waiian Islands. 10. Olfersiafumipennis (Sahlberg). a. Male, type of Pseudolfersia mycetifera Speiser, on an “ eagle,” presumably the osprey, Pandion haliaetus , from Senafir Island, on the coast of Arabia, in the Red Sea. Subfamily Alloboscinae 11. Allobosca crassipes Speiser. a. Female on Lepilemur sp., from Ambavoloana, Madagascar. The fungi have now been found on 11 species belonging to 6 genera in 3 of the 6 recognized subfamilies. It is doubtful, how- ever, that they occur on the Hippoboscinae and Melophaginae, as very many specimens of these two subfamilies have been examined. Practically no Ortholfersiinae were available for this particular search. The material will be fully studied and described by Mr. Benja- min in the near future ; but he has kindly furnished the following information of more general interest, with permission to use it in my work. Probably 3 or 4 distinct species are represented in the present collection. It is of particular importance that they all be- long to the one genus Trenomyces, of the family Peyritschiellaceae, and that this genus was previously known only from the biting lice (Mallophaga). All of the hippoboscid fungi are minute, even for the group to which they belong, being among the smallest Laboul- beniales known. The largest species barely reaches 0.6 mm. in length when fully developed and most of them are considerably smaller. They are dioecious, as usual in Trenomyces. They show, however, the unusual peculiarity of being anchored to the integu- ment by means of a long rhizoid (or root), which is sometimes as 141 ENTOMOLOGICA AMERICANA long as the free fungus and penetrates beyond the cuticle in the epidermal cells of the fly. In most other Laboulbeniales the rhizoid is very short and limited to the cuticle of chitin and sclerotin. These parasites do not seem to prefer any particular location on the fly. For instance, of the two infested Lynchia pollicipes, one bore them on the left side at the base of the abdomen, the other on the base of the left wing. On the type of Lynchia simplex, they grew from the dorsal side of the metathorax, right behind the scutellum. The Ornithoica pusilla from Samar carried a thick growth of fungi along the hind margin of the abdomen. It may also be noted that the Trenomyces are known thus far only on hippoboscids from the tropical parts of the Old as well as the New World. Although very many flies were examined from North America and Europe, none were found infested. 4. Parasitic Mites (Acarina). Certain Acarina are the most common ectoparasites of the Hippoboscidae, but are known thus far only from bird-flies (Ornithoicinae and Ornithomyiinae) . They are of more than passing interest as they seem to be connected with specific parasitic mites of birds, although the details of the relation are at present obscure. The life-history has not been elucidated for any of the species and the published accounts are confused. The oldest genus reported from bird-flies is Microlichus Troues- sart and Neumann (1888, p. 134), originally proposed as a sub- genus of Symbiotes and based on Chorioptes avus Trouessart (1887, p. 923). The female of Microlichus avus, more fully described and figured in 1888, was found imbedded in the skin of a domestic sparrow, presumably in France, no exact locality being given. Trouessart and Neumann stated that they found absolutely similar female mites in the skin of three other species of birds from Europe, South Africa and the Antilles ; that all these mites were cospecific seems difficult to believe. They also referred to M. avus a single male taken from a dead sparrow at Vegesack, near Bremen, Ger- many. Vitzthum (1934) doubted that this male was cospecific with the original female and thought that the true male, as well as the eggs and immature stages, of M. avus were as yet unknown. He referred to M. avus a female from Belgium, as it agreed with Trouessart and Neumann’s figure of the species. This female was found by Collart on the body, not on the wings, of an Ornitho- myia avicularia and was not surrounded by eggs. For this reason Vitzthum surmised that M. avus was not a parasite of Ornithomyia, but that it used the fly only to reach a new bird host, where the mite digs in the skin and starts a brood. According to M. Leclercq 142 Volume XXXII (in lift.), the fly on which Collar t found M. avus was the true 0. avicularia and was collected at Glain, Belgium, on a blackbird, Turdus merula. Another mite of the same genus, Microlichus uncus Vitzthum (1934), is somewhat better known. Females were found fixed to the wings of 9 Ornithomyia biloba near Liege, Belgium, no doubt from Hirundo rustica, the normal host of this fly. Vitzthum (1934) and Collart (1934) called the flies Ornithomyia fringillina ; but M. Leclercq, who examined them at the Brussels Museum, recognized that they were 0. biloba Dufour, as Eichler (19395, p. 222) had surmised. The additional Ornithomyia from Brussels, August 8, 1918, on which Collart (1934, p. 2) said he found more M. uncus, was likewise 0. biloba. All were listed as 0. biloba by Bequaert and Leclercq (1947, p. 81). In spite of published statements to the contrary (Ash, 1952, p. 30), the male, larva and nymphal stages of M. uncus have not been described and it is not definitely known where they occur. Col- lart (1934, p. 2) stated that he found on the flies, amidst the eggs of M. uncus, a few larvae, but these were not described nor mentioned by Vitzthum. Collart stressed that all 21 female mites were fixed on his 9 flies to the under side of both wings, invariably at one of the following three definite, clearer points of the veins : about midway on the upper margin of the second basal cell, at the extreme basal elbow of the same cell, and a short distance beyond the base on the lower margin of the anal cell. Most of the female mites were surrounded or hidden by compact clusters of from 1 to 25 eggs. Usually one, more rarely 2 or 3 such clusters occurred on each wing and on one or both wings. Collart believed that the mites select the wing veins deliberately, neglecting the body, and that they even prefer the clear basal elbow of the second basal cell ; perhaps the clearer, less sclerotized stretches of the veins are more easily pierced with the hypostome in order to suck the blood circu- lating in the wing veins. In general, however, the mites infesting the Hippoboscidae do not show such precise selectivity on the body of the fly, as will be shown later for the American species. A female 0. biloba from a cliff swallow, Biparia rupestris, at Banyuls-sur- Mer, France, recently sent by Mr. J. Theodorides, carried a few mites with eggs on the under side of one wing, but many more on one side of the thorax behind. Mr. Theodorides informs me (in litt., 1951) that Mr. J. Cooreman named these mites Microlichus uncus. In order to determine whether female mites prefer certain parts 143 ENTOMOLOGICA AMERICANA of the flies, only attached mite clusters with eggs should be noted. Upon reaching the fly from a bird, the young female mites no doubt wander at random in search of a proper locus of attachment. They may even seek temporary shelter in certain protected areas, which explains why they often occur without eggs in the parascutellar grooves of the thorax. Thirty-five North American Ornithomyia fringillina bearing mite and egg clusters gave the following results : clusters on basal under side of both wings only, 21 ; on basal under side of one wing only, 7 ; on basal under side of one wing and on abdomen, 2 ; on abdomen only, 5. I suggest that mite clusters are restricted on the wings to the under side due to the repeated clean- ing of the upper surface of wings and body with the fly’s legs, which may also account for the scarcity or absence of clusters on the dorsum of the thorax. I have noted that young, non-gravid and unengorged female mites are sometimes found on the basal upper side of the wings, presumably before they are brushed off by the fly. Thompson (19366, p. 319) reports that a female Ornithomyia fringillina , from Anthus pratensis, at Skokholm Id., Wales, car- ried a female mite, surrounded by a mass of eggs, on the basal under side of each wing. The mite was referred by Vitzthum to his Micro- lichus uncus , and, if this identification is correct, the species is not restricted to 0. biloba. Biittiker (1948; 1949, p. 76, figs. 3-4, mislabelled 0. avicularia ) records and figures mite clusters, which he refers to Microlichus uncus, fixed to the basal veins on the under side of the wing of an Ornithomyia biloba, taken from Hirundo rustica at Langenthal, Switzerland. On the other hand, the same author (1949, p. 75, fig. 1) refers to Pterolichus aquilinum (a feather mite of the family Dermoglyphidae) mites from 4 Ornithomyia avicularia on Buteo buteo, in Switzerland: one fly, from Gnosca (Canton Tessin), car- ried a single larval mite on the hind femur ; 3 others, from Zurich, carried 4 female mites with eggs on the wing veins, in the same posi- tion as the mite clusters of M. uncus on 0. biloba. Nothing further is known of the association of Pterolichus with hippoboscids.12 12 1 cannot refrain from suspecting that these so-called (( Ptero- lichus aquilinum” were misidentified and that the four flies from Buteo buteo were infested with Microlichus, like those from Hi- rundo rustica. The size given for the mite attached to the leg ( ca . 1 mm.) is too large for a larva of a feather mite. The figure of the supposed larva could well be that of a gravid female Microlichus with the usual very short and thick first pair of legs of that genus more or less hidden. 144 Volume XXXII Johnsen (1948, pp. 288 and 290), in Denmark, refers doubtfully to M. uncus mites surrounded by eggs, particularly at the under side of the wing of several Ornithomyia avicularia from a rook, Corvus frugilegus, and of 3 0. fringillina from unknown hosts. Ash (1952, p. 30) states that, according to identifications he re- ceived from T. Hughes, M. uncus occurred in 1950 at Sunninghill, Berkshire, England, on both Ornithomyia avicularia and 0. fringil- lina, and M. avus on 0. avicularia only; but, in material taken the same year in Sweden, both species of mites were found together on each of these flies. R. Edwards (1951) refers mites found in Eng- land on 0. fringillina, from wheatear, Oenanthe oenanthe, rock pipit, Anthus spinoletta, English sparrow, Passer domesticus, and starling, Sturnus vulgaris, to both M. uncus and M. avus. These flies were obtained at the Pair Isle Bird Observatory near the Shet- lands. Quite recently, Ash and Hughes (1952) discuss in detail the mites previously reported by Ash (1952) from England and Sweden, listing the bird hosts of the infested flies ; they also com- ment upon some of the structures used to define the species of mites of bird-flies and Mallophaga. Prior to Vitzthum’s paper of 1934, all mites found on hippo- boscids were referred to Myialges Sergent and Trouessart (1907). The type of this genus, Myialges anchora Sergent and Trouessart (1907), was first described and figured from female mites and larvae found on the pigeon-fly, Pseudolynchia canariensis (Mac- quart) (= Lynchia maura Bigot) , in Algiers, the females surrounded by eggs being fixed to the head, thorax and abdomen. The species was later reported from the same host at Pietermaritzburg, Natal (Thompson, 19365), in Southwest Africa (Reichert, 1939), in Mauritius (Thompson, 19365), and in California (Herman, 1945). A. C. Oudemans (1935) published a revised illustrated description of the female and larval types. M. anchora was described again from the pigeon-fly, at Recife, Brazil, as Myialges pseud, olinchiae, by de Pigueredo and Simoes Barbosa (1944). It seems to be found wherever its fly host occurs, and even flies taken from Avild bird hosts in the Old World are often infested. Sometimes it is very common. Of 31 P. canariensis (15$ and 16^) collected at random from domestic pigeons at Aba, Belgian Congo, 17 (9$ and 8^) car- ried female mites, a few without, but most of them with a cluster of eggs. The majority were fixed to the abdomen, but some also to the thorax, particularly on the prosternum between the fore coxae. In 5 flies a female mite, either with or without eggs, was attached to the base of the rostrum membrane, on the under side of the head. 145 ENTOMOLOGICA AMERICANA Only one of the flies carried one mite cluster on the under side in the basal area of one wing, and it was also infested on the abdomen, thorax, and rostrum membrane. At Tucuman, Argentina, accord- ing to Dr. Schuurmans Stekhoven (in lift., 1952), the percentage of mite-infested P. canariensis was about 18 per cent for the females and 12.5 per cent for the males in the most heavily parasitized lot ; the mites were attached mostly either to the head at the base of the palpi or to the sides near the tip of the abdomen, occasionally also in the space between the thorax and the abdomen. In the original description, Sergent and Tronessart noted the absence of a caruncle or adhesive pad on the first pair of legs, the tarsus ending instead in a heavy, two-hooked, anchor-shaped claw, which fastens the mite in the fly’s cuticle. As Dr. D. Furman pointed out to me (in lift ., 1952), the stalked eggs are fixed to the fly ’s cuticle by a thin base or plaque probably secreted by the female mite at the time the egg is voided. When removing egg clusters from a fly, this thin base is often removed intact. Ferris (19285) published some new figures and critical notes of M. anchora, based on specimens from three species of Ornithoica, 0. vicina (Walker) in North America, and 0. pusilla (Schiner) and 0. philippinensis Ferris in the Philippines.13 Thompson (19365) found 2 females with egg masses of M. anchora (as named by Yitzthum) fixed to the posterior lobes of the abdomen of a female Ornithomyia fringillina, from Dryocopus m. martins, in Esthonia; I recently confirmed the identification of these flies. A second species of the genus, Myialges caulotoon Speiser (1907, pp. 9-10), was based on several female mites surrounded by eggs, found mostly on the abdomen and in one case also on a leg of 2 Lynchia albipennis (Say) (= Olfersia ardeae Macquart), from Ardea purpurea, in Tanganyika Territory (Kibonoto on Mt. Kili- manjaro and Tanga) . It was not figured by the author. The eggs, 13 Originally Ferris (19285) recorded M. anchora “from Orni- thoica confluent a (=0. promiscua), Pasadena, California, and Puerto Princesa, Palawan, Philippine Islands, and from 0. philip- pinensis, Mati, Davao, Mindanao, Philippine Islands.” The first of these flies, from Pasadena, was what I call Ornithoica vicina (Walker). The second, from Puerto Princesa, was later referred to 0. pusilla (Schiner) by Ferris (1929, p. 284). As for 0. phil- ippinensis Ferris, it appears to be a peculiar species, to be discussed in Part II of the present work. Ferris did not give the bird hosts of the infested flies. 146 Volume XXXII several dozen in each mass, were placed upright and fixed by a long, very thin, hyaline, stiff but curved stalk to the integument of the fly. Ferris (19285) redescribed and figured as M. caulotoon fe- males surrounded by eggs, found mostly on the abdomen, in one case also on the thorax, of Ornithoica pusilla (Schiner) (cited as 0. confluent a) and 0. philippinensis Ferris from the Philippines, the same flies also carrying mites referred to M. anchora , as men- tioned before. But he was not certain that the mites were correctly named and even doubted that they were two distinct species. He voiced the suspicion that “actually the two forms have some con- nection with each other” and that “one is simply a developmental form of the other.” E. P. Peed (1932) recorded Myialges caulo- toon (from Ewing’s identification) on several Ornithomyia remota Walker (as 0. chilensis Reed, a synonym), at Valparaiso, Chile, from unknown bird hosts. Spencer (1928; 1938), Thompson (19366) and I (1933e; 1935a) referred to the genus Myialges mites from several other bird-flies. Mention should also be made of mites found by Shipley (1911, 1, p. 360) on 3 Ornithomyia frin- gillina (as 0. lagopodis) from red grouse, Lagopus scoticus, in Eng- land. These Waterston referred to the genus Canestrinia, but doubtless they were either Microlichus or Myialges. Edwards (1952) reports from the Fair Isle Bird Observatory near the Shet- lands an Ornithomyia avicularia, taken on a twite, Carduelis flavi- rostris , and carrying on the abdomen several mite clusters tenta- tively referred to Myialges. Thompson (19365, p. 316) reported mites, similar to those of the bird-flies, fixed to biting lice (Mallophaga) from wild ducks (Ana- tidae) in Uganda; Vitzthum referred these to Myialges caulotoon. Later (1939) he observed such mites also on lice from Anatidae of North America and Ceylon, and published some excellent photo- graphs of the infestations. Cooreman (1944) proposed for mites fixed to biting lice from a wild duck, Mergus merganser, in Bel- gium, a new genus Myialgopsis, with Myialgopsis trinotoni Coore- man as the type, said to differ from Myialges in the presence of a tarsal caruncle on the first pair of legs. According to Radford (1949), Thompson’s supposed Myialges caulotoon, from lice on Anatidae, were likewise Myialgopsis trinotoni. However, the pres- ence or absence of a tarsal caruncle appears to be an unreliable character, so that Myialgopsis is perhaps not generically separable from Myialges. From the point of view of the life-history and ecology of the mites, the true relationship of those of the Hippoboscidae and of 147 ENTOMOLOGICA AMERICANA the Mallophaga and their correct place in the general classifica- tion of the Acarina are of great importance. Sergent and Troues- sart (1907) originally placed Myialges in the family Sarcoptidae; but, as now defined, the Sarcoptidae always have a pair of anterior vertical setae on the dorsal propodosoma (Baker and Wharton, 1952, p. 362), which setae are lacking in Myialges. At one time, Vitzthum (1934, pp. 12-13) regarded Microlichus and Myialges as closely related and was even doubtful about their being generically distinct. In his later comprehensive work on the Acarina (1942, pp. 890 and 898), however, he placed them in two different fami- lies : Myialges in the Myialgesidae and Microlichus in the Epider- moptidae. The Myialgesidae, which he said are “ parasitic on Hippoboscidae, exceptionally on Mallophaga,” were characterized by the tarsi of the first pair of legs lacking caruncles, which are present only on the second, third and fourth pairs. On the other hand, he stated that the Epidermoptidae live “on the skin of birds” and have caruncles on all four pairs of tarsi. In their recent “ In- troduction to Acarology,” Baker and Wharton (1952, pp. 324, 369 and 374) follow Vitzthum ’s classification. However, they include in the Myialgesidae Cooreman ’s genus Myialgopsis, although it was defined by the presence of caruncles on the first pair of tarsi.14 The presence or absence of a caruncle (adhesive pad or ambu- lacrum) on the tarsi of the first legs is a fictitious character. Vitz- thum (1934, p. 13) at first suspected that the caruncle might have been merely overlooked in Myialges ; while Cooreman (1944, p. 3) suggested that the caruncle may be fugacious, being present in the newly-emerged female and lost after the mite anchors in the cuticle. That the caruncle is occasionally present on all four pairs of tarsi in Myialges anchora of the pigeon-fly, was recently observed by Dr. D. P. Furman (in lift ., 1952). Myialges, Myialgopsis, and Microlichus may therefore all be included in the Epidermoptidae and the family Myialgesidae should be dropped. Moreover, the remaining differences between these three so-called genera are very weak and perhaps not of more than specific value, in which case all the mites found attached to Hippoboscidae and Mallophaga could 14 Vitzthum (1942, p. 890) and Baker and Wharton (1952, p. 369) credit the family name Myialgesidae to Trouessart, 1907 ; but Trouessart did not place Myialges in a distinct family nor propose the family name, which dates from Vitzthum, 1942. It may be added that Cooreman (1944) proposed, “at least provisionally,” an additional family Myialgopsidae for Myialgopsis. 148 Volume XXXII be placed in the one genus Microlichus. It is fairly certain that several species of mites are involved; how many of those now de- scribed are valid cannot be decided from the available evidence. With the relationships of the mites of the Hippoboscidae some- what clarified, it becomes more probable that in all of them the major part of the life-history proceeds on or in the skin of birds, which normally harbor the larvae after they leave the fly, the sev- eral nymphal stages and the newly emerged adults of both sexes. After mating on the bird, some or possibly all females migrate to Hippoboscidae, where they attach to the integument, become fully gravid and eventually oviposit, the eggs clustering around the mother mite. After hatching from the egg, the mite larva probably returns to a bird before starting to feed. In this manner a fly in- fested with mites may transfer larvae to a new bird, particularly to nestlings, and this may possibly be the normal mode of dispersal of most species of Microlichus. If this view is correct, Microlichus should be included among the few cases of true hyperparasitism known among the Acarina. While the female mites are not merely “ riders, ” but undoubt- edly true parasites, since they extract nourishment from the body fluids of the fly, it is difficult to estimate the damage they do to the insect. Perris (19285) called attention to the peculiar scars formed where the mites are fixed to the integument. Mr. K. W. MacAr- thur {in litt., 1951) also noted these strongly sclerotized scars pro- duced by the mites, which persist even after prolonged treatment with potash. They may be seen in a photograph of the tip of the abdomen of Ornitliomyia fringillina received from Mr. MacArthur and published here with his permission (Pig. 19). Heavy infesta- tion, particularly when the wings carry many egg clusters, might also impair flight by the mere weight of the parasites. Often a substantial proportion of the general bird-fly popula- tion is mite-infested. For instance, during the summer of 1951, at the Fair Isle Bird Observatory near the Shetlands, according to Edwards (1952), mites were found on 47 of 362 specimens, or 13 per cent, of Ornithomyia (mostly 0. fringillina) taken from 165 birds of 4 species. The following partial list of New World Hippoboscidae observed carrying mites, probably all referable to the genus Microlichus , could be considerably extended. The term “mite cluster” is here used for a female mite surrounded by her eggs. The generic and specific names of the mites are retained as given by the several authors cited, although eventually they will probably all be placed 149 ENTOMOLOGICA AMERICANA Fig. 19. Ornithomyia fringillina Curtis, female from Demarest, New Jersey, on Dumetella carolinensis. Cleared tip of abdomen with gravid female mites attached, showing the sclerotized scars made by the mites. Photograph by Mr. K. MacArthur. 150 Volume XXXII in the genus Microlichus. The list now contains mite infestations on 20 American species of flies belonging to 8 genera. It shows that these parasites occur everywhere, in temperate as well as in tropical areas, and on many flies from a variety of bird hosts. Their prevalence strongly suggests that in most species of Micro- lichus preliminary feeding of the female on bird-flies, followed by oviposition on the insect, is a regular and essential feature of the mite’s life-history. If so, the flies should be regarded as required intermediate hosts as well as carriers of the mites, and a normal link in their life cycle. A. Ornithoica vicina (Walker) (= 0. confluent a of authors, not of Say). 1. Ferris (1928 b, p. 139) referred to Myialges anchora mites attached to the abdomen of a fly (cited as 0. confluent a) from an unknown host at Pasadena, California. 2. Herman (1937 b, p. 165) mentions mites attached to a fly (cited as 0. confluenta) at North Eastham, Massachusetts; host unknown. 3. Female on Pipilo fuscus falcifer , from Berkeley, California, with a mite cluster on the under side of a vein in the basal third of the wing. 4. Fly from Valparaiso, Chile, on unknown host, with several mite clusters. 5. Female on Buteo magnirostris, from Juquia, State of Sao Paulo, Brazil, with several mite clusters. 6. Female on juvenile Turdus m. migratorius, from Demarest, New Jersey, with one large mite cluster at the right side of the tip of the abdomen. 7. Two females from near Valparaiso, Chile, on “zorzal,” Turdus magellanicus, in the E. P. Reed Collection, carried mite clusters at the side of the dorsum of the abdomen, one on one fly, two on the other. 8. Out of a total of 29 flies collected from 6 Junco phaeonotus dorsalis, near Flagstaff, Arizona, by Mr. A. R. Phillips, 5 females (from 2 birds) carried mites, all attached to the veins on the under side of one or both wings; some of the mites were surrounded by eggs, others not. 9. Among 7 flies taken from young domestic fowl by S. L. Yoder at Annandale, Virginia, one female carried 2 mite clusters on the sides of the abdomen, one near the tip on the right side, the other about mid-length on the left side. 10. Female on English sparrow, Passer domesticus (introduced 351 ENTOMOLOGICA AMERICANA from Europe), taken by Dr. C. M. Herman at Silver Spring, Mary- land, with one mite cluster on the under side of the right wing, fixed to the vein closing the base of the anal cell. B. Ornitkomyia fringillina Curtis. 1. Female on Pipilo fuscus falcifer, from Berkeley, California, with several mite clusters on the dorsum of the abdomen. An Or- nithoica vicina, taken from the same individual bird, was also in- fested with mites, as recorded above. 2. G. J. Spencer (1928, p. 257) reports mite clusters in the basal area of both wings of a fly taken on Cyanocitta s. stelleri at Tofinb, Vancouver Id. Cooreman (1944, p. 8) lists the mite of this case under Microlichus, a generic name which Spencer did not mention, however. The same fly (called Ornitkomyia avicularia by Spencer) carried also 16 Mallophaga. 3. G. J. Spencer (1938, p. 43) mentions a mite cluster, sup- posedly of Myialges sp., on a female from a hawk, at Jesmond, British Columbia, fixed to the tip of the abdomen. 4. G. J. Spencer (1938, p. 43) reports several clusters, sup- posedly of Myialges sp., on a fly from Acridotheres crist at ellus (in- troduced from the Far East, now feral in British Columbia), at Vancouver, fixed on the dorsum of the abdomen. 5. Female on Dumetella carolinensis, from Demarest, New Jer- sey, with several mite clusters attached to the hind margin of the abdomen. The scars made by the mites are shown in Fig. 19. 6. Female on Hylocickla mustelina , from Demarest, New Jer- sey, with several mite clusters on the under side of the basal area of both wings. This fly also carried 2 Mallophaga. 7. Female on Zonotrickia albicollis, from Demarest, New Jer- sey, heavily infested with at least 7 egg-laying mites, along the en- tire periphery at the sides of the abdomen. 8. Female on Pipilo e. erythropkthalmus, from Demarest, New Jersey, with one mite cluster at the under side of the basal area of each wing. 9. Female on Geothlypis trichas brachidactyla, from East West- moreland, New Hampshire, with mite clusters in both wings. 10. Female on Geothlypis trichas brachidactyla , from Demarest, New Jersey, with one small mite cluster in the basal area of the left wing, on the under side. 11. Female on Molothrus a. ater, from Demarest, New Jersey, with one small mite cluster in the basal area of the right wing, on the under side. 12. Male on Cyanocitta c. crist ata, from Demarest, New Jersey, 152 Volume XXXII with a small mite cluster on the under side of each wing, in the basal area. 13. Female on Dumetella carolinensis, from Demarest, New Jersey, with one mite cluster in the basal area of the right wing, on the under side. 14. Male on Dumetella carolinensis, from Demarest, New Jer- sey, with several small mite clusters on the under side of each wing in the basal area. 15. Female on Hylocichla mustelina, from Demarest, New Jer- sey, with 2 mite clusters in the basal area of the left wing, on the under side. 16. Female on Melospiza m. melodia, from Barre, Vermont, with one mite cluster in the basal area of the left wing, on the under side, and another on the left side near the tip of the abdomen. 17. Male on Hylocichla mustelina, from Demarest, New Jersey, with a few non-gravid female mites, without eggs, near the base on the upper side of both wings. 18. Male on Icterus galbula, from Demarest, New Jersey, with one mite cluster in the basal area of each wing, on the under side. 19. Male on Icterus galbula, from Demarest, New Jersey, with several mite clusters in the basal area of each wing on the under side. 20. Male on Hedymeles ludovicianus , from Demarest, New Jer- sey, with several non-gravid female mites, without eggs, on various areas of the wings, thorax and abdomen ; 3 of the mites in the left parascutellar groove. 21. Female on Icterus galbida, from Demarest, New Jersey, with several non-gravid female mites, without eggs, on various areas of the thorax and on the upper side in the base of the wing. 22. Female on Melospiza m. melodia, from Demarest, New Jer- sey, with several mite clusters in the basal area of both wings, on the under side. 23. Female on Dumetella carolinensis, from Demarest, New Jersey, with a few non-gravid female mites, without eggs, scattered over the body and apparently not attached. 24. Male on Dumetella carolinensis, from Demarest, New Jer- sey, with mites as in 23. 25. Female on Dumetella carolinensis, from Demarest, New Jersey, with mites as in 23. 26. Female on Dumetella carolinensis, from Demarest, New Jersey, with several non-gravid female mites, without eggs, wedged in the parascutellar grooves, on both sides. 153 ENTOMOLOGICA AMERICANA 27. Female on Pipilo e. erythrophthalmus, from Demarest, New Jersey, with mite clusters on the under side in the basal area of both wings, one in one wing, two in the other. 28. Female on Toxostoma rufum, from Demarest, New Jersey, with 2 mite clusters on the under side in the basal area of the right wing, and a few non-gravid female mites, without eggs, in both parascutellar grooves. 29. Male on Pipilo e. erythrophthalmus , from Demarest, New Jersey, with only non-gravid female mites, without eggs, a few on the basal area at the under side of both wings and several in both parascutellar grooves. 30. Male without host, from Barre, Vermont, with 2 mite clus- ters on the under side in the basal area of the right wing and 3 clusters on the abdomen. 31. Female on Hylocichla mustelina , from Demarest, New Jer- sey, with one mite cluster on the under side in the basal area of the right wing. 32. Male without host, from Barre, Vermont, with several non- gravid females, without eggs, in the basal area of the upper and under side of the left wing. 33. Female on Icterus galbula, from Demarest, New Jersey, with several non-gravid females, without eggs, in various areas of the abdomen and a few on the under side of the basal area of both wings. 34. Male on Icterus galbula, from Demarest, New Jersey, with several non-gravid females, without eggs, in various areas of the body, particularly in the parascutellar grooves, and a few on the upper surface in the basal area of both wings. 35. Male on Dumetella carolinensis, from Demarest, New Jer- sey, with many non-gravid female mites, without eggs, in various areas of the body and a few in the basal area on the upper side of both wings. 36. Female on Quiscalus q. quiscula, from Pequannock, New Jersey, with many non-gravid female mites, without eggs, most of them flattened against the integument and closely packed together on the back of the left side of the abdomen. 37. Male on Dendragapus obscurus fuliginosus, from Kamloops, British Columbia, with 2 mite clusters on the dorsum of the ab- domen. 38. Male on Ixoreus n. naevius, from Trinity Valley, Vernon, British Columbia, with mite clusters in the basal area on the under side of both wings, 1 in one wing, 2 in the other. 154 Volume XXXII 39. Female on Bonasa umbellus umbelloides, from Okanagan Landing, British Columbia, with several mite clusters in the basal area on the under side of both wings. 40. Female on Hylocichla guttata faxoni, from Mt. Desert Is- land, Maine, with mite clusters on the under side in the basal area of both wings. 41. Female on Molothrus a. ater, from Old Deerfield, Massa- chusetts, with mite clusters on the under side in the basal area of the left wing. 42. Female on Dendragapus obscurus richardsoni, from Bridger Mountains, Montana, with several mite clusters on the under side in the basal area of both wings. 43. Female on Zonotrichia albicollis, from Columbus, Ohio, with mite clusters on the under side in the basal area of both wings, 4 in one wing, 2 in the other. In this species of Ornithomyia, when female mites with egg clus- ters are present on the wings, they are always fixed to the veins surrounding the basal and anal cells and on the under side. The foregoing list records infested flies from 20 different species of birds (16 Passeriformes, 3 Galliformes and 1 Falconiformes). The rate of infestation seems to vary from year to year and is presumably correlated with similar variations of the mite infestation of the bird population. In the summer and fall of 1950 mite infestation was unusually low on the flies at Demarest, New Jersey, only 2 being infested among 46 fringillina taken by Mr. B. S. Bowdish on 37 passerine birds of 14 species, all trapped alive. C. Ornithomyia remota Walker. As mentioned before, E. P. Reed (1932) recorded an infestation of Myialges caidotoon on a fly of this species in Chile. 1. Three females on Nesocichla eremita gordoni, from Inacces- sible Id., Tristan da Cunha Group, each with several mite clusters on the under side of the veins in the basal areas of both wings. D. Stilbometopa podopostyla Speiser. 1. A female on Leptotila verreauxi decipiens, from Barra do Tibagl, Rio Paranapanema, State of Sao Paulo, Brazil, carried small mite clusters on the hind part of the thorax, to the side of the scutellum. Herman (1945, p. 24) noted that among several hundred speci- mens of Stilbometopa impressa (Bigot), a common fly on Lophortyx calif ornica vallicola in California, he never found one with mites, while they were not uncommon on Lynchia hirsuta taken on the 155 ENTOMOLOGICA AMERICANA same host. Mr. I. B. Tarshis (in lift., 1951) confirmed this for both flies from his own observations. E. Ornithoctona erythrocephala (Leach). 1. Two females on Oreopeleia linearis infusca, from Yista Nieve, Sa. Marta, Colombia, each with mite clusters on the abdomen, mostly near the tip ; one cluster ventrally near the base. 2. Four females on one Accipiter s. velox, from Whitefish Pt., Michigan, with mite clusters on the dorsum of the abdomen; the dorsum of one fly was nearly covered with them. 3. Female on a species of Falco , from Hansa near Humboldt, Sa. Catharina, Brazil, with 6 female mites, some of them with a few eggs, on the dorsal side of the abdomen. F. Ornithoctona fusciventris (Wiedemann). 1. Two females on Calospiza arthus aurulenta, from Yirolin, Santander, Colombia, with mite clusters on the thorax, chiefly wedged in the deep parascutellar grooves ; one of the flies also with a cluster on the sternum, in the notch between the prosternal lobes, and with another cluster on the under side of the articulation of a wing. G. Lynchia albipennis (Say). 1. Female on Botaurus lentiginosus, from St. Paul, Minnesota, heavily infested. In addition to 2 large clusters on the under side of the basal area of the left wing, 4 clusters on both side margins of the abdomen, some females starting to oviposit on the dorsum of the abdomen, and one female with a few eggs on the anterior face of the right fore femur. 2. Male on Botaurus lentiginosus, from Patterson Lake, Michi- gan, with 4 mites : one with one egg on the under side in the basal area of the right wing ; one without eggs on the left side near the tip of the abdomen; one without eggs on the left mid femur; and one with a few eggs on the gula of the head. 3. Female on Botaurus lentiginosus, from Bald Eagle Lake, Minnesota, very heavily infested. This fly was briefly mentioned by MacArthur (1948, p. 387). Nearly all the clusters on the ab- domen, the wings being entirely free from mites. Seven females with egg masses on the venter, 4 of them in the basal median area ; at least 9 more females, mostly imbedded in large clusters of eggs at the sides of the dorsum, 2 females as yet without eggs. In addi- tion one female, without eggs, fixed to the integument of the right fore femur. 4. Two females on Botaurus lentiginosus, from Racine, Wis- consin, each with several mite clusters. 156 Volume XXXII 5. Male on Botaurus lentiginosus, from Philadelphia Neck, Pennsylvania, with mite clusters. It seems remarkable that 5 specimens of this common fly ob- served carrying mites, were taken on Botaurus lentiginosus. 6. Female from an unknown host at Iowa City, Iowa, with 3 large mite clusters on the dorsal sides of the abdomen and one small cluster on the under side of the base of the left wing. H. Lynchia angustifrons van der Wulp. I. A female fly on Accipiter s. velox, from Vineland, Ontario, May 13, 1924, is the most heavily infested hippoboscid I have ob- served. Both upper and under surfaces of the abdomen are nearly completely covered with crowded mite clusters. It is impossible to determine exactly how many females deposited the eggs, but they numbered at least fifty and the eggs run into the hundreds. In addition, there is a large mite cluster on the head, attached beneath the base of the palpi, in such a way that it would seem the fly was eventually unable to feed. The wings, thorax and legs, however, are entirely free of mites. I. Lynchia nigra (Perty). 1. Three females on Buteo jamaicensis borealis, from Grindrod, British Columbia, each with a single female mite, without eggs, on one side of the abdomen. 2. Female on Accipiter bicolor pileatus, from Fazenda Becreio, Coxim, Matto Grosso, Brazil, with one mite cluster on the right side at the hind third of the abdomen. 3. Male on Milvago c. chimachima, from Bio das Mortes, Posto Pindaiba, Matto Grosso, Brazil, with 3 mite clusters on the sides and venter of the abdomen, a small group at the under side in the basal area of one wing, and a large cluster on the right hind tibia. 4. Female on Buteo jamaicensis calurus, from Tucson, Arizona, April 5, 1916, with a gravid female mite, without eggs, fixed on the dorsal left side of the abdomen. 5. Female on Cathartes a. aura, from east of Tucson, Arizona, May 23, 1949 (A. B. Phillips), with 2 female mites fixed on the right mid femur, one without and the other with eggs. J. Lynchia fusca (Macquart). Herman (1945, p. 23) states that he saw mite clusters on several specimens from hawks and owls in California. He has sent me more definite information on the following two cases. 1. Eleven flies on one Bubo virginianus subsp., from Napa Co., were all infested ; but flies emerging from 7 puparia deposited by these infested flies, were free of mites. The clusters usually con- 157 ENTOMOLOGICA AMERICANA sisted each of a female mite surrounded by a large mass of eggs, and were fixed to the under side of the wing veins and to the venter of the abdomen near the tip. See fig. 12 in Herman’s paper. 2. Two males and 6 females among 19 flies on one Bubo vir- ginianus subsp., from Lassen Co., were infested with mites. 3. Two flies on Otus asio floridanus, from Auburn, Alabama, with several mite clusters on the dorsum of the abdomen. . 4. Two females on an unknown host, from Ipiranga, State of Sao Paulo, Brazil, each with 2 mite clusters on the abdomen, dor- sally on one fly and ventrally on the other. 5. One male and one female on Tyto alba tuidara, from Ipi- ranga, State of Sao Paulo, Brazil, with mite clusters on the dorsum of the abdomen, 8 on the male and 5 on the female. 6. Female from an unknown host, from Monte Alegre, State of Sao Paulo, Brazil, with 4 mite clusters on the sides of the abdomen. K. Lynchia kirsuta Ferris. 1. Herman (1945, p. 23) reports seeing mite clusters on one of 11 flies on Lophortyx calif ornica vallicola, in California. He writes me that he examined since 36 more of these flies without finding mites. 2. Mr. I. B. Tarshis (in lift., 1951) observed mites on about a dozen flies on Lophortyx calif ornica vallicola , from various Califor- nian localities. 3. One female and 3 males on Lophortyx calif ornica, from the Hastings Natural History Reservation near Jamesburg, Monterey Co., California, each with from 1 to 3 mite clusters on the abdomen. Mr. Tarshis informs that at the Hunter Liggett Military Reser- vation, San Benito Co., California, about 2.2 per cent of some 900 L. hirsuta, taken from L. c. vallicola, were infested with mites. L. Pseudolynchia canariensis (Macquart) (= Lynchia maura Bigot). It may be of interest that one of Macquart ’s two cotypes of canariensis, from the Canary Islands, now at the Paris Museum, carries 6 mite clusters on the hind part of the abdomen (J. Be- quaert, 1935a, p. 399). 1. I have mentioned before Myialges pseudolinchiae de Figu- eredo and Simoes Barbosa (1944), described from a fly on domestic pigeon at Recife, Brazil. I regard this mite as a synonym of Myialges anchora. 2. Fly on domestic pigeon, fram Havana, Cuba, with mite clus- ters, according to Dr. J. Perez Vigueras (J. Bequaert, 1935a, p. 397). 3. Bishopp (19295, p. 980) reports that mites, probably of the 158 Volume XXXII germs Myialges, were attached in great numbers to some flies on domestic pigeons at Lockhart, Florida. 4. Dr. C. M. Herman informs me {in lift., 1951) that 2 females out of a lot of 3 males and 2 females, on domestic pigeons, from San Francisco, California, were infested with mites. 5. Out of 5 flies taken on domestic pigeons, at Rio de Janeiro, Brazil, 3 females were infested with mites. The heaviest infesta- tion consisted of 4 clusters on the sides and venter of the abdomen of one fly. The other 2 flies each had one cluster on the abdomen, and one of them carried besides a female mite, without eggs, on the rostrum membrane of the head. 6. A female on a domestic pigeon at Atlanta, Georgia, received from Mr. P. W. Fattig, carried one small mite cluster near the dorsal tip of the abdomen. 7. The high rate of mite infestation observed by Dr. J. C. Schuurmans Stekhoven and his co-workers at Tucuman, Argentina, was mentioned before. 8. Male from a domestic pigeon, at Ames, Iowa, August, 1929 (H. M. Harris), with 2 female mites on the abdomen: one non- gravid, dorso-laterally on the left side, some distance before the tip ; the other with a large package of eggs on the right side near the tip ; in this cluster only the female mite was attached to the integu- ment, all the eggs being fixed to the venter of the mite, some of them being stalked and others sessile. M. Pseudolynchia brunnea (Latreille). 1. Male on Chordeiles m. minor , from Ipiranga, State of Sao Paulo, Brazil, with several mite clusters on the basal veins at the under side of one wing. N. Microlynchia pusilla (Speiser). 1. Dr. C. M. Herman {in lift 1951) found mites on a female on Zenaidura macroura marginella, from Brawley, California, col- lected by G. F. Augustson. This case was inadvertently referred by Herman (1945, p. 23) to Pseudolynchia canariensis, the author informs me. O. Microlynchia crypturelli J. Bequaert. 1. Female on Columba rufina sylvestris, from Nova Teutonia, State of Santa Catharina, Brazil, with 2 small mite clusters on the thorax, one on each pleurotergite, immediately behind the base of the wing. P. Olfersia spinifera (Leach). 1. Two flies on Fregata minor ridgwayi , from Tower Island, Galapagos : one fly with a female mite on the upper side of the first 159 ENTOMOLOGICA AMERICANA longitudinal vein, close to the base of the left wing; the other fly with a female surrounded by many stalked eggs on the left meso- pleuron, immediately behind the base of the fore leg. At one time (J. Bequaert, 1933e), I referred these mites to Myialges. 2. The male type of Ornithomyia unicolor Walker, a synonym of 0. spinifera, from Fregata magnificens rothschildi in Jamaica, bears a mite cluster on the sides of the metathorax, immediately above the insertion of the right hind leg. Q. Olfersia fossulata Macquart. 1. Thompson (193 61), p. 316) records a fly on Pelecanus sp., from South America, with one mite cluster, referred to Myialges sp., on the under side of the metathorax. R. Olfersia coriacea van der Wulp. 1. Male on Crax fasciolata, from Goyaz, Brazil, with a single female mite, without eggs, fixed by the hypostome to the left side of the abdomen. S. Olfersia bisulcata Macquart. 1. Male on Coragyps atratus foetens, from Rio Tapajos, State of Para, Brazil, with one large mite cluster near the mid-dorsum of the abdomen. T. Olfersia fumipennis (Sahlberg). 1. Female from British Honduras (Newman), on Pandion haliaeetus carolinensis, with a cluster of eggs and a female mite on the under side of the basal veins of the right wing and a gravid female mite without eggs on the right side of the metathorax below the insertion of the wing. 5. Parasitic Hymenoptera. The early instars of most Diptera are frequently destroyed by various parasitic Hymenoptera or more rarely by certain other Diptera. The Hippoboscidae are un- usually free from such attacks, no doubt due chiefly to the very short duration of the extra-uterine development, the only time at which parasitic insects could oviposit in the newly-voided, soft- skinned larva before it hardens into a puparium. The fact that only single larvae are deposited at rather long intervals by the gravid female flies, which, moreover, usually scatter them at ran- dom or may even hide them, further decreases the parasite ’s chances of encountering them at the proper oviposition time. The few in- sects known to parasitize hippoboscid puparia are minute Hymen- optera, probably all of the group Chalcidoidea. None are specific or restricted to the Hippoboscidae, but they are instead polyphagous parasites with a wide host range. Modeer (1785, p. 37), in Sweden, first mentioned that he bred 160 Volume XXXII both sexes of an “ichneumon” (a term then used for all parasitic wasps) from several puparia of Stenepteryx hirundinis (= Hippo- bosca hirundinis). He stated that about 30 ferruginous wasps, a little larger than fleas, emerged from each puparium. They ap- peared early in August from puparia collected in swallows’ nests the same spring, but only about mid- June of the next year from puparia taken in the autumn. Zetterstedt (1848, 7, p. 2909) sug- gested that Modeer’s parasite might have been one of the Encyr- tidae, although he did not see the specimens. It should be added that quite possibly Modeer bred his parasites from puparia of Crataerina pallida and not of Stenepteryx hirundinis, these two flies being commonly confused by the early entomologists. In any case, no insect parasites have since been obtained from Stenepteryx. Cornelius (1869, p. 407) bred a “Pteromaline chalcid fly” from puparia of Crataerina pallida found in the fall in a nest of Euro- pean swifts, Apust apus, near Eberfeld, Germany, the parasite emerging the next April ; no further identification was attempted. Bagnall (1915) reared C. pallida from puparia in a swift’s nest at Newcastle, England ; but one of the puparia yielded instead a small chalcid. This was possibly the parasite which Waterston later found in Bagnall ’s collection and identified as Mormoniella vitri- pennis (Walker), according to Gahan (1927, p. 6) ; although Waterston stated, perhaps by an oversight, that the specimens had been “reared from a puparium of Stenopteryx found in a house- martin’s nest.” M. vitripennis (= Nasonia brevicornis Ashmead), one of the Pteromalidae, is a widespread parasite of many Diptera, particularly muscoids. Recently Mr. G. E. Woodroffe sent me some chalcids he bred from a puparium of Crataerina pallida in England. These were kindly identified by Dr. D. B. Burks, of the U. S. Bureau of Entomology and Plant Quarantine, as Dibrachys cavus (Walker), also one of the Pteromalidae and likewise a cos- mopolitan parasite of a variety of insects in several orders. Stadler (1948, p. 52) states incidentally that “an entirely yellow braconid” issued from a puparium of an undetermined Ornitho- myia found in Germany in the nest of a sand martin, Biparia riparia. It is unfortunate that nothing more is known of this in- teresting parasite. The host was, I suspect, Ornithomyia biloba. Schuurmans Stekhoven and his co-workers, in Tucuman, Ar- gentina, found 4 specimens of a chalcid in a pupaauum of the pigeon-fly, Pseudolynchia canariensis (unpublished observations). The parasites were identified by Dr. Luis De Santis, of the Museum of La Plata, as Mormoniella vitripennis (Walker). 161 ENTOMOLOGICA AMERICANA Fig. 20. Ornithomyia fringillina Curtis, male from Demarest, New Jersey, on Dumetella carolinensis, with Mallophagan attached. From MacArthur, 1S48, fig. 233. 162 Volume XXXII It is interesting to note that the only insect parasites of louse- flies which have been definitely identified are two cosmopolitan species of Pteromalidae with a wide range of hosts. Furthermore, they have been bred thus far only from species of bird-flies whose puparia are often abundant in the nesting sites of their hosts, swal- lows, martins, swifts and domestic pigeons, where many birds breed together. These factors tend to increase the local louse-fly popu- lation and particularly the number of larvae deposited in a rela- tively small area, thus favoring parasitism by polyphagous chaleid wasps. Phoresy of Mallophaga. Certain Hippoboscidae of birds oc- casionally carry one or more biting (or chewing) lice (Mallophaga), attached by the mandibles to the integument, usually at the base of a body seta, more rarely to a vein of the wing.15 Such cases of so-called “phoresy” are on the border line of true parasitism. Each fly usually carries only one or two lice ; exceptionally there may be more, respectively 15, 16 and even 31 in three observed cases. K. W. MacArthur (1948) mentions an Ornithomyia fringil- lina, from T urdus m. migratorius, with 8 lice placed close together in a semi-circle, along the sides of the abdomen, all heads pointing to the center of the circle. The lice often select the postero-lateral areas of the abdomen; and, if two lice are present, they may be placed more or less symmetrically, with the heads directed forward and the remainder of the bodies backward, flattened against the integument of the fly. Possibly the lice are more commonly seen attached to the abdomen because in other positions their hold is so precarious that they usually drop off before being noticed. So many cases are now on record of biting lice being carried by bird-flies in all parts of the world, that this type of phoresy can no longer be regarded as abnormal, particularly in view of the fact that the Hippoboscidae are rarely collected in large numbers. Its significance and determining factors have been the subject of much speculation. The problem cannot be merely dismissed with the statement that the “Mallophaga which use Pupipara to go from one bird to another” are cases of “phoresy due to a trophic stimu- lus” (Yachon, 1941, p. 12), as will appear from the subjoined dis- cussion. 15 Throughout this account the term ‘ ‘ lice ’ ’ will refer only to “biting lice,” the generally accepted English name for Mallophaga, but which Hopkins (1949, p. 388) wishes to replace by “chewing lice.” There is no evidence that Anoplura ever attach to Hippo- boscidae. 163 ENTOMOLOGICA AMERICANA Observations by Clay and Meinertzhagen (1943), in England, seemed to lend some support to the view that lice might attach to a fly only after the bird host dies. They relate how on two occasions a single Ornithomyia fringillina (=0. lagopodis) , free of Mallo- phaga, was taken on a freshly killed Shetland starling, Sturnus vulgaris zetlandicus, in the Orkneys. In each case the fly was re- turned to the dead host, which was chloroformed an hour later. The dead flies were then recovered bearing specimens of Philopterus sturni attached to the abdomen, 3 females in one case, 7 males and 21 females in the other. A few other similar experiments gave negative results, however. The one positive result seemed to agree with Harrison’s (1913, p. 109) conclusion “that the mallophagous insects which find a hippoboscid upon the body of a dead host, fasten upon it, as its body temperature is above that of the defunct bird, without any intent, conscious or otherwise, of seeking trans- port to a new host. This would seem the more reasonable view to take, and it still allows the possibility of infection of a new host by parasites carried by a hippoboscid. ’ ’ Observations by bird-banders, who collect flies from living, healthy birds, do not support this ex- planation, however. Among 63 flies (46 Ornithomyia fringillina and 17 Ornithoica vicina) taken by B. S. Bowdish during the 1950 season, in New Jersey, on 50 birds belonging to 13 species, 5 carried Mallophaga (3 0. fringillina and 2 0. vicina). Ash (1952, p. 29), in England, likewise found that out of 108 Ornithomyia avicularia and 0. fringillina taken from live birds, 7, or 6.5 per cent, all 0. fringillina , carried lice. It is doubtful, therefore, that the main factor inducing lice to attach to a fly is associated with the death of the host or with the greater body heat of the fly, even though these may be sometimes added incentives. After the host dies, all ecto- parasites possibly tend to congregate in a few areas of the bird’s body which retain heat longer than others, thus increasing the chances of lice coming in contact with flies. Ewing (1927) offered some other possible explanations, none of them very satisfactory. He supposed, for instance, that bird-lice might attempt biting the integument of a fly in search of food, mistaking it for the bird’s skin; or that they might be induced to attach by some peculiar scent emanating from the fly, by some attractive secretion of the fly’s integument, or by the heat temporarily acquired by the fly’s body while in contact with the skin under the protective layer of feathers. These and other suggestions may well be worth investi- gating; meanwhile the instinctive or environmental stimuli induc- ing the lice to attach remain uncertain. 164 Volume XXXII The most attractive and purely teleological explanation of the phoresy of Hippoboscidae is, of course, that lice attach intention- ally to the flies in order to obtain ready transportation to a new host, which would imply instinctive foresight on the part of the louse. Most naturalists will find it difficult to admit such pur- poseful behavior, obviously not induced by physiological or mor- phological peculiarities of either louse or fly. It would seem ex- tremely doubtful that phoresy is now or could ever have been of sufficient benefit to the species of louse to have much survival value. It does not occur often enough to ensure transportation to a new host when needed, particularly in view of the very small proportion of birds of a given species harboring flies in the total bird popula- tion. The probability that a winged hippoboscid leaving a dead or live bird, away from the nest, might reach another bird suitable to the particular louse carried by the fly, is even smaller. As Mac- Arthur (1948) points out, most species of Mallophaga are narrowly restricted in the choice of a host, while the majority of Hippo- boscidae, and particularly most species of Ornithomyia and Or - nithoica, show very little host specificity. The chances of a given species of louse reaching the proper host by fly transport appear therefore to be remote. Nevertheless, Miss Clay (1949, p. 293) considers it possible that transference by hippoboscid flies is one of the normal ways in which the cuckoo, Cuculus canorus, obtains its louse population, as except during copulation, there is no contact between individual cuckoos. I have difficulty in accepting this suggestion. It should also be noted that very few of the numerous species of biting lice occurring on birds have been observed using this supposed roundabout method of survival, most of them man- aging well without it. All considered, it seems reasonable to regard the phoretic asso- ciations of lice and bird-flies as fortuitous, though rather frequent occurrences. Chance encounters of the parasites occur on the live bird when they seek out the same areas of the plumage, and on a dying bird when all parasites tend to congregate where more of the body heat is retained. It may be significant in this connection that most reported cases of phoresy are from flies on small or medium- sized birds, where different parasites have a better chance to meet. Possibly also the attachment of the louse to the fly is fortuitous, the louse merely grabbing the fly’s setae with the mandibles, as it is wont to do with the barbules of the feathers. There is no evidence that this type of phoresy is ever of any 165 ENTOMOLOGICA AMERICANA benefit to the hippoboscid, which merely tolerates it, because the lice appear to do little if any harm and the fly has no means of keeping them off: or removing them. There is also no proof that the lice obtain any nourishment from the flies. No doubt a hippoboscid leaving a bird with one or more of the host’s lice attached, reaches occasionally another bird, which will be usually of a different species. This might explain certain cases of so-called “straggling” or the occurrence in nature of species of Mallophaga on abnormal hosts. Recorded below is the capture on a vulture in Oregon of an Ornithomyia fringillina, normally not found on this type of bird, carrying a species of Brilelia characteris- tic of Fringillidae. Ash (1952, p. 29) reports a similar case in England of an Ornithomyia avicularia from a live great spotted woodpecker, Dryobates major, transporting Brilelia marginata, a specific louse of the European blackbird, Turdus merula. Miss Clay (1949, p. 293) discusses the possible role which phoresy by hippoboscids may have played in the speciation of the Mallophaga. Very few of the lice observed on louse-flies have been named as to species or even genera, mainly owing to the present unsatisfactory state of mallophagan taxonomy. Guimaraes (1944) reviewed criti- cally 20 cases in which a specific identification of the louse was at- tempted. He found that in 11 of them the lice belonged to species known to occur normally on the bird species carrying the fly. The identification of the louse was obviously erroneous in one case; while in 4 cases the bird on which the flies were taken is unknown. The following additional cases, all from the Old World, should be added to Guimaraes’ list. Eichler (1939, p. 223) reports Sturni- doecus sturni on Crataerina pallida from Apus apus, Philopterus pastoris on Ornitheza metallica from Pastor roseus, and Brilelia merulensis on Ornithomyia avicularia from Cuculus c. canorus. Clay and Meinertzhagen (1943) repeat 2 of Eichler ’s cases and add 3 more : Brilelia glandarii on Ornithomyia avicularia from Gar- rulus glandarius rufitergum, and Sturnidoecus sturni on Ornitho- myia fringillina (=0. lagopodis ) from & Humus vulgaris zetlandicus (on 2 birds). Eichler (1946) has one more case of Docophorulus lannii on Ornithomyia avicularia from Lanius collurio. Ansari (1947) records the interesting occurrence of Columbicola columbae on Pseudolynchia cgnariensis from the wild rock pigeon, Columba livia intermedia , in the Punjab. Phoresy of Brilelia glandarii by Ornithomyia avicularia on Garrulus g. glandarius was reported by Callot (1946) from France and by Buttiker (1949, p. 75) from Finland. Ash (1952, p. 29) cites Brilelia merulensis on Ornitho- 166 Volume XXXII myia avicularia from T urdus merula (on 3 birds; also recorded from this fly on the same host by Thompson, 19525, p. 38), and Bruelia marginata on the same fly from Dryobates major and T Ur- dus merula. Hopkins (1947, p. 170) mentions a species of Hohor- stiella carried by an undetermined hippoboscid from an unidentified dove, at Sterkfontein, Transvaal. It may be of some interest that, as Miss Clay (1949, p. 293) pointed out, all the Mallophaga found on hippoboscids and identified thus far, belong to the superfamily Ischnocera. The following summary of the known cases of phoresy of lice by American Hippoboscidae includes all published and some new records. Unless the contrary is stated, I have personally examined the flies of those previously published, correcting the identifica- tions if needed. A. Ornithoica vicina (Walker). This is the species generally called 0. confluenta in American literature (see Part II for a dis- cussion of the nomenclature). Although it is nearly as common on North American birds as Ornithomyia fringillina, it is more rarely taken carrying lice, probably because the fly’s small size makes it difficult for a louse to grasp it. 1 and 2. The first two cases of phoresy for 0. vicina were re- corded by Clay and Meinertzhagen (1943; as Ornithoica confluenta and Ornithomyia confluenta) , from California. The more precise localities are added here from my own records. A fly from Pasa- dena, on Aphelocoma c. calif ornica, carried a female louse of the genus Bruelia, attached to a vein of the wing.16 As shown in the authors’ illustration, one mandible of the louse was beneath the junction of the costa and first longitudinal, pressing the latter up- wards against the other mandible which had pierced the wing mem- brane from above. The second fly, from Altadena, on Pipilo macu- latus megalonyx, also carried a female Bruelia sp. fixed on a vein of the wing. 3 and 4. B. S. Bowdish collected the following two flies from living, trapped birds at Demarest, New Jersey, in 1950. A male fly on Turdus m. migratorius, August 6, had one louse attached. Another male fly on Cyanocitta c. crist at a, August 2, carried 3 lice, one fixed to the wing and the others to the abdomen. 5 and 6. Two flies on a “thrush,” taken by W. Clarke-Macln- tyre, at Abitagua, Ecuador, each carried a mallophagan. 16 The correct generic name for most of the biting lice formerly recorded as Degeeriella appears to be Bruelia. 167 ENTOMOLOGICA AMERICANA 7 to 9. Three flies from Nova Teutonia, Santa Catharina, Brazil, taken by F. Plaumann on Uroleuca cristatella, each carried a louse on the abdomen. 10. One fly from Nova Teutonia, Santa Catharina, Brazil, taken by P. Plaumann on Pyroderus scutatus, had a louse attached to the tip of the abdomen. B. Ornithomyia fringillina Curtis. By far the largest number of known North American cases, 42 out of total of 69, are from this species, the flies being taken on 18 species of birds (Passeriformes with 2 exceptions). Thirty of the cases I observed myself among some 500 0. fringillina seen from North America, about 6 per cent of the flies carrying lice, the normal percentage for this type of phoresy. In the Old World also the majority of the reported cases are from either of the two common species of Ornithomyia, 0. avicu- laria and 0. fringillina. The first observation of the kind, by C. Aube (1857) concerned an undetermined species of Ornithomyia, from a magpie, Pica p. pica, in Prance, which carried 2 “lice. ” Thompson (1947) noted that of 112 0. avicularia collected from trapped birds by one bird-bander in the same British locality, 11, or roughly 10 per cent, had lice attached. Ash (1952, p. 29), also in England, found that of 108 0. avicularia and 0. fringillina collected from trapped birds in one season, 7 0. avicularia, or 6.5 per cent, bore lice. The proportion was even lower in a collection of 46 0. fringillina taken from live trapped birds at Demarest, New Jersey, by Mr. B. S. Bowdish in 1950, only 2, or less than 5 per cent carrying lice. According to Edwards (1952), during the summer of 1951 phoresy was unusually common on Ornithomyia fringillina taken from live starlings, Sturnus vulgaris, at the Pair Isle Bird Observatory near the Shetlands, 17 of 78 flies, or 21.8 per cent, carrying lice ; it was, however, very rare on flies taken from wheat- ear, Oenanthe oenanthe (only on 1 of 205 flies), and not seen on flies from other birds. 1. N. Banks (1920) reported the first known case of phoresy of lice for an American hippoboscid. A male Ornithomyia, later re- ferred to 0. anchineuria (a synonym of 0. fringillina) by Johnson (1922), from Perisoreus canadensis, carried 2 lice, one on each side near the tip of the abdomen. (The supposed new species, Peri- soreus ~barhouri, was never described.) Thompson (1937) deter- mined the 2 lice as females of a species of Briielia. 2 and 3. W. L. McAtee (1922) reported 2 cases. One louse, later identified by Ewing (1927) as Briielia rotundata (Osborn), was attached by the mandibles to the upper right surface at the tip 168 Volume XXXII of the abdomen of a female fly from an unknown bird, collected at the mouth of the Macfarlane River, Lake Athabaska, Saskatchewan, by F. Harper, Aug. 11, 1920. Another specimen of Bruelia rotun- data was fixed in the same manner on a fly from Corvus brachy- rhynchos hesperis, at Ontario, Oregon. 4 and 5. H. E. Ewing (1927) added 2 more cases. A fly on Melospiza m. melodia, from Gates Mills, Ohio, carried a male and a female of Bruelia inter posit a (Kellogg), placed symmetrically at the sides near the tip of the abdomen. Another fly from the same locality, on Dumetella carolinensis , had a single Bruelia interposita. Ewing published a photograph of this case. He called the flies Ornithomyia avicularia ; although I have not seen them, there can be little doubt that they were the common North American species, 0. f ring illina. 6. G. J. Spencer (1928) found a fly on Cyanocitta s. stelleri, at Tofino, British Columbia, carrying 16 Bruelia deficiens (Piaget), attached by the mandibles to the abdominal tergites. Spencer records the fly as 0. avicularia ; although I have not seen it, I refer it to 0. fringillina. 7 and 8. G. B. Thompson (1936) recorded 2 female Bruelia sp. attached to the terminal segment of the abdomen of a male fly on Melospiza m. melodia , on Martha’s Vineyard, Massachusetts; also 2 females of Bruelia simplex (Kellogg) fixed to the dorsal sur- face of the abdomen of a female fly on Turdus m. migratorius, at Elmhurst, New York. 9 and 10. G. B. Thompson (1937) reported 2 more cases: a fly on Hylocichla u. ustulata, at Vancouver, British Columbia, car- rying 4 female and 1 immature Bruelia interposita (same speci- men recorded by G. J. Spencer, 1938) ; and a fly from Spruce Brook, Newfoundland, on Hylocichla f. fuscescens, with 1 female Bruelia sp. attached to the side of the abdomen. 11. C. M. Herman (1937) mentioned a fly from North East- ham, Massachusetts, on Molothrus a. ater, bearing 2 Mallophaga, one of which was later identified by G. B. Thompson (1947) as a species of Philopterus. 12. T. Clay and R. Meinertzhagen (1943) reported a fly from Groton, Massachusetts, on Turdus m. migratorius , bearing two female Philopterus sp., one attached to each side of the abdomen. 13. A specimen from Forest Hill Borough near Pittsburgh, Pennsylvania, on Hylocichla mustelina, carried a louse when it was taken, according to the collector, A. D. Kirk. 14 and 15. At Woodacre, Marin Co., California, J. Maillard 169 ENTOMOLOGICA AMERICANA collected the same day on Zonotrichia coronata, presumably on one bird, 4 flies, two of which carried lice : a male had 4 Brilelia sp. fixed on the dorsum of the abdomen, near the tip ; a female had 1 Brilelia sp. in the same position. 16 to 29. I have seen the following specimens carrying lice, all collected from living, trapped birds by B. S. Bowdish at Demarest, New Jersey. Male on Dumetella carolinensis, with 1 louse near the middle of the hind dorsal margin of the abdomen ; a photograph of this specimen, published by MacArthur (1948, p. 386, fig. 233), is reproduced in my Fig. 20. Male on Turdus m. migratorius, with 3 lice, 2 on the left and 1 on the right side of the abdomen. Female on Hylocichla mustelina, with 2 lice, one on each side, ventrally and close to the sides of the abdomen. Female on Turdus m. migra- torius, with 2 lice, one on each side of the abdomen. Female on Turdus m. migratorius, with 1 louse on the dorsal left side of the abdomen, close to the base. Female on Melospiza m. melodia, with 2 lice, one on each side margin, about mid-length of the abdomen. Female on juvenile Turdus m. migratorius, with 2 lice on the abdo- men, one on the left side near the base, the other on the right side near the apex. Female on Melospiza m. melodia, with 2 lice, one on each side margin of the abdomen. Female on Toxostoma rufum, with 2 lice on the dorsum of the abdomen, one on the left side at about mid-length, the other on the right side near the apex. Four females on three Cyanocitta c. cristata, each with 1 louse on the side of the abdomen. Female on Cyanocitta c. cristata, with 2 lice on the abdomen. 30. MacArthur (1948, p. 385) records a male from Turdus m. migratorius at Demarest, New Jersey (B. S. Bowdish), with 8 lice “clinging to the rounded lateral abdominal surface, arranged side by side, their heads all directed toward the center of the ab- domen. ’ 7 31 to 34. Mr. K. MacArthur sent me for publication the fol- lowing additional cases, not seen by myself, based on flies collected also by B. S. Bowdish at Demarest, New Jersey, from living, trapped birds. Male on Dumetella carolinensis, with 1 louse on the left side of the abdomen. Male on juvenile Turdus m. migratorius , with 7 lice on the abdomen, 3 dorsally, 3 ventrally and 1 on the right side. Female on Icterus galbula, with 1 louse on the right side of the ab- domen. Female on Hylocichla /. fuscescens, with 1 louse on the right side of the abdomen. 35. Female taken by H. A. Drew on Dryobates v. villosus, at 170 Volume XXXII Barre, Vermont, carried 1 louse on one side about mid-length on the dorsum of the abdomen. 36 and 37. At Oak Bluffs, Martha’s Vineyard, Massachusetts, a female on Melospiza m. melodia had 1 louse ( Bruelia sp.) fixed to the back of the abdomen; another fly on Spizella pusilla pas- serina also carried 1 louse of the same genus. 38. Male taken by A. E. Brower, July 29, 1932, on Mt. Desert Island, Maine, on Hylocichla guttata faxoni, had 2 lice, one fixed on each side near the tip of the abdomen. 39. Male taken by S. K. Carnie, October 12, 1950, at Berkeley, California, on Zonotrichia coronata, carried a louse on the side of the dorsum of the abdomen close to the basal laterotergite. 40. Female taken by Boy Wing at Euchre Mt., Lincoln Co., Oregon, on Cathartes aura teter, June 3, 1948, carried 4 Mallo- phaga at the tip of the abdomen, 3 on one side, 1 on the other. These lice were identified by Mr. K. L. Edwards as a species of Bruelia normally found on some fringillid passerine host. This seems to indicate that the Ornithomyia had strayed very recently from some passerine prey caught by the vulture, the latter not being known otherwise as a host of 0. fringillina. The case is also a definite example of a louse straying onto the wrong type of host following transport by a hippoboscid. 41. Female on Pipilo fuscus falcifer, taken at Berkeley, Cali- fornia, August 28, 1949, by H. E. Childs, Jr., carried a louse on each side of the abdomen, one before mid-length, the other close to the tip. 42. Female on Melospiza m. melodia , taken at Lake of Two Rivers, Algonquin Park, Ontario, August 20, 1950, by D. M. Davies, carried one louse on the right side edge, near the base of the ab- domen. C. Ornithomyia remota Walker (=0. paricella Speiser; 0. chilensis Reed). Although only some 50 flies of this species were seen, 6 of them carried Mallophaga, the proportion being higher than usual for 0. fringillina. 1. Inaccessible Island, Tristan da Cunha Group : female from Nesocichla eremita wilkinsi , collected by Y. Hagen, January 20, 1938, carried two lice fixed to the tip of the abdomen. 2 to 5. Valparaiso, Chile: four specimens, collected and re- ported by E. P. Reed (1932), carried lice which were determined by Ewing as a species of Bruelia. Male on Diuca d. diuca (= Diuca grisea), with one louse. Female cotype of 0. chilensis, from an unknown host, with 3 lice, one on the left and 2 on the right side 171 ENTOMOLOGICA AMERICANA of the abdomen. Female from an unknown host, with 2 lice, one on each ventral side of the abdomen. Female from an unknown host, with one louse at about mid-length on the right ventral side. 6. Maquehue, Temuco, Chile : female on Colaptes p. pitius, col- lected by R. M. Middleton, with one louse attached near the tip of the abdomen (case published by MacArthur, 1948, p. 385). D. Ornithoctona erythrocephala (Leach). Only 3 cases of phoresy are known thus far for this species, although it is common and widely distributed, with a variety of hosts. 1. A female taken on Momotus momota chlorolaemus by W. Weyrauch, at Huacapistana, Rio Tarma, Peru, carried one louse. 2. A female taken on Pionus sordidus saturatus by M. A. Car- riker, Jr., at Cincinnati, Sa. Marta, Colombia, August 29, 1945, carried on the abdomen no less than 15 lice, apparently all of one species ; 8 of them ( 7 adult and 1 immature ) were clustered around the apical anal plate ; the remainder were fixed to both sides near the base. The lice are a species of Paragoniocotes ( ? costalimai Guimaraes). 3. A female from Cuba, on Oreopeleia montana, bears a louse fixed by the mandibles to a hair, some distance from the integument, at the hind margin of the abdomen. E. Ornithoctona fuscivejitris (Wiedemann). Only one case known. 1. A female from Boquete, Chiriqui Province, Panama, on Psel- liophorus tibialis , carried 2 lice attached to the legs. F. Stilbometopa ramphastonis Ferris. Two cases are known thus far. 1. Guimaraes (1944) recorded and figured a fly on Trogonurus aurantius, from Rio S. Jose, State of Espirito Santo, Brazil, with 3 specimens of Bruelia odontopleuron (Guimaraes) fixed to the sides at the apical half of the abdomen. 2. A female taken on an undetermined toucan ( Pamphastos sp.), at Sto. Tomas, Izabal, Guatemala, carried one biting louse. G. Lynchia americana (Leach). Only one case of phoresy was ever recorded for this common North American fly. H. S. Peters (1935) reported upon a fly of this species from Hartstown, Penn- sylvania, on Casmerodius albus egretta, carrying 31 immature speci- mens of Ardeicola (reported as Esthiopterum) botauri (Osborn), a specific louse of certain wading birds. The lice were fixed to the thorax and abdomen, the fly appearing “to be covered with a white powder. ’ ’ This case presents several unusual features. It is note- worthy for the very heavy infestation with lice, all of which were, 172 Volume XXXII moreover, immature. In addition, the fly had strayed onto an ab- normal host, this being the only case to my knowledge of Lynchia americana having been taken on a wading bird. The fly was de- termined by me. H. Lynchia albipennis (Say). G. B. Thompson (1936) re- corded a specimen of this species (as L. botaurinorum) , from an unknown host, at Orizaba, Mexico, carrying 5 lice, ArdeicQla bo- tauri (Osborn). I have also seen this fly. I. Pseudolynchia canariensis (Macquart), the pigeon-fly, some- times called Lynchia maura Bigot, has been reported in the Old World (India) as carrying lice by Helen Adie (1915, p. 679) and Ansari (1947). Ansari ’s case was particularly interesting, as the fly came from a wild stock of the domestic pigeon, Columba livia intermedia. The following American cases refer to flies from do- mestic pigeons. 1 and 2. Margaret Martin (1934) observed two flies at Houston, Texas, carrying the pigeon louse, Columbicola columbae (Lin- naeus) : in one case with 3 lice (2J and lj'), in the other with one male louse. 3. Hathaway (1943) recorded and figured a fly from Rio de Janeiro, Brazil, carrying 7 specimens of Columbicola columbae. The cases of phoresy of Mallophaga known from the Old World concern the following species of flies: Ornithomyia avicularia (Lin- naeus), 0. fringillina Curtis (=0. lagopodis Sharp; 0. chloropus Bergroth), 0. roubaudi Seguy (probably a synonym of 0. fur Schiner), 0. perfuga Speiser, Crataerina pallida (Latreille), Ornithoica pusilla (Schiner), Ornitheza metallica (Schiner), and Pseudolynchia canariensis (Macquart). They were summarized by Thompson (1937; 1947), Clay and Meinertzhagen (1943), and Ansari (1947). A few more recent cases were reported by Hopkins (1947) from South Africa, and by Ash (1952, p. 29), Williamson (1950, p. 23), R. Edwards (1951; 1952) and Thompson (19525, p. 38) from England. Phoresy of Psocoptera. The following unique observation by Villeneuve (1913), at Rambouillet, France, may be of interest in view of the close relationship of the book-lice, or Psocoptera, to the Mallophaga. ‘ ‘ One autumn day I saw Lipoptena cervi running in some numbers from a freshly-killed roebuck [Capreolus capre- olus], brought to me damp and covered with mud. Most of them carried on the ventral side a small lengthened mass which puzzled me greatly. I have since learned from Mr. Speiser, that this mass 173 ENTOMOLOGICA AMERICANA was the dead body of a psocid of the genus Amphigerontia. I do not know how to interpret this observation. ’ ’ Phoresy of Thysanoptera. MacArthur (1948, p. 387) reports that a species of thrips was found on the abdomen of a male Or- nithomyia fringillina, taken on a live Toxostoma rtifum by Mr. B. S. Bowdish at Demarest, New Jersey. The occurrence was prob- ably a case of accidental transport; although it is known that cer- tain Thysanoptera are predacious, while others have been observed biting Man or animals. Phoresy of Pseudoscorpions. Seguy (1936, p. 117 ; 1939, p. DCCXXVI) states briefly that Crataerina pallida transports the small pseudoscorpion ( Chelifer ) that lives in the nests of “swal- lows.” As the author kindly informs me, this remark is based on a personal observation presumably made in the region of Fontaine- bleau, France. Although the pseudoscorpion is no longer avail- able for precise identification, it may have been the species re- ported, as Chelifer cancroides (Linnaeus), by Buttiker (1944, p. 35), who found it in Switzerland in the nest of a swift, Apus apus, the usual host of C. pallida. Miss Miriam Rothschild informs me (in lift., 1952) that she found C. cancroides in many nests of house martins and swallows in England. III. Reproduction The peculiar reproduction by integral or adenotrophic vivipar- ity (pupiparity), characteristic of the Hippoboscidae, dominates most activities of these flies and accounts for many features of their structure and physiology. Its origin and further evolution are puzzling problems, so far without satisfactory solution. Keilin (1916) emphasized the fundamental difference between the relatively simple ovoviviparity of many muscoid Diptera and the pupiparity of the Glossinidae, Nycteribiidae, Streblidae and Hippoboscidae. The first type of viviparity calls for no special method of feeding the offspring in the female’s uterus, since the larva develops mostly outside her body. As this type is found nor- mally or accidentally in several unrelated groups of insects, also outside the Order Diptera, it was induced presumably by a factor or combination of factors occurring rather frequently in nature. Yet it is by no means clear what this determining incentive may have been at the start ; nor is it certain that it was always the same. Compared With this, the origin of the incubating or adenotrophic type of viviparity of the pupiparous flies is an even more complex problem. It involves intricate adjustments of structure and func- 174 Volume XXXII tion insuring the full development of all larval instars within the uterus, where the larva is fed solely on the secretion of special milk-glands, until it is ready to pupate. As Keilin expresses it: “In order to achieve this, it is not enough to postulate an adapta- tion of the adult fly, in some way or other ; but it is also necessary to consider the specific and correlated adaptation of the newly-hatched larva, which, originally an independent, free organism undergoing its developmental cycle outside the mother’s body, must now be- come adapted to the new and very special conditions of intra-uterine life. The determinism of this mutual adaptation of larva and mother is no less complicated than that of a parasitic larva and its hosts. ’ ’ I agree with Keilin that no satisfactory explanation has as yet been offered for the origin of the several types of viviparity of the Diptera. Neither Portschinsky ’s surmise that all viviparous flies were derived from types with coprophagous larvae, nor Roubaud’s theory that pupiparity was induced by the combination of a very rich diet, namely blood, and a high temperature activating the gen- eral metabolism of the adult, suffice to account for it. Meanwhile, there may be some value in Keilin ’s suggestion that, in the Cyclor- rhapha at any rate, the larval adaptations required for successful viviparity possibly appeared first among flies with larvae parasitic upon the tissues of insects or vertebrates. But this leaves the ques- tion unanswered as to how the correlated adaptive changes of the female reproductive system actually started. Granted that such changes were initiated in the proper direction, their further de- velopment into the near-perfect condition now observed in the pupi- parous Diptera, would seem to be explained best by Roubaud’s theory. In particular, to be successful, integral viviparity re- quires, at least in free-living flies, a tropical or near-tropical en- vironment, such as is found nowadays in the intertropical zone and such as seemingly prevailed over much of the Earth’s surface dur- ing late Mesozoic and early Tertiary times. Sexual Dimorphism. Apart from the external terminalia, there are no striking secondary (C. H. T. Townsend’s tertiary) sexual characters in the Hippoboscidae. The sexes are often diffi- cult to distinguish, especially in pinned specimens where the shrivelled abdomen obscures the terminalia. The few differences are mostly in the number, size or shape of the abdominal tergal sclerites; sometimes also in the relative width of the interocular face, although the eyes are always broadly separated in both sexes, even at the postvertex; and occasionally in the chaetotaxy of the 175 ENTOMOLOGICA AMERICANA body and of the legs. Ornithoctona plicata (v. Olfers), of the Old World, is exceptional in the genus as well as in the family, in that the tip of the fore tibia of the female ends in a broad, rounded plate, which is replaced by a tuft of stiff setae in the male. As Ferris (1929) pointed out, sexual dimorphism is more pronounced in Ornithoica than usual, which is of interest because this genus is the most primitive of the family. In the female, the abdomen bears 5 large, transverse tergal plates, the 5th sometimes partly divided by a median constriction, followed by an apical pair of smaller sclerites. The male has only 4 median plates, followed by the apical pair. In addition, the hind trochanters of the male of most species bear one or more stout, short, spine-like setae, never present in the female. There is no convincing evidence that any of the few secondary sexual characters of the louse-flies are of special use in copulation or are connected with pre-mating behavior. The slight morpholog- ical differentiation of the Sexes 'and the simplified male terminalia may be correlated with permanent ectoparasitism, since both sexes have the same sedentary habits and similar requirements for shelter and food. They meet, therefore, without much difficulty, and sexual behavior is reduced to a minimum. In this connection it may be of interest that the free-living tsetse-flies ( Glossinidae ) , which are also blood-sucking and pupiparous, show a similar dearth of secondary sexual characters, but are nevertheless endowed with an extremely complex male copulatory apparatus. Sex Ratio. The exact proportion of the sexes in the normally reproducing population of an insect is difficult to determine. The best approach to the problem is to breed out numerous puparia produced under natural conditions, taking into account possible sexual differences in the pupal mortality. The few available data which come near fulfilling these requirements seem to show that in the Hippoboscidae both sexes are produced in about equal numbers. Coatney (1931), in Iowa, found that of 221 Pseudolynchia canarien- sis bred from puparia, 96, or 42.9 per cent, were males and 125, or 57 per cent, females. Schuurmans Stekhoven and his co-workers, in Argentina, obtained for the same species among 269 bred flies, 142, or 52.8 per cent, males and 127, or 47.2 per cent, females (un- published observations). The slight observed differences in the proportion of the sexes are cancelled out by these two experiments. Schuurmans Stekhoven (1923a, p. 29), in Indonesia, bred 968 flies from puparia of Hippobosca variegata (= H. maculata) , 473, or 48.8 per cent, being males and 495, or 51.2 per cent, females. Herman 176 Volume XXXII (1944, p. 117), in California, also obtained a sex ratio of approxi- mately 1 :1 from puparia of 8 'tilbometopa impressa reared in the laboratory : in 1941, of 69 bred flies, 39 were females and 30 males ; in 1942, of 19 flies, 8 were females and 11 males. Sex counts of adults collected at random on the hosts sometimes give similar results. Thus Schuurmans Stekhoven (1923a, p. 29), in Indonesia, found both sexes of Hippobosca variegata (= H. maculata ) in about equal numbers on horses: of 1562 caught flies, 804, or 51.4 per cent, were males and 658, or 48.6 per cent, females. More often, however, females tend to predominate on the hosts, because the males are either actually short-lived or more exposed to accidents. Sometimes males are rare, even in fairly large lots taken from a single host, particularly during the colder months, when reproduction may be slowed down or at a standstill. For instance, only one male was found among 29 dealated Lipoptena depressa taken from one black-tailed deer, Odocoileus hemionus columbianus , at Tilden Park, Berkeley, California, by Mr. I. B. Tarshis, January 14, 1949 (personal communication). G. J. Spencer (1938, p. 42) noted 51 females and 30 males in a lot of 81 L. depressa taken from one black-tailed deer in British Columbia. Hardenberg (1927, p. 13; 1929, p. 509) found 83 females and 46 males in a lot of L. cervi collected on deer in the Netherlands. The true sex ratio of Melophagus ovinus does not appear to have been determined by breeding puparia. W. C. Miller (1925), in England, stated that in random counts of adults on sheep fe- males outnumbered males about 2 to 1. Hardenberg (1927 ; 1929), in the Netherlands, found that 14 different lots taken from sheep contained in all 501 females and 338 males, but that males pre- dominated in some lots and that both sexes were equally represented in one. In three random collections made in Germany by Kemper (1951, p. 236), the females outnumbered the males in 2 cases and the males were more numerous in one ; the totals were 135, or 48.7 per cent, females and 142, or 51.3 per cent, males. Hardenberg (1927; 1929), in the Netherlands, bred 22 males and 20 females of Stenepteryx kirundinis from puparia found in nests of the house martin, Delichon u. urbica; while of 111 adult flies collected in the same nests 72 were females and only 39 males. Huzimatu (1938) obtained similar results for this species (which he calls S. nipponica) , on the black-chinned martin, Delichon urbica dasypus, in Japan: of 188 adults taken from 12 nests, 106 were females and 88 males. Kemper (1951, p. 235), in Germany, collected at the nesting 177 ENTOMOLOGICA AMERICANA sites of swifts, early in June, 54 female and 23 male Crataerina pallida; at the end of July and in early August, 43 females and only 9 males. He concluded that the females outlive the males, but granted that the numbers were too small to warrant definite con- clusions. The sex ratios of the common species of Ornithoica, Ornithomyia, and Ornithoctona have not been determined by breeding. In Ornithomyia the females usually outnumber the males on the hosts. Thompson (1940, p. 114) states that, in a series of 0. avicularia and 0. fringillina taken in England from 75 trapped live birds of 14 species, from June to September, 61.5 per cent were females and 38.5 per cent males. Ilis table shows that of 89 0. avicularia col- lected, 55 were females and 34 males, and of 36 0. fringillina , 28 were females and 8 males. The discrepancy was less pronounced among flies collected from trapped live birds in 1949-1950 by Ash (1952), in Berkshire, England: of 55 0. avicularia sexed, 47.3 per cent were males and 52.7 per cent females; of 53 0. fringillina, 60.4 per cent were males and 39.6 per cent females. Ash (1951) reported also a preponderance of females in Oeland, southern Sweden : 15 0. fringillina taken from 13 species of migrants trapped between August 27 and September 15, 1950, were all females ; while only one male was found among 16 0. avicularia from 3 species of birds; but the numbers are too small to be significant, owing to the scarcity of flies on migratory birds in that locality, since only 35 of the 1482 trapped birds carried flies. The disproportion of the sexes was slight in a lot of 0. fringillina collected by Mr. B. S. Bowdish from trapped birds at Demarest, New Jersey, between July 7 and October 29, 1950 : of 46 flies taken on 13 species of birds, 28 were females and 18 males ; in 1947, of 29 flies takeft on 11 species of trapped live birds, between July 29 and August 19, 15 were females and 14 males. According to Schuurmans Stekhoven and his co-workers (in lift., 1952), at Tucuman, Argentina, of 382 adult Pseudolynchia canariensis collected on domestic pigeons, 213, or 55.7 per cent, were males and 169, or 44.3 per cent, females. A series of 61 Lynchia americana from one Bubo v. virginianus near Ithaca, New York, consisted of 39 females and 22 males (J. Bequaert, 1945