LI B R.AFLY OF THE UNIVERSITY OF ILLINOIS. 595.2 W48e cop. 2 Biology // 7 /- / •'' L/'ljfc. Ol is book on or before the Date stamPed below. L161— Q-1096 ECTOPARASITES OF PANAMA ECTOPARASITES OF PANAMA RUPERT L. WENZEL VERNON J. TIPTON Editors FIELD MUSEUM OF NATURAL HISTORY CHICAGO, ILLINOIS NOVEMBER 22, 1966 Published with the assistance of Grants DA-MD-49-193-63-G73, G9211 United States Army Medical Research and Development Command As of March 1, 1966, Chicago Natural History Museum reas- sumed its former name, Field Museum of Natural History. As this volume was already in press, references to the museum remain unchanged throughout the text. Library of Congress Catalogue Card No. 64-25393 PRINTED IN THE UNITED STATES OF AMERICA 5-9 IT, . JL Contributors ROBERT M. ALTMAN, Ph. D., Lieutenant Colonel, Medical Service Corps, Entomology Consultant, Office of the Surgeon General, Department of the Army, Washington, D. C.; formerly Chief, Environmental Branch, Office of the Chief Surgeon, United States Army, Caribbean, Fort Amador, Canal Zone. ALFREDO BARRERA, Ph. D., Jefe, Laboratorio de Biologia Economica, Escuela Nacional de Ciencias Biologicas, Institute Politecnico Nacional, Mexico, D. F. JAMES M. BRENNAN, Ph. D., Research Entomologist (Medical), Department of Health, Education and Welfare, Public Health Service, National Institutes of Health, Na- tional Institute of Allergy and Infectious Diseases, Rocky Mountain Laboratory, Hamilton, Montana, and Middle America Research Unit, Canal Zone. K. C. EMERSON, Ph. D., Stillwater, Oklahoma; Research Associate, United States Na- tional Museum, Washington, D. C. GRAHAM B. FAIRCHILD, Ph. D., Medical Entomologist, Gorgas Memorial Laboratory, Panama, Panama. DEANE P. FURMAN, Ph. D., Professor of Parasitology and Chairman of Division, Uni- versity of California, Berkeley. LINDOLPHO R. GUIMARAES, Ph. D., Director (Retired), Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil. CHARLES O. HANDLEY, JR., Ph. D., Associate Curator, Mammals, Smithsonian Institution, Washington, D. C. PHILIP HERSHKOVITZ, M. S., Research Curator, Division of Mammals, Chicago Natural History Museum, Chicago, Illinois. PHYLLIS T. JOHNSON, Ph. D., Assistant Research Pathobiologist, Division of Biological Sciences, University of California at Irvine ; formerly with Gorgas Memorial Labora- tory, Panama, Panama. CHARLES M. KEENAN, B. G. S., Supervisory Biologist, Environmental Health Branch, Office of the Chief Surgeon, United States Army Forces Southern Command, Fort Amador, Canal Zone. :V>ALICJA KIEWLICZ, B.Sc., Assistant, Division of Insects, Chicago Natural History Museum, Chicago, Illinois. GLEN M. KOHLS, M. S., Sanitarian Director, Department of Health, Education and Wel- fare, Public Health Service, National Institutes of Health, National Institute of Al- lergy and Infectious Diseases, Rocky Mountain Laboratory, Hamilton, Montana. EUSTORGIO MENDEZ, M. S., Medical Entomologist, Gorgas Memorial Laboratory, Panama, Panama. HAROLD D. NEWSON, Ph. D., Lieutenant Colonel, Medical Service Corps, Chief, Entomol- ogy Research Section, United States Army Medical Research and Development Com- mand, Office of the Surgeon General, Department of the Army, Washington, D. C. FRANK J. RADOVSKY, Ph. D., Postgraduate Research Parasitologist, The George Williams Hooper Foundation, San Francisco, California ; formerly Acting Assistant Professor, Department of Parasitology and Entomology, University of California, Berkeley. RUSSELL W. STRANDTMANN, Ph. D., Professor of Biology, Texas Technological College, Lubbock. VERNON J. TIPTON, Ph. D., Lieutenant Colonel, Medical Service Corps; Chief, Entomology Branch, Medical Field Service School, Brooke Army Medical Center, Fort Sam Houston, Texas; formerly Chief, Environmental Health Branch, Office of the Chief Surgeon, United States Army Caribbean, Fort Amador, Canal Zone. RUPERT L. WENZEL, Ph. D., Curator, Division of Insects, Chicago Natural History Mu- seum, Chicago, Illinois; Lecturer, Department of Zoology, University of Chicago; Research Associate, Department of Biology, Northwestern University, Evanston, Illinois. CONRAD E. YUNKER, Ph. D., Senior Scientist, Department of Health, Education and Wel- fare, Public Health Service, National Institutes of Health, National Institute of Al- lergy and Infectious Diseases, Rocky Mountain Laboratory, Hamilton, Montana, and Middle America Research Unit, Canal Zone. " " " " ERRATA ECTOPARASITES OF PANAMA p. x, line 28 for "authorized" read: "authored" p. 200, line 28 for "singe" read: "single" p. 420, line 7 for "posterior" read: "proximal (anterior)" " " 10 " "anterior" " "distal (posterior)" 1 A. " " " " " 30-31 " "posterior" " "proximal (anterior)" p. 436, line 12 for "116" read: "96" " 13 " "43" " "22" p. 518, line 8 for "Almirante" read: "Changuinola" p. 528, 9 of text ' "setae in dunni" read: "setae. In dunni" p. 570, line 7 for "Playo" read: "Playa" p. 575, line 16 p. 598, lineS for "1" read: "6" p. 602, line 19 add: "Paratypes deposited in the collections listed on p. 410." p. 609, line 5 for "Guanote" read: "Guanota" 39 add: "Paratypes deposited in the collections of Field Museum of Natural History, the United States National Museum, Universidad Central de Venezuela, and the Environmental Health Branch, USAFSC, at Corozal (Canal Zone)." p. 620, last line delete: "11 (1 bat) Juan Mina (Canal Zone), 28 July 1960" p. 622, line 30 add: "Paratypes deposited in the collections listed on p. 410." p. 622, last line delete: " . From Artibeus lituratus" p. 623, line 1 for "2 August" read: "30 August" p. 698, lines 13, 14 for "50" and " ±300" read : "40" and " ±200" p. 709, lines 24, 25 for "III" and "IV", read: "IV and "III" p. 716, line 40 for "50" and "±300" read: "40" and "±200" p. 734, line 39 " " " " " " " p. 797, line 21 for "Coleopterg" read: "Coleoptera" Foreword The well-known difficulties presented by malaria and yellow fever dur- ing the construction of the Panama Canal served to focus the interest of medical entomologists upon this area of the world. The continuing presence of these and other arthropod-borne diseases has maintained this interest at a high level in the years following the completion of the canal. Much of the work in this interim period has been concerned with those aspects of tropical diseases that are directly associated with the attempts of humans to estab- lish and maintain a healthy environment in the tropics for themselves and their domestic animals. As more information is obtained concerning the natural ecology of arthropod-borne human and animal diseases in tropical areas, however, the need for a more complete understanding of host-parasite relationships becomes more apparent and the potential importance of both vertebrates and invertebrates as vectors or reservoirs of disease is ap- preciated more fully. The geographical location of the Panamanian Isthmus has made it a region of interchange between the fauna of North and South America as well as being a part of the area in which the autochthenous fauna of Middle America has developed. The climate, geological history, and varied physiography of Panama have combined to produce varied environments having a rather rich ectoparasite fauna in which unique and complex host- parasite relationships have developed. It is with these that the investiga- tions reported here are concerned. This volume presents the results of rather extensive studies concerning ectoparasites of the vertebrate fauna of Panama, and to a lesser degree, their hosts. While the results of these studies have indicated the variety of ectoparasites to be found in this rather limited geographical area and some- thing of the complexities of the host-parasite relationships that occur there, it is recognized that the information available for some of the groups is meager and incomplete. There still are remote areas of Panama in which the ectoparasite fauna remains virtually unstudied and it is quite certain that future work will result in additions being made to the species now known to be present. It is the authors' hope that this publication will be use- ful to those concerned with tropical environments and will stimulate other investigators to continue to work toward a more complete understanding of the many problems still remaining in these areas. HAROLD D. NEWSON Lieutenant Colonel, Medical Service Corps United States Army Medical Research and Development Command Washington, D. C. vii Preface This volume deals with major groups of ectoparasites of vertebrates, primarily those of mammals, in Panama. The Mallophaga of birds, the Analgesidae, and the smaller families of mites — such as the Myobiidae, Sarcoptidae, and Listrophoridae are not included. The Spelaeorhynchidae (bat mites) and Polyctenidae (bat bugs), although not treated in separate papers, have been identified to genus and included under their hosts in the comprehensive host parasite list near the end of the volume. Records that have been published for these in the past have also been included in the list. The contributors are aware that many described and undescribed species in the groups treated are yet to be collected in Panama and that many taxonomic problems are yet to be solved. This work should not be considered as a definitive treatise on the groups discussed, but rather as a starting point for the study of the complex taxonomic, biological, and epidemiological prob- lems that are encountered in this important area of the world. In editing the book, a number of unique problems arose, chiefly because of the diverse subject material and differing taxonomic formats used by the various contributors, the use made of a complete volume as contrasted with that of reprints, and because some of the papers had been prepared before the decision was made to incorporate them in one volume. Thus, consistency of treatment was not always possible. Acknowledgments The preparation of this volume has required the active participation and cooperation of so many agencies and individuals that it would be difficult to list them all here or to acknowledge and assess their individual contributions. The role played by many of them is acknowledged by Dr. Fairchild and others in the Introduction and elsewhere. The field investigations and study of the material collected were effected through the close collaboration of personnel of various federal agencies in Panama, notably the Gorgas Memorial Laboratory, the Middle America Re- search Unit (National Institutes of Health), the United States Army, and the Smithsonian Institution. The Army Transportation Corps provided much of the transportation for collecting trips, some of them to remote and virtually inaccessible areas. Some of the collaborating specialists not asso- ciated with the above agencies also participated in field investigations. Spe- ix PREFACE cial acknowledgment is due Mr. Charles M. Keenan of the Environmental Health Branch, United States Army Caribbean, and Dr. Charles 0. Handley, of the Smithsonian Institution, whose knowledge, field experience, and ener- gies contributed greatly to the success of the field program ; and Mr. Pedro Galindo of the Gorgas Memorial Laboratory whose assistance in expediting the field surveys was invaluable. The collaboration of Dr. Handley, who has concurrently been preparing a monograph on the mammals of Panama, was indispensable to many phases of the project. The encouragement and support of Colonel Robert Traub, Medical Service Corps (Retired) — formerly Chief, Entomology Research Section, United States Army Medical Research and Development Command, Office of the Surgeon General, Department of the Army — and of his successor, Lieutenant Colonel Harold D. Newson, Medical Service Corps, played a major role in the realization of the volume. Much of the field work of Lt. Col. Vernon J. Tipton, Charles M. Keenan, and Charles O. Handley was made possible through a research grant to Lt. Col. Tipton by the United States Army Medical Research and Development Command. Publication costs were assisted by similar grants to the Chicago Natural History Museum, with Rupert L. Wenzel as Principal Investigator and Lt. Col. Tipton as Principal Professional Assistant. The editorial work, too, required the active cooperation of many indi- viduals. Grateful acknowledgment is given the following : Dr. Graham B. Fairchild, who reviewed most of the manuscripts before submitting them to be prepared for publication ; various of the staff of the Division of Insects, Chicago Natural History Museum, including Miss Ella Fojtik, former secretary, who retyped major parts of the manuscripts sub- mitted for publication ; Mary Ryan Wenzel, who typed the index and all of the manuscripts authorized jointly by the editors; and Alicja Kiewlicz My- szkowski, technician, who carefully and painstakingly did a large part of the indexing ; and especially Christina Johnson Fowler who was responsible for much of the editing, proofreading, and make-up, and for the typographic de- sign of the book. She also assisted in other phases of the work, including the compilation of the comprehensive host-parasite list, a complex and difficult task. RUPERT L. WENZEL VERNON J. TIPTON Editors Contents FOREWARD Vii by Harold D. Newson PREFACE ix by Rupert L. Wenzel and Vernon J. Tipton INTRODUCTION 1 by Graham B. Fairchild GAZETTEER OF COLLECTING LOCALITIES IN PANAMA 9 by Graham B. Fairchild and Charles O. Handley, Jr. MITES OF THE SUBFAMILY LAELAPTINAE IN PANAMA (ACARINA: LAELAPTIDAE) 23 by Vernon J. Tipton, Robert M. Altman, and Charles M. Keenan THE DERMANYSSID MITES OF PANAMA (ACARINA: DERMANYSSIDAE) 83 by Conrad E. Yunker and Frank J. Radovsky THE GENUS Hirstionyssus FONSECA IN PANAMA (ACARINA: DERMANYSSIDAE) 105 by Russell W. Strandtmann and Conrad E. Yunker THE SPINTURNICID MITES OF PANAMA (ACARINA: SPINTURNICIDAE) 125 by Deane P. Furman THE TICKS OF PANAMA (ACARINA : IXODOIDEA) 167 by Graham B. Fairchild, Glen M. Kohls, and Vernon J. Tipton THE CHIGGERS OF PANAMA (ACARINA: TROMBICULIDAE) 221 by James M. Brennan and Conrad E. Yunker MALLOPHAGA OF THE MAMMALS OF PANAMA 267 by K. C. Emerson CHECKLIST OF THE SUCKING LICE OF PANAMA (ANOPLURA) 273 by Rupert L. Wenzel and Phyllis T. Johnson NEW SPECIES OF THE GENUS Amblyopinus SOLSKY FROM PANAMA AND MEXICO (COLEOPTERA: STAPHYLINIDAE) 281 by Alfredo Barrera THE FLEAS ( SIPHONAPTERA) OF PANAMA 289 by Vernon J. Tipton and Eustorgio Mendez xi CONTENTS CHECKLIST OF THE HIPPOBOSCIDAE OF PANAMA (DIPTERA: HIPPOBOSCIDAE) 387 by Graham B. Fairchild NYCTEBIBIID BATFLIES FROM PANAMA (DIPTERA: NYCTERIBIIDAE) 393 by Lindolpho R. Guimaraes THE STREBLID BATFLIES OF PANAMA (DIPTERA: STREBLIDAE) 405 by Rupert L. Wenzel, Vernon J. Tipton, and Alicja Kiewlicz SOME RELATIONSHIPS BETWEEN MAMMAL HOSTS AND THEIR ECTOPARASITES 677 by Rupert L. Wenzel and Vernon J. Tipton MICE, LAND BRIDGES AND LATIN AMERICAN FAUNAL INTERCHANGE 725 by Philip Hershkovitz CHECKLIST OF THE MAMMALS OF PANAMA 753 by Charles O. Handley, Jr. APPENDIX. CLASSIFIED LIST OF HOSTS AND PARASITES 797 INDEX . . . 825 xii Introduction GRAHAM B. FAiRCHiLD1 Study of the ectoparasites of vertebrates has been greatly intensified since World War II, chiefly because of an increased interest in zoonoses, diseases of animals communicable to man. Particular emphasis has been placed on the underdeveloped areas of the world, including large parts of tropical America, whose ectoparasite fauna is still poorly known. The ex- perience of medical entomologists has shown that taxonomic and ecological studies of animal reservoirs and vectors are essential to investigating the epidemiology of arthropod-borne diseases and must precede any organized effort to control them. The importance of ectoparasites in the epidemiology of such diseases is aptly illustrated by the role of fleas in the transmission of plague, of trombiculid mites in scrub typhus, of ticks in Rocky Mountain spotted fever. The attention of medical entomologists was first focused on Panama by the classical work with yellow fever and malaria during the building of the Canal. Through the control of these and other diseases, a safe and sanitary environment was rapidly established in the Canal Zone, and shortly thereafter in Panama. This enabled trained personnel to reside in the area for long periods of time and to conduct extended field and laboratory investigations of other arthropod-borne diseases and of the natural his- tory and resources of Panama. The providing of adequate facilities and of an atmosphere congenial to research at the Board of Health Laboratory at Ancon, under the leader- ship of research-minded Samuel Taylor Darling, directed the interest of several persons toward medical entomology. Notable among them was Lawrence H. Dunn, who produced the first work of any consequence on the ectoparasites of Panama (1916, et seq.). The initiation of biological surveys of the Canal Zone under the aus- pices of the Smithsonian Institution further stimulated interest in the 1 Gorgas Memorial Laboratory, Panama, Panama. 1 2 ECTOPARASITES OF PANAMA area and led to the publication of Goldman's Mammals of Panama in 1910 and Standley's Flora of the Panama Canal Zone in 1928. In this period, too, the volumes by Meek and Hildebrand on the fishes of Panama, both fresh water and marine, were published by the Field Museum (now the Chicago Natural History Museum) . In 1924, the opening of the field lab- oratory on Barro Colorado Island provided much-needed facilities for work in the area. Many of the scientists who worked on the island collected ectoparasites. In 1929, the Gorgas Memorial Laboratory was established as a research institute for tropical medicine. From its beginning, it was a center for studies in medical entomology; staff members published papers on ecto- parasites and furnished material for specialists elsewhere. In 1943, Fair- child listed most of the then-known ectoparasites of Panama (excluding the Mallophaga and the pupiparous Diptera) , a total of 63 species and subspecies belonging to 21 genera. He also gave a partial bibliography of relevant publications. In 1956, a systematic survey of the ectoparasite fauna of Panama was initiated by Major (now Lt. Colonel) Robert M. Altman. By 1959 a con- siderable backlog of information and unworked material relating to Pan- amanian ectoparasites had accumulated, both in Panama and in various collections elsewhere. The fortuitous circumstances which brought to- gether Drs. Conrad E. Yunker and James M. Brennan at the Middle America Research Unit, Major Vernon J. Tipton with the Army Environ- mental Health Branch, and Dr. Phyllis T. Johnson, Mr. Eustorgio Mendez, and Dr. Graham B. Fairchild at the Gorgas Memorial Laboratory, all in- terested in ectoparasites, proved mutually stimulating. These workers con- tinued the project begun by Major Altman. Field work by Dr. Alexander Wetmore on the birds of Panama and by Dr. Charles O. Handley on the mammals, together with field investigations of arthropod-borne virus diseases that were being carried out by Mr. Pedro Galindo of the Gorgas Memorial Laboratory, offered unrivaled opportunities to make extensive collections in remote areas that would have otherwise been difficult to reach. Thus, the ectoparasites of Panama have been more thoroughly collected than those of any other area of comparable size in trop- ical America. In the course of this survey, over 360 species of blood-sucking ectopara- sites, representing more than 120 genera were collected. Of these, 15 genera and more than 115 species were new. As a consequence it became necessary to enlist the aid of other specialists to study and report on the material collected. Because of the difficulties encountered in identifying the numerous species in diverse groups and the widely scattered literature on the subject, Major Vernon J. Tipton and Dr. Rupert L. Wenzel suggested the desir- ability of bringing together the papers resulting from these studies and, thus, to incorporate in one volume most of what is known about the ecto- parasites of a single area. It seemed particularly appropriate that this should be done for Panama because of its significance in the history of medical entomology. The enthusiastic cooperation of the participating FAIRCHILD: INTRODUCTION 3 specialists, the officials of the Chicago Natural History Museum, and the United States Army (Medical Research and Development Command, Office of the Surgeon General) made this possible. The fauna of Panama is of special interest, because the isthmian region is the only dry-land bridge between North and South America. Knowledge of the present Panama fauna is of great importance in understanding the movements and distribution of the animals of both continents. Further- more, the relatively small land area, with its great diversity of habitats and climatic zones, contains an unusually rich fauna of manageable size. It is believed, therefore, that the following papers, although primarily taxonomic in purpose, will prove of basic usefulness not only to those en- gaged in studies of zoonoses in this area, but also those with broader zoological interests. The Republic of Panama occupies the isthmus between North and South America. It is a roughly S-shaped area with its long axis running approximately east and west. It lies between 7° 09' and 9° 37' North lati- tude, and 77° 09' and 85° 01' West longitude, hence wholly within the trop- ics and about 1200 miles directly south of Miami, Florida. It is bounded on the west by the Republic of Costa Rica, on the north by the Caribbean sea, on the east by the Republic of Colombia, and on the south by the Pacific Ocean. It has a maximum east-west extension of about 475 miles, and a north-south extension of about 225 miles, but because of its shape no straight east-west line passes wholly over land and no point within the country is much more than 30 miles from salt water. The exact area is not known ; published figures vary from 28,000 to 33,000 square miles. Physically the country is dominated by a backbone range of moun- tains, the continental divide, consisting largely of igneous rocks and in- cluding a number of now extinct volcanos. This range is highest in west- ern Panama; elevations of over 11,000 feet are reached, the highest point being El Baru (Volcan Chiriqui), at 11,410 feet. Eastward, the divide drops to lower elevations ; the low point of 316 feet is reached at the Canal Zone, whence the range continues at around 2500 feet to near the Col- ombian border, where elevations of over 5000 feet are reached. For the most part the continental divide is closer to the Caribbean coast than to the Pacific, with the result that most of the agriculturally developed land is on the Pacific side, as are all but one of the major rivers. Geologically the underlying structure is complex, reflecting a history of repeated changes, although at the present time the land is comparatively stable, and no serious earthquakes have occurred in recent times. Sed- imentary rocks of tertiary age alternate with igneous intrusions and flows of lava and beds of ash. There is much folding and faulting. The soils are in general rather acid clays of low fertility, although areas with com- paratively good soils do exist, especially in Chiriqui Province and the low- lands of Darien and Bocas del Toro Provinces. There are no large bodies of fresh water except for the artificial Gatun and Madden Lakes in the Canal Zone. Except for the lower reaches of the Tuira in Darien Province, rivers are rapid and shallow and not navigable except by dugout canoe and similar craft. There are extensive freshwater and tidal coastal swamps 4 ECTOPARASITES OF PANAMA on both coasts, especially at the mouths of the numerous rivers. On the Pacific coast, notably within the Gulf of Panama, the tidal range is great, reaching 18 feet or more, while on the Caribbean coast it is barely two feet. Much of the coast line is rocky and precipitous or fringed with man- grove swamp and mud flats, though sand beaches occur in favorable local- ities on both coasts. Originally most of the country was covered with forest, except for some drier areas along the Pacific coast which appear to have been grassland since prehistoric times. Man's agricultural activities in the past few cen- turies have changed much of this, especially on the more densely populated Pacific coastal plain, so that forest in this area is reduced to scrub on steep hillsides and narrow gallery forest along streams. The predominant and somewhat primitive agricultural practice consists of cutting and burning forest in the dry season to plant a crop in the ashes. A new patch of forest is destroyed each year, so that virgin forest in all accessible areas is rapidly disappearing. Repeated burnings prevent reforestation. This results in increasing areas of grassland, unsuitable for further agricultural use and with very depauperate fauna. Panama has a hot, tropical climate which is somewhat tempered by the proximity of the sea. Temperatures at sea level in the Canal Zone area seldom go below 70°F. or above 90°F. though extremes of about 60° and 96° are sometimes encountered. Information is scanty for many areas, but in general extremes are slightly greater on the drier Pacific coast than on the Caribbean, and lower temperatures prevail at higher elevations. The diurnal temperature range is not great, seldom more than 10°F. Hot nights are rare. The year is divided into two climatic seasons, wet and dry, which are much more pronounced on the Pacific side than on the Atlantic. The wet or rainy season, known locally as invierno, or winter, generally extends from about the middle of April to the middle of December, al- though it varies greatly in duration and intensity locally and from year to year. During the rainy season, rain is to be expected on any day, most often in the afternoon and generally in the form of heavy local thunderstorms. Occasionally there are widespread rains of long duration, especially from October to December. Rainfall may be exceedingly heavy, up to 2.48 inches in 5 minutes and over 10 inches in a single 24-hour period. Annual average rainfall varies from year to year and place to place, but it is generally heaviest along the Caribbean coast on the mountain slopes and least along the Pacific coast west of the Canal Zone. As much as 247 inches in a single year have been recorded from Porto Bello on the Caribbean and as little as 25 inches at Naos Island at the Pacific entrance of the Canal. Greater rainfall probably occurs in some mountain areas. Relative humidity is generally high, seldom below 60 percent during the day and almost always above 90 percent at night. During the dry season, the lo- cal verano or summer, there may be no rain, at least on the Pacific side, for as much as four months, and seldom is more than an inch a month recorded for the months from January through March. The Caribbean slope and moun- tains usually receive occasional showers, and in the wetter areas there is no real dry season. FAIRCHILD: INTRODUCTION 5 Winds are seldom strong, and windspeeds of over 30 m.p.h. are excep- tional. Panama is outside the hurricane belt. During the rainy season light and variable winds from the south are common or on occasion there may be strong winds of short duration, accompanying thunderstorms. During the dry season, the North East tradewinds may blow fairly steadily from January through March. Politically the country is divided into nine provinces and the Intendencia of San Bias. Of these, Bocas del Toro, Colon, and San Bias lie wholly on the Caribbean coast; Veraguas fronts on both coasts, and the remaining provinces are wholly on the Pacific side of the continental divide. The Canal Zone is a strip about 10 miles wide from coast to coast dividing Colon and Panama provinces. Detailed delineation of life zones in Panama has hardly begun. Gold- man in his Mammals of Panama (1920) discusses the problem and gives a provisional map. Standley in his Flora of the Panama Canal Zone (1928) devotes several pages to general descriptions of the plant associations in the Canal Zone. Holdridge and Budowski in a recent report (1955) discuss life zones in more detail, giving a map showing four life zones : Tropical, up to 600 meters altitude on the Atlantic coast, to 700 meters on the Pacific ; Subtropical, from 600 or 700 to 1500 meters ; Lower Montane, up to 2600 meters and Montane, over 2600 meters. These zones are further divided into dry, wet, and transition divisions. In general, the Tropical zone com- prises 76 percent of the republic, the Subtropical about 18 percent, the Lower Montane and Montane together about 5 percent, the last occurring only on the highest mountains in Chiriqui Province. The classification is based largely on considerations of temperature, precipitation and forest asso- ciations. The mountainous nature of much of the country results in limited areas of high rainfall with associated rain shadows. This, coupled with edaphic and underlying geological features of circumscribed extent, cause abrupt changes in the vegetation cover over relatively short distances and render general statements subject to numerous exceptions. Thus, relatively low hills within a few miles of each other may have quite different vegetation and associated faunas. The same mountain may be clothed in grass and xero- phytic scrub on its leeward side and dense fog forest on the windward side with practically no zone of transition. For present purposes, and until much more detailed information is available, the country may be divided into three zones : below 1000 feet ; from 1000 to 5000 feet ; and over 5000 feet. These correspond very roughly to the Tropical, Subtropical and combined Submontane and Montane zones of Holdridge and Budowski. The collections reported in this group of papers have come very largely from elevations below 5000 feet, with some rep- resentation from areas in Chiriqui Province from above 10,000 ft. The ac- companying map indicates these three zones. A few collecting localities that are representative of these zones are described below. Cerro Punta (Chiriqui) is a small village located on the slopes of Volcan Baru at an elevation of about 6000 feet. Extensive collections of mammals and ectoparasites were made in this vicinity (Bambito, Finca Lara, Casa 6 ECTOPARASITES OF PANAMA Tilley, Finca Martinz) at elevations of from 4800 feet to over 8000 feet. Rainfall is moderately heavy, and the dry season is distinct though not intense. The heavy forest is representative of the humid upper Tropical Zone, with oaks and bamboo occurring at the higher levels. The area is rapidly being cleared for coffee and vegetable crops. Finca Lerida (Chiriqui) is a coffee farm between Boquete and Cerro Punta on the slopes of Volcan Baru at an elevation of between 5000 and 6000 feet. An area of virgin forest near the farm was preserved until 1955, but most of the adjacent area is now used for coffee and vegetable crops and for pasture. The former owner and his wife, Mr. and Mrs. T. B. Monniche, were interested in natural history, collected birds, and were hosts to many visiting naturalists, a number of whom collected ectoparasites. Almirante (Bocas del Toro) is the headquarters town for large banana and cacao plantations operated by the Chiriqui Land Company. Extensive collections were made from August 1951 through May 1953 at a camp es- tablished about 12 miles southwest of the town for the study of sylvan yellow fever. From 1959 to 1962, collections were made at several other localities within a few miles of the town. All collecting localities are below 500 feet, and most are lower. They include virgin forest as well as swamp forest, second growth, and land under cacao and banana cultivation. Pre- cipitation is high throughout the area. There is no pronounced dry season and no month regularly has less than three inches of rainfall. The town is situated on the Chiriqui Lagoon. Isla Bastimentos and Cayo Agua (Water Key) are islands in the Lagoon. Rio Changena, lower camp (Bocas del Toro) was a temporary camp on an eastern tributary of the Rio Changuinola, about 20 miles inland, at an elevation of 2400 feet. The area is one of heavy tropical forest with continu- ous high rainfall throughout the year. Collections were made during Sep- tember 1961, within a radius of two miles of the camp, and up to eleva- tions of 3000 feet. Rain fell on more than half the days during this time. The ground was continuously wet, in degrees ranging from moist to sat- urated. Rio Changena, upper camp, or Rancho Mojica (Bocas del Toro), a small coffee farm, is located about 10 miles from the continental divide on the Atlantic side, at an elevation of 4800 feet. The collecting area is a tropical rain forest surrounding a four or five acre plot, once cleared but now sup- porting some secondary growth. A stream runs near the lower fringe of the clearing. Collections were made along the stream and along a trail on a high ridge ranging in elevation from 5000 to 5650 feet. The climate during the collecting period seemed drier than that at the lower camp. Cerro Hoya (Los Santos) is a mountain area behind Las Palmitas. The higher elevations are heavily forested and appear to have abundant rain- fall. Mammal and ectoparasite collections were made here from 11 January 1962 to 2 March 1962 at elevations between about 1000 and 3000 feet. Las Palmitas (Los Santos) is a small settlement at the foot of Cerro Hoya, near the Pacific coast. The area is relatively dry, with an intense dry season. The land has been long under cultivation and little of the original vegetation remains. FAIRCHILD: INTRODUCTION 7 Canal Zone. Numerous collections, all at low elevations, have been made on both sides of the isthmus in the Zone. Because of restrictions on agricultural use of land within the Zone, there are many areas of nearly un- disturbed forest. On the Caribbean coast, in the Camp Pina and Mohinga Swamp areas, there are patches of virgin rain forest. Barro Colorado Is- land and Madden Forest provide successively drier habitats. In general, the rainfall pattern in this area is modified by the low elevation. The Caribbean coast is somewhat drier than adjacent areas and the Pacific coast has a less intense dry season than areas farther west. Considerable trapping was done in and about townsites and military installations, in small patches of second growth forest, scrub and grassland. Many collections have come from animals found dead on highways and from bats taken in abandoned buildings, road culverts, and old mine tunnels. Cerro Pirre or upper Rio Seteganti (Darien) is the location of a tempo- rary camp, used during January and February 1961. It was situated on the western edge of a sloping valley or plateau, between the Cerro Pirre range and Cerro Setetule, one mile south of the Rio Seteganti and 26 air miles to the south of El Real. The elevation of this camp was about 1500 feet, in an area of tropical rain forest broken by occasional marshy meadows and fields of wild cane. Collections were made in the valley itself and along a long, narrow, gently sloping spur, perpendicular to the Cerro Pirre range, up to an elevation of 3500 feet. Except for occasional trails of rubber col- lectors leading toward the nearby Colombian border, there is no evidence of human habitation of this area since the nearby Cana mines were closed about 45 years ago. Weather data for this area is fragmentary. Judging from the dearth of deciduous trees and the abundance of surface water noted during the dry season, this area is more like the modified rain forest of the Atlantic Coast than the tropical wet and dry or Savanna Forest of the Pacific slopes. References FAIRCHILD, G. B. 1943. An annotated list of the bloodsucking insects, ticks and mites known from Panama. Amer. Jour. Trop. Med., 23, (6), pp. 569-591. GOLDMAN, E. A. 1920. Mammals of Panama. Smiths. Misc. Coll., 69, (5) , (Publ. 2498) , pp. 1-309, pis. 1-39, text figs. 1-24. HOLDRIDGE, L. R., AND BUDOWSKI, G. 1955. Report of an ecological survey of the Republic of Panama. (Technical Coop- eration program of the I.A.I.A.S. of the O.A.S.). Unpublished. STANDLEY, P. C. 1928. Flora of the Panama Canal Zone. Contrib. U. S. Nat. Herb., 27, pp. i-x, 1-416, pis. 1-66, text figs. 1-7. Gazetteer of Collecting Localities in Panama GRAHAM B. FAIRCHILD l and CHARLES 0. HANDLEY, JR.2 The Panamanian collecting localities mentioned in this volume are in- dexed in this gazetteer. Section I is an index to province of all collecting lo- calities. In Section II, the localities are arranged by province and have been identified as precisely as possible, usually to the nearest minute of latitude and longitude.3 However, it should be realized that the exact collecting sites in many instances may be miles away from the center of the town, or summit of the mountain, or bank of the stream indexed. Names of mountains, rivers, ranches, forts, islands, etc., are to be found under the specific name. For example, Cerro Mali will be found under Mali, Cerro, and Rio Chagres under Chagres, Rio. Elevations in feet above sea level have been given for all collecting sites above 1000 feet. In this instance the figures relate as nearly as possible to the elevation of actual collecting sites rather than to the elevation of the town or mountain indexed. To facilitate rapid discovery of the general position of collecting locali- ties, all localities have been related to 50 key localities which are identified by number in the gazetteer and on the accompanying map. This map and the gazetteer should be used in conjunction with more detailed maps, pref- erably the one used as a basis for determining the coordinates (see foot- note) or the Millionth Map of Hispanic America (American Geographical Society, Publication No. 5). The Road Map of Canal Zone and Vicinity, prepared by the Engineer Service (Corozal, Canal Zone), United States Army Caribbean, is also very useful, though much less detailed. The gazetteer was compiled after most other portions of the volume were in galley proof. Consequently it is possible that some of the errors in spell- ing or altitude that existed in the systematic reports may not have been corrected. If any are discovered, or if additional localities can be more pre- cisely identified, information on them will be received with thanks by the compilers and the editors. 1 Gorgas Memorial Laboratory, Panama, Panama. - Smithsonian Institution, Washington, D. C. 3 The three part map of the Republic of Panama, scale 1:500,000, produced by USARCARIB, was used as a base for the determination of all coordinates. 10 ECTOPARASITES OF PANAMA Section I. Index of Localities to Province Achiote, Colon Afuera, Isla, Veraguas Agua, Cayo, Bocas del Toro Aguadulce (15), Cocle Aguas Buenas, Panama Albrook Field, Canal Zone Alhajuela, Canal Zone Almijas, Isla, Chiriqui Almirante (2), Bocas del Toro Altos Cacao, Veraguas Amador, Fort, Canal Zone Amagal, Darien Ancon, Canal Zone Anton, Cocle Armila (49), San Bias Arraijan, Panama Aruza, Darien Aspinwall, Colon Avaso, Rio, Panama Azuero, Peninsula de, Herrera, Los Santos, Veraguas Azul, Cerro (37), Panama Balboa, Canal Zone Bambito, Chiriqui Barro Colorado Island (29), Canal Zone Baru, Cerro, Chiriqui Bas Obispo, Canal Zone Bastimentos, Isla, Bocas del Toro Bayano, Rio, Panama Bejuco, Panama Boca de Cupe, Darien Boca del Drago, Bocas del Toro Bogavo, Chiriqui Bohio, Canal Zone Bonita, Quebrada, Colon Boqueron, Chiriqui Boquete (5), Chiriqui Boquete-Volcan Trail, Chiriqui Boracho, Loma, Canal Zone Borinquen Road, Canal Zone Brava, Isla, Chiriqui Bruja, Cerro, Colon Bubi, Rio, Veraguas Buena Vista, Colon Bugaba, Chiriqui Burica, Punta, Chiriqui Butz, Finca, Chiriqui Caballero, Rancho, Bocas del Toro Cabima, Panama Cacao Plantation, Canal Zone Calidonia, Panama Calobre, Veraguas Calovebora, Veraguas Calzada Larga (26), Panama Camoganti, Darien Campana, Cerro, Panama Cana, Darien Cana, Loma, Darien Canal de Afuera, Isla, Veraguas Candela, Rio, Chiriqui Candelaria Hydrographic Station, Panama Cangandi, Rio, San Bias Capeti, Darien Capina, Herrera Capira, Panama Carasquilla, Panama Cardenas, Canal Zone Casa Larga (26), Panama Casaya, Rio, Canal Zone Casita, Darien Casita Alta, Chiriqui Cativa (or Catival), Colon Cativo, Panama Cebaco, Isla, Veraguas Cement Plant, Colon Cerro Punta (4), Chiriqui Cerro Punta-Boquete Trail, Chiriqui Chagres, Camp, Canal Zone Chagres, Rio, Canal Zone Chame (19), Panama Changena, Rio (3), Bocas del Toro Changuinola, Bocas del Toro Changuinola, Rio, Bocas del Toro Chapera, Isla, Panama Charco del Toro, Panama Chepigana, Darien Chepo (40), Panama Chico, Canal Zone Chilibre, Panama Chilibrillo Caves, Panama Chiman (42), Panama Chiriqui, Volcan de, Chiriqui Chiriqui Viejo, Rio, Chiriqui Chiriquicito, Bocas del Toro Chiva Chiva, Canal Zone Chucunaque, Rio, Darien Cituro, Darien Clayton, Fort (24), Canal Zone Coco Solo, Canal Zone Cocoli, Canal Zone Cocos, Punta, Panama Coiba, Isla (9), Veraguas 4 Section I is an index to province of all collecting localities. In Section II, the localities are arranged by province. FAIRCHILD AND HANDLEY : GAZETTEER 11 Colon, Colon Colon, Isla, Bocas del Toro Colorado, Rio, Chiriqui Concepcion (6), Chiriqui Corozal, Canal Zone Corte Culebra Road, Canal Zone Cotito, Rio, Chiriqui Goto Region, Chiriqui Cristobal, Canal Zone Culebra, Canal Zone Curundu, Canal Zone Cylindro, Bocas del Toro Davala, Chiriqui David (7), Chiriqui Davis, Fort, Canal Zone Divala, Chiriqui Divisa, Herrera Donoso, Colon El Banco, Chiriqui El Baru, Chiriqui El Cope, Code El Hato del Volcan, Chiriqui El Limon, Colon El Potrero, Code El Real (43), Darien El Valle (18), Code El Vijia, Canal Zone El Volcan, Chiriqui Emperador, Canal Zone Empire, Canal Zone Escobal (30), Colon Escudo de Veraguas, Isla (1), Bocas del Toro Esnape, Rio (48), Darien Farfan, Canal Zone France Field, Canal Zone Frijoles, Canal Zone Frijolito, Rio, Colon Galeta Island, Canal Zone Galeta Point, Canal Zone Gamboa (28), Canal Zone Gariche, Rio, Chiriqui Gatun, Canal Zone Gobernadora, Isla, Veraguas Goofy Lake, Panama Guanico, Los Santos Guayabalito, Colon Guayabito, Panama Guayabo, Darien Gulick, Fort (33), Canal Zone Howard Field, Canal Zone Hoya, Cerro (12), Los Santos Huile, Panama Indio, Rio, Canal Zone Insolita, Isla, Chiriqui Jaque (47), Darien Jefe, Cerro, Panama Jesucito, Rio, Darien Juan Diaz, Panama Juan Mina, Canal Zone Kobbe, Fort (22), Canal Zone K-9 Road, Canal Zone K-10 Road, Canal Zone La Boca, Canal Zone La Chorrera (20), Panama La Concepcion, Chiriqui La Laguna, Darien La Palma, Darien La Vaca, Rio de, Chiriqui La Zumbadora, Panama Lagartera, Canal Zone Lara, Finca, Chiriqui Las Cascadas, Canal Zone Las Cruces, Canal Zone Las Cruces Trail, Canal Zone Las Cumbres, Panama Las Palmitas, Los Santos Lava Flow, Chiriqui Lerida, Finca, Chiriqui Lewis, Casa, Chiriqui Limon, Rio, Darien Lion Hill, Canal Zone Llano Verde, Chiriqui Madden Airstrip, Panama Madden Dam (27), Canal Zone Madden Forest (25), Canal Zone Madden Road, Canal Zone Madden Wye, Canal Zone Maje, Rio, Panama Mali, Cerro, Darien Mamoni, Rio, Panama Mandinga (38), San Bias Mandinga, Rio, Canal Zone Margarita, Canal Zone Mariato, Rio, Veraguas Marraganti, Darien Martinz, Finca, Chiriqui Martinz Dairy, Chiriqui Maxon Ranch, Panama Miraflores, Canal Zone Mohinga Valley, Canal Zone Moja Polla, Rio, Panama Mojica, Rancho, Bocas del Toro Mojinga, Rio, Canal Zone Mono, Rio, Darien Monte Obscuro, Panama Mount Hope, Canal Zone 12 ECTOPARASITES OF PANAMA Nueva Colonia, Chiriqui Nueva Gorgona, Panama Nuevo Emperador, Panama * Nuevo Limon (34), Colon Orchid Island, Canal Zone Pacheca, Isla, Panama Pacora (39), Panama Paitilla, Punta, Panama Paja, Panama Palenque, Colon Palo Santo, Chiriqui Panama (23), Panama Panama Viejo, Panama Pando, Cerro, Chiriqui Paracote, Veraguas Paraiso, Canal Zone Parida, Isla, Chiriqui Parita (14), Herrera Paya, Boca de Rio (44), Darien Paya Village, Darien Pearl Islands (41), Panama Pedasi (13), Los Santos Pedregal, Chiriqui Pedro Gonzales, Isla, Panama Pedro Miguel, Canal Zone Pelado, Cerro, Canal Zone Pelisa, Darien Peluca, Rio, Panama Pena, Punta de, Bocas del Toro Penonome (16), Code Pequeni, Rio (36), Panama Perlas, Archipielago de las (41), Panama Pese, Herrera Pina (31), Colon Pina, Camp, Canal Zone Pina, Punta, Darien Pinogana, Darien Pirre, Cerro, Darien Pital, Camp (6a), Chiriqui Pito, Rio, San Bias Porcada, Isla, Chiriqui Portobelo (or Porto Bello) (35), Colon Potuga, Herrera Prominente, Cerro, Panama Pucro, Rio, Darien Pueblo Nuevo, Chiriqui Pueblo Nuevo, Panama Puente, Rio, Panama Puerto Limon, Colon Puerto Obaldia, San Bias Puma Island, Canal Zone Punusa, Boca de Rio, Darien Quarry Heights, Canal Zone Randolph, Fort, Canal Zone Real de Santa Maria, Darien Red Tank, Canal Zone Remedies (8), Chiriqui Represo, Canal Zone Rey, Isla del, Panama Rio Aba jo, Panama Rio Chico Hydrographic Station, Panama Rio Hato (17), Code Risco, Boca de Rio, Bocas del Toro Rodman Naval Station, Canal Zone Sabanas, Panama Saboga, Isla, Panama Salamanca Hydrographic Station, Canal Zone Salto de Madrono, Panama Salud, Colon San Felix, Chiriqui San Francisco de la Caleta, Panama San Jose, Isla, Panama San Juan, Canal Zone San Lorenzo, Fort, Canal Zone San Miguel, Panama San Miguel, Isla, Panama San Pablo, Canal Zone Santa Clara, Chiriqui Santa Clara, Code Santa Clara, Quebrada, Chiriqui Santa Clara, Rio, Chiriqui Santa Cruz de Cana, Darien Santa Fe (11), Veraguas Santa Rosa, Colon Santiago (10), Veraguas Sapo, Cerro, Darien Sereno, Chiriqui Seteganti, Rio (46), Darien Sevilla, Isla, Chiriqui Sherman, Fort (32), Canal Zone Sibube, Bocas del Toro Siolo (or Siola), Chiriqui Sona, Veraguas Summit, Canal Zone Tabernilla, Canal Zone Taboga, Isla (21 ), Panama Taboguilla, Isla, Panama * Brennan and Yunker (see The Chiggers of Panama, elsewhere in this volume) refer collections from Nuevo Emperador to the Canal Zone. These collections were made in the Canal Zone near Nuevo Emperador. FAIRCHILD AND HANDLEY : GAZETTEER 13 Tacarcuna, Cerro, Darien Trinidad, Rio, Panama Tacarcuna Casita, Darien Tuira, Rio, Darien Tacarcuna Laguna, Darien Tacarcuna Village (45), Darien Venado Beach, Canal Zone Tacarcuna Yellow Fever Station, Darien Vieja, Punta, Bocas del Toro Tapalisa, Darien Viejo, Cerro, Veraguas Tapia, Panama Viejo, Rio, Veraguas Terebe, Rio, Bocas del Toro Vijia, Canal Zone Teribe, Rio, Bocas del Toro Villa Rosario, Panama Tigre, Cerro, Canal Zone Vique, Punta, Panama Tilley, Casa, Chiriqui Timi de Boa, Rio Teribe, Bocas del Toro Wald, Chiriqui Timishik, upper Rio Teribe, Bocas del Toro Tocumen, Panama Yaviza, Darien Section II. Index to Localities by Province BOCAS DEL TORO Agua, Cayo, 9°10'N-82°02'W (near Almirante-2) Almirante (2), 9°18'N-82°24'W Bastimentos, Isla, 9°19'N-82°08'W (near Almirante-2) Boca del Drago, 9°26'N-82°20'W (near Almirante-2) Caballero, Rancho, near 9°02'N-82°41'W (near Rio Changena-3), 5000 feet Changena, Rio (3), 9°06'N-82°34'W, 2300-2600 feet Changuinola, 9°27'N-82°31'W (near Almirante-2) Changuinola, Rio, 9°22'N-82°31'W (near Almirante-2) Chiriquicito, 8°57'N-82°10'W (near Almirante-2) Colon, Isla, 9°24'N-82°16'W (near Almirante-2) Cylindro (NE of Boquete on upper Caribbean slope, near Boquete-5), above 4000 feet Escudo de Veraguas, Isla (1), 9°06'N-81°33'W Mojica, Rancho, near 9°02'N-82°41'W (near Rio Changena-3), 4800-5600 feet Pena, Punta de (a point on Laguna de Chiriqui?) Risco, Boca de Rio, 9°16'N-82°28'W (near Almirante-2) Sibube, 9°36'N-82°47'W (near Almirante-2) Terebe, Rio (See: Rio Teribe) Teribe, Rio (=Rio Terabe), 9°24'N-82°33'W (near Almirante-2) Timi de Boa, Rio Teribe (near Almirante-2) Timishik, Upper Rio Teribe (near Almirante-2) Vieja, Punta (=Punta Patino, 9°18'N-82°04'W, or a point on Laguna de Chiriqui?) CANAL ZONE Albrook Field, 8°59'N-79°34'W (near Fort Clayton-24) Alhajuela, 9°11'N-79°38'W (old hydrographic station between highway bridge and Mad- den Dam-27) Amador, Fort, 8°55'N-79°33'W (near Panama-23) Ancon, 8°57'N-79°34'W (near Panama-23) Balboa, 8°57'N-79°35'W (near Panama-23) Barro Colorado Island (29), 9°09'N-79°51'W Bas Obispo, 9°06'N-79°42'W (old village, now abandoned, on Panama Railroad, near Gamboa-28) Bohio, 9°10'N-79°51'W (old station on Panama Railroad, now under water, near Barro Colorado Island-29) Boracho, Loma, 9°17'N-79°56'W (near Fort Sherman-32) 14 ECTOPARASITES OF PANAMA Borinquen Road (=K-2 Road), west bank, Pacific Side (near Fort Kobbe-22) Cacao Plantation, 9°06'N-79°41'W (near Gamboa-28) Cardenas, 8°59'N-79°35'W (near Fort Clayton-24) Casaya, Rio, 9°06'N-79°41'W (near Gamboa-28) Chagres, Camp, 9°13'N-79°37'W (near Madden Dam-27; records of 20-30 years ago may refer to a locality within Fort Sherman-32) Chagres, Rio, 9°08'N-79°41'W (near Gamboa-28) Chico (probably Rio Chico Hydrographic Station, Panama, near Calzada Larga-26) Chiva Chiva, 9°01'N-79°35'W (near Fort Clayton-24) Clayton, Fort (24), 8°59'N-79°36'W Coco Solo, 9°21'N-79°54'W (near Fort Gulick-33) Cocoli, 8°58'N-79°36'W (near Fort Kobbe-22) Corozal, 8°58'N-79°35'W (near Fort Clayton-24) Corte Culebra Road Cristobal, 9°20'N-79°55'W (near Fort Gulick-33) Culebra, 9°03'N-79°40'W (near Gamboa-28) Curundu, 8°59'N-79°33'W (near Fort Clayton-24) Davis, Fort, 9°15'N-79°56'W (near Fort Gulick-33) El Vijia (=Vijia), 9°12'N-79°36'W (village, now under water, near Madden Dam-27) Emperador (See: Empire) Empire (= Emperador), 9°03'N-79°41'W (old administrative center of Canal, on west bank, about halfway between Paraiso and Gamboa-28) Farfan, 8°55'N-79°36'W (Near Fort Kobbe-22) France Field, 9°21'N-79°53'W (near Fort Gulick-33) Frijoles, 9°10'N-79°49'W (near Barro Colorado Island-29) Galeta Island, 9°23'N-79°53'W (near Fort Gulick-33) Galeta Point, 9°23'N-79°52'W (near Fort Gulick-33) Gamboa (28), 9°06'N-79°42'W Gatun, 9°15'N-79°56'W (near Fort Gulick-33) Gulick, Fort (33), 9°18'N-79°53'W Howard Field, 8°54'N-79°37'W (near Fort Kobbe-22) Indio, Rio, 9°15'N-79°59'W (near Fort Sherman-32) Juan Mina, 9°09'N-79°40'W (near Gamboa-28) Kobbe, Fort (22), 8°54'N-79°36'W K-9 Road (parallels south bank of Rio Cocoli for 3.3 mi. between K-2 and K-6 roads; near Fort Kobbe-22) K-10 Road (extends 6.6 mi. NW from Arraijan to head of Rio Mandinga; near Fort Kobbe-22) La Boca, 8°56'N-79°34'W (near Panama-23) Lagartera, 9°07'N-79°58'W (village, now under water, near Escobal-30) Las Cascadas, 9°05'N-79°42'W (near Gamboa-28) Las Cruces, 9°07'N-79°41'W (village, now under water, near Gamboa-28) Las Cruces Trail (extends from Rio Chagres, above mouth of Rio Casaya, through Mad- den Forest, Chiva Chiva, Cardenas, and Curundu to Panama) Lion Hill, 9°13'N-79°54'W (now an island in Gatun Lake, near Barro Colorado Island-29) Madden Dam (27), 9°13'N-79°38'W Madden Forest (25), 9°05'N-79°38'W Madden Road (=C-25 Road, extending between Paraiso and Madden Dam, and passing through Madden Forest) Madden Wye, 9°03'N-79°39'W (near Madden Forest-25) Mandinga, Rio, 9°05'N-79°42'W (near Gamboa-28) Margarita, 9°18'N-79°54'W (near Fort Gulick-33) Miraflores, 8°59'N-79°36'W (near Fort Clayton-24) Mohinga Valley (=Rio Mojinga), 9°18'N-79°59'W (near Fort Sherman-32) Mojinga, Rio (See: Mohinga Valley) Mount Hope, 9°19'N-79°54'W (near Fort Gulick-33) Orchid Island, 9°10'N-79°52'W (near Barro Colorado Island-29) FAIRCHILD AND HANDLEY : GAZETTEER 15 Paraiso, 9°02'N-79°39'W (near Madden Forest-25) Pedro Miguel, 9°01'N-79°37'W (near Fort Clayton-24) Pelado, Cerro, 9°07'N-79°43'W (near Gamboa-28) Pina, Camp, 9°16'N-80°00'W (near Fort Sherman-32) Puma Island, 9°13'N-79°55'W (near Barro Colorado Island-29) Quarry Heights, 8°57'N-79°34'W (near Panama-23) Randolph, Fort, 9°22'N-79°54'W (near Fort Gulick-33) Red Tank, 9°00'N-79°36'W (near Fort Clayton-24) Represo (near Barro Colorado Island-29) Rodman Naval Station, 8°56'N-79°35'W (near Fort Kobbe-22) Salamanca Hydrographic Station, 9°17'N-79°36'W (near Rio Pequeni-36) San Juan, 9°15'N-79°36'W (village, now under water, near Madden Dam-27) San Lorenzo, Fort, 9°18'N-80°01'W (near Fort Sherman-32) San Lorenzo Caves (See: Fort San Lorenzo) San Pablo, 9°06'N-79°48'W (old station on Panama Railroad, now under water, near Barro Colorado Island-29) Sherman, Fort (32), 9°21'N-79°57'W Summit, 9°03'N-79°40'W (near Madden Forest-25) Tabernilla, 9°07'N-79°49'W (old station on Panama Railroad, now under water, near Barro Colorado Island-29) Tigre, Cerro, 9°04'N-79°39'W (near Madden Forest-25) Venado Beach, 8°53'N-79°37'W (near Fort Kobbe-22) Vijia (See: ElVijia) CHIRIQUf Almijas, Isla, 8°16'N-82°24'W (near David-7) Bambito, 8°15'N-82°37'W (near Cerro Punta-4), 5000-6000 feet Baru, Cerro (See: Volcan de Chiriqui) Bogavo (See: Bugaba) Boqueron, 8°31'N-82°34'W (near Concepcion-6) Boquete (5), 8°47'N-82°25'W, 2000-7500 feet Boquete- Volcan Trail (near Boquete-5), above 6500 feet Brava, Isla, 8°12'N-82°16'W (near David-7) Bugaba (=Bogavo), 8°29'N-82°37'W (near Concepcion-6) Burica, Punta, 8°02'N-82°52'W (near Camp Pital-6a) Butz, Finca, 8°50'N-82°37'W (near Cerro Punta-4), 5000 feet Candela, Rio, 8°51'N-82°49'W (near Cerro Punta-4), above 3500 feet Casita Alta (See: Finca Lerida) Cerro Punta (4), 8°53'N-82°34'W, 5000-7800 feet Cerro Punta-Boquete Trail (between Cerro Punta-4 and Boquete-5), 6800-7800 feet Chiriqui, Volcan de (=Cerro Baru and El Baru), 8°49'N-82°32'W (near Cerro Punta-4), 6000-11,400 feet Chiriqui Viejo, Rio, 8°49'N-82°40'W (near Cerro Punta-4), above 3000 feet Colorado, Rio, 8°51'N-82°44'W (near Cerro Punta-4), 4000 feet Concepcion (=La Concepcion-6), 8°31'N-82°37'W Cotito, Rio, 8°51'N-82°45'W (near Cerro Punta-4), 4900 feet Goto Region (=Rio de la Vaca, base of Burica Peninsula, near Concepcion-6) Davala (See: Divala) David (7),8°26'N-82°26'W Divala (=Davala), 8°25'N-82°43'W (near Concepcion-6) El Banco, 8°42'N-82°31'W (near Boquete-5), near 3500 feet El Baru (See: Volcan de Chiriqui) El Hato del Volcan (See: El Volcan) El Volcan (=E1 Hato del Volcan and Lava Flow) , 8°47'N-82°38'W (near Cerro Punta-4) , 4000-6000 feet Gariche, Rio, 8°44'N-82°41'W (near Cerro Punta-4), 3200-5300 feet 16 ECTOPARASITES OF PANAMA Insolita, Isla (=Isla Porcada), 8°08'N-81°44'W (near Remedios-8) La Concepcion (See: Concepcion) La Vaca, Rio (See: Goto Region) Lara, Finca, 8°51'N-82°36'W (near Cerro Punta-4), 5600-5800 feet Lava Flow (See: El Volcan) Lerida, Finca (=Casita Alta), 8°49'N-82°29'W (near Boquete-5), 5000-7400 feet Lewis, Casa, 8°52'N-82°36'W (near Cerro Punta-4), 5600-5700 feet Llano Verde, 8°48'N-82°37'W (near Cerro Punta-4), 5000 feet Martinz, Finca (=Martinz Dairy), 8°52'N-82°34'W (near Cerro Punta-4), 6500-6800 feet Martinz Dairy (See: Finca Martinz) Nueva Colonia Palo Santo, 8°49'N-82°40'W (near Cerro Punta-4), 4200 feet Pando, Cerro, 8°55'N-82°43'W (near Cerro Punta-4), 3800-5600 feet Parida, Isla, 8°07'N-82°19'W (near David-7) Pedregal, 8°22'N-82°26'W (near David-7) Pital, Camp (between Puerto Armuelles and Costa Rican Boundary-6a) Porcada, Isla (See: Isla Insolita) Pueblo Nuevo, 8°06'N-81°42'W (near Remedios-8) Remedies (8), 8°14'N-81°50'W San Felix, 8°19'N-81°52'W (near Remedios-8) Santa Clara (=Quebrada Santa Clara and Rio Santa Clara), 8°51'N-82°46'W (near Cerro Punta-4), 3600-4200 feet Santa Clara, Quebrada (See: Santa Clara) Santa Clara, Rio (See: Santa Clara) Sereno, 8°51'N-82°51'W (near Cerro Punta-4), 3600-3700 feet Sevilla, Isla, 8°14'N-82°23'W (near David-7) Siolo (or Siola), 8°51'N-82°44'W (near Cerro Punta-4), 4100-4300 feet Tilley, Casa, 8°51'N-82°36'W (near Cerro Punta-4), 5300-5600 feet Wald (Rio Chiriqui Viejo, near Cerro Punta-4), 3800 feet COCLE Aguadulce (15), 8°14'N-80°33'W Anton, 8°24'N-80°16'W (near Rio Hato-17) El Cope, 8°37'N-80°35'W (near Penonome-16), 1500 feet El Potrero, 8°32'N-80°33'W (near Penonome-16) El Valle (18), 8°36'N-80°08'W, 2000-3000 feet Penonome (16), 8°31'N-80°22'W RioHato (17), 8°22'N-80°11'W Santa Clara, 8°22'N-80°07'W (near Rio Hato-17) COL6N Achiote, 9°12'N-80°01'W (near Pina-31) Aspinwall (See: Colon) Bonita, Quebrada (on Transisthmian Highway; near Madden Dam-27) Bruja, Cerro, 9°29'N-79°34'W (near Portobelo-35), 1000-2000 feet Buena Vista, 9°16'N-79°42'W (near Madden Dam-27) Cativa (or Catival), 9°21'N-79°51'W (near Fort Gulick-33) Cement Plant, 9°15'N-79°40'W (near Madden Dam-27) Colon (=Aspinwall),9021'N-79°55'W (near Fort Gulick-33) Donoso, 9°09'N-80°19'W (near Pina-31) El Limon (See: Nuevo Limon) Escobal (30),9°08'N-79°58'W Frijolito, Rio, 9°12'N-79°46'W (near Nuevo Limon-34) Guayabalito, 9°11'N-79°40'W (near Madden Dam-27) FAIRCHILD AND HANDLEY I GAZETTEER 17 Nuevo Limon (=E1 Limon and Puerto Limon) (34), 9°14'N-79°49'W Palenque, 9°42'N-79°22'W (near Portobelo-35) Pina (31),9°16'N-80°03'W Portobelo (or Porto Bello) (35), 9°41'N-79°41'W Puerto Limon (See: Nuevo Limon) Salud, 9°12'N-80°08'W (near Pina-31) Santa Rosa, 9°10'N-79°40'W (near Madden Dam-27) DARIEN Amagal, 7°24'N-78°02'W (near Jaque-47), 1000 feet Aruza, 8°02'N-77°39'W (near El Real-43) Boca de Cupe, 8°02'N-77°36'W (near El Real-43) Camoganti, 8°08'N-77°54'W (near El Real-43) Cana (=Santa Cruz de Cana), 7°47'N-77°42'W (near Rio Setsganti-46), 1800-3500 feet Cana, Loma (near Cerro Pirre and Rio Seteganti-46), 4900 feet Capeti, 8°04'N-77°33'W (near El Real-43) Casita (=Tacarcuna Casita), 8°01'N-77°22'W (near Tacarcuna Village-45), 1500 feet Chepigana, 8°17'N-78°04'W (near El Real-43) Chucunaque, Rio, 8°23'N-77°49'W (near El Real-43) Cituro, 8°00'N-77°36'W (near Boca de Rio Paya-44) El Real (=Real de Santa Maria) (43), 8°06'N-77°45'W Esnape, Rio (48), 8°05'N-78°13'W Guayabo, 7°23'N-78°02'W (near Jaque-47) Jaque (47) and Rio Jaque, 7°31'N-78°10'W Jesucito, Rio, 8°02'N-78°18'W (near Rio Esnape-48) La Laguna (=Tacarcuna Laguna), 8°04'N-77°19'W (near Tacarcuna Villaga-45), 3200 feet La Palma, 8°24'N-78°09'W (near El Real-43) Limon, Rio (between Cana and Cerro Pirre, near Rio Seteganti-46), 5100 feet Mali, Cerro, 8°07'N-77°14'W (near Tacarcuna Village-45) , 4100-4800 feet Marraganti, 8°08'N-77°44'W (near El Real-43) Mono, Rio, 7°43'N-77°33'W (near Boca de Rio Paya-44) Paya, Boca de Rio (44), 7°55'N-77°31'W Paya Village, 7°53'N-77°24'W (near Boca de Rio Paya-44) Pelisa (on Rio Pavarando?) Pina, Punta (or Pina Point), 7°34'N-78°13'W (near Jaque-47) Pinogana, 8°07'N-77°41'W (near El Real-43) Pirre, Cerro (or Mount Pirre), 7°51'N-77°44'W (near Rio Seteganti-46), 4500-5300 feet Pucro, Rio, 7°59'N-77°34'W (near Boca de Rio Paya-44) Punusa, Boca de Rio, 7°48'N-77°32'W (near Boca de Rio Paya-44) Real de Santa Maria (See: El Real) Santa Cruz de Cana (See: Cana) Sapo, Cerro (or Mount Sapo), 7°58'N-78°22'W (near Rio Esnape-48), 1000-3000 feet Seteganti, Rio (46), 7°46'N-77°40'W, 1600-3000 feet Tacarcuna, Cerro, 8°10'N-77°18'W (near Tacarcuna Village-45), 4100-4800 feet (Note: the "Mount Tacarcuna" of Anthony, Bull. American Mus. Nat. Hist., 35:357-376, 1916, and Goldman, Smithsonian Misc. Coll., 69, no. 5, 1920, is Cerro Mali) Tacarcuna Casita (See: Casita) Tacarcuna Laguna (See: La Laguna) Tacarcuna Village (=Tacarcuna Yellow Fever Station) (45), 8°05'N-77°17'W, 1800-3000 feet Tacarcuna Yellow Fever Station (See: Tacarcuna Village) Tapalisa (and Rio Tapalisa), 7°59'N-77°26'W (near Boca de Rio Paya-44) Tuira, Rio (or Rio Tuyra), 8°08'N-77°45'W (near El Real-43) Yaviza (or Yavisa), 8°09'N-77°42'W (near El Real-43) 18 ECTOPARASITES OF PANAMA HERRERA Azuero, Peninsula de (Herrera and Los Santos and part of Veraguas provinces) Capina, 7°53'N-80°33'W (near Parita-14) Divisa, 8°07'N-80°42'W (near Parita-14) Parita (14), 7°59'N-80°32'W Pese, 7°54'N-80°38'W (near Parita-14) Potuga, 8°04'N-80°38'W (near Parita-14) Los SANTOS Azuero, Peninsula de (Herrera and Los Santos and part of Veraguas provinces) Guanico (=Guanico Arriba and Las Palmitas), 7°2(XN-80°30'W (near Cerro Hoya-12) Hoya, Cerro (12), 7°18'N-80°42'W, 1500-3200 feet Las Palmitas (See: Guanico) Pedasi (13), 7°32'N-80°03'W PANAMA Aguas Buenas, 9°07'N-79°37'W (near Madden Forest-25) Arraijan, 8°57'N-79°41'W (near Fort Kobbe-22) Avaso, Rio (probably a misprint for Rio Abajo) Azul, Cerro (37) (See: Cerro Prominente, Goofy Lake, and La Zumbadora) Bayano, Rio, 9°06'N-79°05'W (near Chepo-40) Bejuco, 8°36'N-79°54'W (near Chame-19) Cabima, near 9°08'N-79°34'W (near Calzada Larga-26) Calidonia, 8°57'N-79°32'W (near Panama-23) Calzada Larga (=Casa Larga) (26), 9°10'N-79°34'W Campana, Cerro, 8°41'N-79°56'W (near Chame-19) Candelaria Hydrographic Station, 9°22'N-79°32'W (near Rio Pequeni-36) Capira, 8°45'N-79°53'W (near La Chorrera-20) Carasquilla (probably La Carasquilla, a suburb of Panama City; Casa Larga (See: Calzada Larga) Cativo, 9°10'N-78°50'W (near Chepo-40) Chame (19), 8°34'N-79°54'W Chapera, Isla, 8°34'N-79°03'W (Archipielago de las Perlas-41) Charco del Toro (head of Rio Maje; near Chiman-42) Chepo (40),9°10'N-79°06'W Chilibre, 9°08'N-79°38'W (near Madden Dam-27) Chilibrillo Caves (near Chilibre on Transisthmian Highway; near Madden Dam-27) Chiman (42), 8°43'N-78°37'W Cocos, Punta, 8°12'N-78°55'W (Archipielago de las Perlas-41) Goofy Lake (—"Cerro Azul" of Environmental Health Branch, U. S. Army), 9°09'N- 79°26'W (Cerro Azul-37), 750-2000 feet Guayabito (probably a locality near Chorrera, formerly Los Guayabitos) Huile, 9°01'N-79°46'W (near La Chorrera-20) Jefe, Cerro (See: La Zumbadora) Juan Diaz, 9°02'N-79°28'W (near Panama-23) La Chorrera (20), 8°52'N-79°48'W La Zumbadora (="Cerro Azul" of Gorgas Memorial Laboratory and C. O. Handley, Jr.; including Cerro Jefe), 9°14'N-79°21'W (Cerro Azul-37), 850-3200 feet Las Cumbres, 9°05'N-79°33'W (near Panama-23) Madden Airstrip, 9°10'N-79°34'W (near Calzada Larga-26) Maje, Rio, 8°40'N-78°32'W (near Chiman-42) Mamoni, Rio (See: Salto de Madrono) Maxon Ranch (See: Rio Trinidad) FAIRCHILD AND HANDLEY : GAZETTEER 19 Moja Polla, Rio (Quebrada), 9°11'N-79°39'W (near Madden Dam-27) Monte Obscuro (suburb of Panama-23) Nueva Gorgona, 8°33'N-79°53'W (near Chame-19) Nuevo Emperador (=Paja), 9°00'N-79°45'W (near La Chorrera-20) Pacheca, Isla, 8°39'N-79°04'W (Archipielago de las Perlas-41) Pacora (39), 9°04'N-79°18'W Paitilla, 8°57'N-79°32'W (suburb of Panama-23) Paja (See: Nuevo Emperador) Panama (=Panama City) (23), 8°58'N-79°32'W Panama Viejo, 8°59'N-79°30'W (near Panama-23) Pearl Islands (See: Archipielago de las Perlas) Pedro Gonzales, Isla, 8°22'N-79°07'W (Archipielago de las Perlas-41) Peluca, Rio, 9°22'N-79°34'W Pequeni, Rio (36), 9°22'N-79°32'W Perlas, Archipielago de las (=Pearl Islands) (41), 8°21'N-79°00'W Prominente, Cerro (="Cerro Azul" of E. A. Goldman), 9°13'N-79°18'W (Cerro Azul-37) , 500-2950 feet Pueblo Nuevo, 9°01'N-79°31'W (near Panama-23) Puente, Rio, 9°11'N-79°34'W (near Calzada Larga-26) Rey, Isla del (=Isla San Miguel), 8°23'N-78°56'W (Archipielago de las Perlas-41) Rio Abajo, 9°01'N-79°31'W (near Panama-23) Rio Chico Hydrographic Station, 9°15'N-79°31'W (near Calzada Larga-26) Sabanas (suburb of Panama-23) Saboga, Isla, 8°36'N-79°05'W (Archipielago de las Perlas-41) Salto de Madrono, Rio Mamoni (near Chepo-40) San Francisco de la Caleta, 8°59'N-79°30'W (near Panama-23) San Jose, Isla, 8°15'N-79°08'W (Archipielago ds las Perlas-41) San Miguel, 8°26'N-78°57'W (Archipielago de las Perlas-41) San Miguel, Isla (See: Isla del Rey) Taboga, Isla (21), 8°47'N-79°35'W Taboguilla, Isla, 8°47'N-79°32'W (near Isla Taboga-21) Tapia, 9°01'N-79°28'W (near Panama-23) Tocumen, 9°04'N-79°24'W (near Pacora-39) Trinidad, Rio (=Maxon Ranch), 8°57'N-80°00'W (near Escobal-30) Villa Rosario, 8°46'N-79°53'W (near La Chorrera-20) Vique, Punta, 8°52'N-79°40'W (near Fort Kobbe-22) SAN BLAS Armila (49) 8°40'N-77°27'W Cangandi, Rio, 9°26'N-79°07'W (near Mandinga-38) Mandinga (38), 9°29'N-79°05'W Pito, Rio, 8°42'N-77°32'W (near Armila-49) Puerto Obaldia, 8°40'N-77°26'W (near Armila-49) VERAGUAS Afuera, Isla (See: Isla Canal de Afuera) Altos Cacao, 7°39'N-80°49'W (near Cerro Hoya-12) 1400 feet Azuero, Peninsula ds (Herrera and Los Santos and part of Veraguas provinces) Bubi, Rio, 8°00'N-81°32'W (near Remedios-8) Calobre, 8°19'N-80°51'W (near Santiago-10) Calovebora, 8°48'N-81°12'W (near Santa Fe-11) Canal de Afuera, Isla (=Isla Afuera), 7°41'N-81°37'W (near Isla Coiba-9) Cebaco, Isla, 7°31'N-81°11'W (near Isla Coiba-9) Coiba, Isla (9), 7°26'N-81°45'W Gobernadora, Isla, 7°33'N-81°12'W (near Isla Coiba-9) 20 ECTOPARASITES OF PANAMA Mariato, Rio, 7°40'N-89°54'W (near Cerro Hoya-12) Paracote, 7°40'N-81°01'W (near Cerro Hoya-12) Santa Fe (11), 8°31'N-81°04'W Santiago (10), 8°05'N-80°59'W Sona, 8°00'N-81°19'W (near Santiago-10) Viejo, Cerro, 7°38'N-80°47'W (near Cerro Hoya-12), 2000-3300 feet Viejo, Rio, 8°03'N-81°34'W (near Remedios-8) Mites of the Subfamily Laelaptinae in Panama (Acarina: Laelaptidae) VERNON J. TIPTON J, ROBERT M. ALTMAN 2, AND CHARLES M. KEENAN 3 During the past few years we have collected several hundred small mam- mals in Panama. The ectoparasites removed from these animals form a most interesting collection, of which the Acarina are by far the most numer- ous. The present study is limited to mites of the subfamily Laelaptinae, since the vast majority of the mesostigmatid mites collected belong to eight genera of this subfamily. Of these, Laelaps Koch, Gigantolaelaps Fonseca, Eubrachylaelaps Ewing, Echinolaelaps Ewing and Mysolaelaps Fonseca, are most commonly associated with myomorph rodents ; TUT Baker and Wharton with hystricomorph rodents; Steptolaelaps Furman with sciuromorph ro- dents, and Haemolaelaps Berlese with both rodents and marsupials. As Furman and Tipton (1961) point out, "The study of neotropical para- sitic mites is of importance because they may fill key roles in epidemiological patterns which for the moment are confusing." Because they are numerous and widespread, it is possible that mites may figure prominently in the solu- tion of some of our more perplexing epidemiological problems in the tropics. We wish to acknowledge with gratitude the assistance given us by Captain Wallace P. Murdoch and Major Gordon Field and their staff of illustrators at the 406th Medical General Laboratory in Tokyo, Japan. All of our illustrations were prepared by this group. Dr. Charles 0. Handley, Smithsonian Institution, identified the host animals. Dr. Flavio da Fonseca, Institute Butantan, Sao Paulo, Brazil, as always, has been very kind in as- sisting us with information about species of the Laelaptinae in South 1 Lieutenant Colonel, Medical Service Corps, United States Army. The senior au- thor assumes complete responsibility for the descriptions of new species in this paper. For purposes of citation in subsequent publications, authorship of species described herein should be attributed to him alone. 2 Lieutenant Colonel, Medical Service Corps, United States Army. 3 Environmental Health Branch, Preventive Medicine Division, Office of the Chief Surgeon, United States Army Forces Southern Command, Fort Amador, Canal Zone. 23 24 ECTOPARASITES OF PANAMA America. We are especially indebted to Dr. Deane P. Furman, University of California at Berkeley, as our format and some of our keys essentially follow those of Furman and Tipton (1961) . KEY TO THE PANAMANIAN GENERA OF LAELAPTINAE FEMALES; FROM FURMAN AND TIPTON, 1961 1. Genito-ventral plate with one pair of setae 2 Genito-ventral plate with more than one pair of setae 4 2. Peritremalia produced posterior to stigmata; posterior seta of coxa II similar to setae of other coxae 3 Peritremalia not produced posterior to stigmata; posterior seta of coxa II longer than setae of other coxae Gigantolaelaps Fonseca 3. Spine-like seta on posterior margin of coxa III ; elongate setae on dorsal apices of femur and genu I; tectum with three apical lobes Eubrachylaelaps Ewing Coxa III without spine-like seta; femur and genu I without elongate setae; tectum a single rounded lobe Haemolaelaps Berlese 4. Genito-ventral plate with three pairs of setae Steptolaelaps Furman Genito-ventral plate with four pairs of setae 5 5. Central setae of dorsal plate minute; coxae without spiniform setae Mysolaelaps Fonseca Central setae of dorsal plate not minute; at least one pair of coxae with spini- form setae 6 6. Large mites, over 1 mm. long; sternal plate as long or longer than wide Echinolaelaps Ewing Medium sized mites, less than 1 mm. long; sternal plate never as long as wide ... .7 7. Peritremal plate broad; epigynial and anal plates fused or in juxtaposition Tur Baker and Wharton Peritremal plate narrow; epigynial and anal plates not fused or in juxtaposition Laelaps Koch We prefer to summarize the morphological characteristics of the genera in a tabular form (table 1) rather than treat each genus separately, since Tipton (1960) and Furman and Tipton (1961) have recently discussed this group of genera. Some of the characters listed in table 1 are variable in occurrence in certain genera. Thus, coarse dorsal setae occur consistently in species of Echinolaelaps, Steptolaelaps and Tur, but only in some species of Eubrachy- laelaps, Gigantolaelaps and Laelaps. There are numerous setae on the un- armed portion of the venter in some species of Haemolaelaps but not in others. The value of the tectum as a generic character has not been fully investigated. In Tur anomalus n. sp., the tectum is very difficult to see, but appears to be a single lobe ; it is trilobed in T. uniscutatus. Genus Echinolaelaps Ewing Macrolaelaps Ewing, 1929, Man. Ext. Parasites, p. 185. Echinolaelaps Ewing, 1929, loc. cit., pp. 185-186. Type-species : Laelaps echidninus Berlese, 1887. This genus is represented in Panama by two species, one of them (E. lowei n. sp.) known only from Panama. Echinolaelaps echidninus (Berlese) Laelaps echidninus Berlese, 1887, Acari, Myriap. Scorp. Italia, pt. 39, p. 157. Echinolaelaps echidninus Ewing, 1929, Man. Ext. Parasites, p. 185. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 25 MATERIAL EXAMINED : 9 females from Rattus rattus, Arraijan (Panama) , 5 April 1961, collected by C. M. Keenan and V. J. Tipton. REMARKS : Our Panamanian material differs very little from specimens in our collections from other parts of the world. We have collected hundreds of specimens of both R. rattus and R. norvegicus but have found only a single rat infested with E. echidninus. However, Laelaps nuttalli was very abun- dant. TABLE 1. COMPARATIVE CHART OF PANAMANIAN GENERA OF LAELAPTINAE (FEMALES) I .3 Character A -o s Idiosoma 0 + o over 1 mm. Pairs of setae 4 on G-V plate 1 1 1 Number of 4 lobes in tectum* 3 1 1 At least 1 ro- bust coxal seta + + o Pilosity of o venter sparse + + Sternal plate as long as wide 0 0 0 Dorsal setae coarse 0 0 0 I vs **4 +£ 1 ^ E-H + 0 0 4 3 4 3 1 3 0 + 'Refers only to Panamanian species Echinolaelaps lowei Tipton, new species. Plate 1. DIAGNOSIS: Like E. boultoni Furman and Tipton, E. lowei n. sp., has epigynial and anal plates which are not in juxtaposition. E. boultoni has bulbous setae on coxa I ; these setae are pilif orm to setiform in E. lowei. DESCRIPTION : Idiosoma. — 1040 /* long, 697 a wide (width measured at level of coxae IV). Dorsum. — Dorsal plate elliptical, with 37 pairs setae; 936 n long, 614 /a wide; covers almost entire dorsum, leaving only narrow, postero-lateral band of soft integument bearing 14 pairs' setae; posterior margin with three pairs longer than medial setae; penultimate setae about one-half length of last pair. Venter. — Tritosternum bifurcate near base, pilose ; sternal plate coarsely reticulate ; 198 ij. long, 161 fj, wide; with three pairs setae approximately same length; one pair slit-like pores just caudad of first pair setae. Endapodal plates well denned; with setae smaller than sternal setae. Metapodal plates small, elongate. Epigynial plate expanded; with prominent transverse lines; with four pairs setae, distance between first pair 130 /a, second pair 198 M, third pair 151 /*, fourth pair 73 n; anal plate widely separated from 26 ECTOPARASITES OF PANAMA epigynial plate; 146 ^ long, 114 n wide; distance between anterior margin of plate and anus much greater than length of anus; adanal setae 52 /j. long; postanal seta 104 ^ long. Legs. — Legs long and slender except for leg II, which is more robust. Setae of coxa I piliform to setiform; a short, bulbous seta on coxa III. Dorsal apices of femur and genu I with setae less robust than sternal setae. Gnathosoma. — Deutosternum with seven rows of two to five teeth per row. Gnathoso- mal setae about one-half length of medial hypostomal setae. Chelae robust, both digits toothed; fixed digit bearing long, bent seta. Hypopharyngeal processes prominent, fim- briate epipharynx sublanceolate, covered with denticles. Tectum not apparent. TYPE MATERIAL : Holotype female from Nectomys alfari (host no. 4082) , Cerro Azul (Panama), 2 February 1958, collected by R. M. Altman and C. M. Keenan. In the collection of the United States National Museum. A paratype female, same data and repository as the holotype. REMARKS: This species is named for Mr. Wilbur Lowe, a remarkable technician and a tireless worker. Genus Eubrachylaelaps Ewing Eubrachylaelaps Ewing, 1929, Man. Ext. Parasites, p. 186. Furman, 1955a, Ann. Ent. Soc. Amer., 48, (1-2), pp. 51-59 (revision and key). Cyclolaelaps Ewing, 1933, Proc. U.S. Nat. Mus., 82, (30), p. 5. Type-species: Laelaps hollisteri Ewing, 1925. Only one species of this genus is recorded from Panama. It would be unusual, however, if E. rotundus Fonseca is not collected in Panama in the future ; it occurs in nearby Venezuela. Eubrachylaelaps jamesoni Furman. Plate 2. Eubrachylaelaps jamesoni Furman, 1955a, Ann. Ent. Soc. Amer., 48, (1-2), pp. 52-54, figs: 1-4. MATERIAL EXAMINED: 153 females from Peromyscus nudipes, Cerro Punta; 83 females from P. nudipes from Rancho Mojica; 2 females from Oryzomys fulvescens, Cerro Punta and Rancho Mojica; 1 female from Reithrodontomys creper; 1 female from Heteromys desmarestianus, Cerro Punta; and hundreds of specimens in alcohol from both localities, mostly from P. nudipes; January 1960 to March 1962, collected by C. O. Handley, C. M. Keenan, V. J. Tipton, and C. E. Yunker. REMARKS : Our measurements of the dorsal and sternal plates of Panama specimens are roughly comparable to those given by Furman (1955a) for type material from Mexico. Specimens from Rancho Mojica are slightly smaller (average length of dorsal plate for 10 specimens, 590 /*) , less heavily sclerotized with the setae of the venter apparently a little more delicate than specimens from Cerro Punta (average length of dorsal plate for 10 speci- mens, 626 p.) . The average ratio of length to width of sternal plate is 2 to 1 for Rancho Mojica specimens and 2.1 to 1 for Cerro Punta specimens. We must add that different mounting techniques were used for speci- mens from these two localities. Additional recently collected specimens from Cerro Punta were treated in much the same manner (using heat as a part of the mounting technique) as the Rancho Mojica specimens and the morphological differences were then not nearly as apparent. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 27 Genus Gigantolaelaps Fonseca Gigantolaelaps Fonseca, 1939a, Mem. Inst. Butantan, 12: 12 (transl., p. 61). Type-species: Gigantolaelaps vitzthumi Fonseca, 1939. The genus Gigantolaelaps Fonseca is well represented in Panama, both by species and individuals. We have tentatively associated our Panamanian specimens with five described species. KEY TO THE PANAMANIAN SPECIES OF GIGANTOLAELAPS FEMALES 1. Sternal plate with two to six (usually three) accessory setae . G. oudemansi Fonseca Sternal plate with no accessory setae 2 2. Proximal seta of coxa I longer than distal seta ; both setae spinif orm G. goyanensis Fonseca Proximal seta of coxa I not longer than distal seta; distal seta always piliform, proximal seta usually piliform 3 3. Dorsal plate bearing approximately 400 setae; a relatively small species with dorsal plate not over 1100 /j. long G. inca Fonseca Dorsal plate bearing 86 setae; a relatively large species with dorsal plate over 1400 fi long 4 4. Antero-median projection of sternal plate scarcely produced beyond insertion of first pair of setae. Small paired setae of posterior margin of dorsal plate reach beyond margin for not more than one-fourth of their length. . . .G. gilmorei Fonseca Antero-median projection of sternal plate produced far beyond insertion of first pair of setae as a pronounced lobe. Small paired setae of posterior margin of dorsal plate reach beyond margin for more than one-fourth of their length .... G. wolffsohni (Oudemans) Gigantolaelaps gilmorei Fonseca. Plates 3 (figs. 4, 8) , 4. Gigantolaelaps gilmorei Fonseca, 1939a, Mem. Inst. Butantan, 12: 22 (transl., p. 71), figs. 6-10 (Goyaz State, Brazil). Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle, 21, (60), pp. 175, 177, pis. 2 (fig. 9), 3 (figs. 1, 2), 4 (fig. 6), 5 (fig. 2), 6 (fig. 2). MATERIAL EXAMINED : A total of 206 females and 1 male as follows : 179 females and 1 male from (35) Oryzomys capito; 27 females from (5) O. alfaroi; 13 females from (2) O. bombycinus; 3 females from (1) O. caligino- sus; 4 females from (1) Nectomys alfari; 3 females from (1) Sigmodon hispidus; 1 female from (1) Reithrodontomys sumichrasti; 1 female from (1) Zygodontomys microtinus; 1 female from (1) Proechimys semi- spinosus. Most specimens from O. capito and O. alfaroi were collected at Cerro Hoya (Los Santos) by C. 0. Handley, February 1962. Most others were collected at Cerro Azul (Panama) and Canal Zone, July 1956 to January 1962, by Robert M. Altman, C. M. Keenan and V. J. Tipton. REMARKS: G. gilmorei Fonseca is a large species (idiosoma well over 2000 /u, in length) , distinct from others of the genus in possessing the follow- ing combination of characters : the bases of the first pair of sternal setae are on or very near the anterior margin of the sternal plate ; both setae of coxa I are piliform ; and the epigynial setae do not extend to the caudal margin of the epigynial plate. 28 ECTOPARASITES OF PANAMA The majority of our specimens were collected from a group of closely related species of the genus Oryzomys at localities of 2000-3000 feet eleva- tion. They conform rather closely to the description and figures given by Fonseca (1939a) except that the epigynial setae are shorter, the setae of the venter are somewhat more sparse, and the anal and epigynial setae slightly more delicate than indicated by Fonseca. Gigantolaelaps goyanensis Fonseca. Plates 3 (figs. 1, 6) , 5. Gigantolaelaps goyanensis Fonseca, 1939a, Mem. Inst. Butantan, 12: 32 (transl., p. 81), figs. 15-18 (Brazil). Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle, 21, (60), p. 177, pis. 3 (fig. 3), 4 (fig. 5), 5 (fig. 8), 6 (fig. 3). MATERIAL EXAMINED: 22 females and 2 males from (1) Zygodontomys microtinus, Cerro Azul (Panama), 16 August 1956, collected by R. M. Alt- man and C. M. Keenan. REMARKS : G. goyanensis Fonseca is the only species of the genus thus far recorded from Panama in which both setae of coxa I are robust and the proximal seta is longer than the distal seta. A rather anomalous situation exists with respect to this species in that we have collected it only once al- though we have examined 74 specimens of Zygodontomys microtinus. Gigantolaelaps inca Fonseca. Plate 6. Gigantolaelaps inca Fonseca, 1960, Acarologia, 2, (1), pp. 11-14, figs. 1, 2 (Peru). Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle, 21, (60), pp. 182-184, pis. 2 (fig. 7), 3 (figs. 7, 8), 4 (fig. 4), 5 (fig. 7), 6 (fig. 6). MATERIAL EXAMINED: A total of 655 females and 2 males as follows: 568 females and 2 males from (25) Oryzomys albigularis ; 63 females from (8) Oryzomys alfaroi; 11 females from (1) Peromyscus flavidus; 9 females from (1) Didelphis marsupialis; 4 females from (1) Peromyscus nudipes; all collected above 5000 feet elevation near Cerro Punta (Chiriqui) or Ran- cho Mojica (Bocas del Toro) during January, February, and May, 1960, September 1961 and March 1962 by C. O. Handley, C. M. Keenan and V. J. Tipton. REMARKS : In Panama, G. inca Fonseca apparently occurs only at eleva- tions above 5000 feet. Like G. oudemansi Fonseca, it is a small species but it lacks the accessory setae on the sternal plate and it has very dense setae on the dorsal plate. Our specimens are not markedly different from those in our collection from Venezuela nor from figures and description given by Fonseca (1960) and Furman and Tipton (1961). The dorsal plates of our specimens appear to cover a greater area in relation to the entire specimen than do those from Venezuela and those from Peru as figured by Fonseca. However, measurements of the dorsal plates conform closely to those given by Fonseca and Furman and Tipton. Gigantolaelaps oudemansi Fonseca. Plates 3 (figs. 2, 7), 7. Gigantolaelaps oudemansi Fonseca, 1939a, Mem. Inst. Butantan, 12: 15 (transl., p. 64), figs. 1-5 (Goyaz State, Brazil). Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle, 21, (60), pp. 173, 175, pis. 3 (figs. 5, 6), 4 (fig. 3), 5 (fig. 6), 6 (fig. 5). TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 29 MATERIAL EXAMINED: A total of 960 females and 7 males as follows: 791 females and 6 males from (48) Oryzomys capito; 73 females from (6) O. alfaroi; 35 females from (2) O. bombycinus; 24 females from (2) O. caliginosus; 7 females from (1) O. bicolor; 1 female from (1) O. fulvescens; 22 females from (1) Nectomys alfari; 4 females, 1 male from (1) Sig- modon hispidus; 1 female from (1) Zygodontomys microtinus; 1 female from (1) Peromyscus flavidus; 1 female from (1) Didelphis marsupialis; from all of our collecting localities that were below 5000 feet elevation, July 1956 to February 1962, collected by R. M. Altman, C. O. Handley, C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS : G. oudemansi Fonseca is the only known species of the genus that has accessory setae on the sternal plate. We have collected G. oudemansi from a number of species of hosts. There are some morphological characters which vary according to host. Generally, there are three accessory setae on the sternal plate but in several specimens from 0. capito there were four or five accessory setae. Other measurable differences are presented in table 2, based on the same criteria used by Furman and Tipton (1961). Thus, populations from different hosts and different geographic locations may be compared. Gigantolaelaps wolffsohni (Oudemans). Plates 3 (figs. 3, 5, 9, 10), 8-11. Laelaps wolffsohni Oudemans, 1910, Rev. Chilena Hist. Nat., 14: 147 (Chile). Gigantolaelaps wolffsohni Morlan, 1951, Jour. Parasit., 37, (3), p. 273. Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle, 21, (60), pp. 179-181; pis. 3 (figs. 4, 9), 4 (figs. 1, 2), 5 (figs. 1, 3-5), 6 (figs. 1, 4). Macrolaelaps peruvianus Ewing, 1933, Proc. U. S. Nat. Mus., 82, (30), p. 7 (Lima, Peru). Gigantolaelaps peruvianus Fonseca, 1939a, Mem. Inst. Butantan, 12: 11 (transl., p. 60). MATERIAL EXAMINED: 140 females from hosts and localities mentioned below, collected from July 1956 to September 1961 by R. M. Altman, C. O. Handley, C. M. Keenan and V. J. Tipton. REMARKS: Furman and Tipton (1961) have discussed the morphological variation in 75 Venezuelan specimens of G. wolffsohni, the majority of which were from eight identified host species. We have 140 Panamanian speci- mens from six host species from six localities. Specimens taken from Oryzomys fulvescens at Rancho Mojica (pi. 11) are obviously quite different from those taken from O. caliginosus at Cerro Campana (pi. 8). On the basis of size alone they would seem to represent two distinct species, but in addition, the Cerro Campana specimens have ro- bust setae on coxa I, and shorter penultimate setae on the dorsal plate. How- ever, specimens from other hosts and other localities intergrade between these two extremes and in spite of vast differences in total size, the ratio be- tween plate sizes is essentially the same in all specimens. We have tended to disregard differences in length of setae because they may be broken off or may extend at an angle through a deep focal plane, rendering measure- ments unreliable and bringing the factor of distortion into an already con- fused picture. 30 ECTOPARASITES OF PANAMA F GIGANTOLAELAPS OU DEM AN SI, BY HOST SPECIES OF ORYZOMYS. (in microns) O. bombycinus O. caliginosus O. capita Venezuela (4) Los Santos (7) Canal Zone (10) Cerro Azul (10) 156-177 198-229 200-220 187-208 187-270 CO 0 CM O TH CM CM L- TH o co eg o I 10 os eg eg eg o CD eg oo os eg eg eg iH 00 eg o 1 CO 05 eg co eg c- o eg co 1 o -* eg O5 TH o CD eg •<* 1 •<* 05 eg co eg 0 CD eg •<* 05 eg eg eg t- t- TH IO eg co IO iH 10 CO iH oo TH 1 TH TH TH o TH TH o CD TH 3 iH CD IO TH s TH «o IO l-H 00 n< TH o CD rH IO TH TH O5 TH 10 Tl< iH 00 o eg 05 1 00 t- TH t- i-H 00 05 TH 00 CDTH CD iH 00 O eg o 1 05 O TH IO TH iH 00 TH IO 05 TH CD 00 O5 TH TH 1 00 CO TH CD iH 00 1— 1 eg o 1 05 t- iH t- TH •«* TH TH CO ^ TH IO 00 o c!S 00 t> 05 y-, _L O5 ^d1 O5 05 to CO00 00 IO TH O5 1 0 10 TH O5 co o 1 c- o eg co o C- eg TH 1 10 oo eg 0 eg TH oo eg eg oo eg TH eg co eg oo 1 eg o eg eg eg eg o co co 1 t- o eg IO eg eg TH co co 1 CD o eg 10 eg CO to TH eg iH iH co 10 iH IO IO TH CO TH CD CD TH O 1 10 0 TH TH O eg TH t- 1 TH TH iH CD TH O IO TH CO iH CD TH CO I CO TH TH specimens measured, ons. 0 «4H '£ EMENTS O O5 K eg £§ 2 O5 . iH TH T-l o co eg co I 10 10 eg eg o CO O 10 IO 00 O eg co 1 05 C- TH t- TH O 10 05 oo TH 00 eg CD I 10 eg eg CO eg IO t- co ••* TH TH J-l * "1 '« 3 cS K 5 O C 1 tn — i— i 1 bo ? 0) bo B C & 0> i O> c V bo B C es o> bo B V bo B B c3 8 -2 'B 'B B Q) eg' & £ «J§ « 1 «s tf § «s P5S •r-l £yO EH Character Length of Posterior t— 1 1— 1 03 X o O "8 0> V Length of Genito- Ventral Plate* Distance between Genital Setae Length of Genital Setae Length of Accessory Setae of Sternal Plate '3 PH -1J 0) iH 4J tii £ G S 4) — (J w Distance between 1st Pair Sternal Setae Numbers in parenthes* * Measured from level < TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 31 Our specimens fall into four groups designated A, B, C, and D. GROUP A (pi. 8), with 100 specimens from Oryzomys caliginosus, 2 specimens from Nectomys alfari, 1 specimen from Zygodontomys microtinus, from Cerro Azul, Cerro Campana, El Valle, and Bocas del Toro, is the largest of the four groups. The length of the dorsal plate ranges from a minimum of 1612 /* in the Bocas del Toro specimens to a maximum of 1956 p. in the Cerro Cam- pana specimens. The length of the sternal plate ranges from a minimum of 374 fj. in the Cerro Azul and Bocas del Toro specimens to a maximum of 455 /A in the Cerro Campana specimens. The setae of coxa I are more robust, the venter is more setose, and the epigynial plate is expanded somewhat more than in Group B. Group A resembles Fonseca's G. vitz- thumi. GROUP B (pi. 9), 12 specimens from Oryzomys capito and 2 specimens from O. fulvescens from El Valle, is characterized as follows : the length of the dorsal plate varies from 1612 to 1664 /x and the sternal plate ranges from 270 to 312 /A in length; the anterior margin of the sternal plate is not pro- jected forward as radically as in Group A ; the setae of coxa I are setiform, and the setation of the venter is somewhat more sparse than in Group A. GROUP C (pi. 10), 12 specimens from Sigmodon hispidus from Bocas del Toro, differs little from the previous group except that the coxal setae are very short and somewhat spiniform. The orientation of the coxae may ac- count for this seeming difference. Also, the venter is more setose than in Group B. GROUP D (pi. 11), 8 specimens from 0. fulvescens from Cerro Punta (Chiriqui) and 13 specimens from O. fulvescens from Rancho Mojica (Bocas del Toro), differs from the other groups in the following characters: size is small, the length of the dorsal plate varying from 1404 to 1580 p and that of the sternal plate from 281 to 292 p. ; the setae of coxa I, especially the distal seta, are pilif orm ; the penultimate setae of the dorsal plate extend beyond the margin of the plate for more than one-half of their length and the epigynial plate is expanded only slightly. This group appears to be Fon- seca's G. ivolffsohni, s. str. We have not overlooked the possibility that these four groups may repre- sent four distinct species, but because of the gradation in size, shown in table 3, we prefer to refer all of our specimens to one species, G. wolffsohni. A detailed study of specimens from its entire range will be necessary before the composition of this particular taxon is understood. However, for the mo- ment we may say that G. wolffsohni, s. lot., is a highly variable species which has both a wide geographical and ecological range. Comments on the Genus Gigantolaelaps Furman and Tipton (1961) have pointed out that the unusually high degree of apparent variation in Gigantolaelaps "may be linked in part to the evolutionary status of the host genera" as well as to a certain amount of dis- tortion associated with the mounting technique used. We have several long series each, from single host animals. The adults in these series apparently are not all the same age ; in addition, it appears that not all have fed as adults. It would be interesting and profitable to investigate the changes in total size 32 ECTOPARASITES OF PANAMA S 3? Cq o escens o a JH -M TH 05 CM CO 1 00 iH CM OO CM CO CO LQ 1 rH CM CO 0 CO "T ^ lO rH co o %& \ CM TJ< rH O O V '-' oo °> t- 0 CM Tf 1 10 00 CM CM co CO Tj< 1 co CM CO CM 10 CM CM 00 1 rH CO CM CM c- TH TH 0 rH rH 1 — i 00 lO t- •j CM CO rH rH CM co CM rH rH CO Oryzomy Rancho Mojica CM OJ CM 05 1 00 rH CM 00 CM CM co co o 1 rH rH CO OJ CM lO rH 00 1 CO CM rH CO rH 10 CM rH 0 \ rH O rH o rjH *o 00 0 CM t- 1 10 O CM 10 CM 312-332 318 os co CM CD 1 rH O CM O CM rH CM 1 co O rH CM TH CO 10 rH Sigmodon hispidus a| o H o pq -^ 73 302-312 308 co CO CM 1 rH rH CO 00 CM rH CD rH OO 00 rH rH CM g _o i — i CM co rH CD oo rH co 10 **3* CD o • 00 00 co «> c- CM CM CM CM co co CM CM rH rH co co CM U5 1 co 0 CM 0 oo os oo ^^ co I—I CM t- rH oS CD c 10 CD O o -^ CM CM CD •HI lO g! r^ S to -* oo 1 co co ^ TH -* CM 1 os -* CO rH CM CO 1 0 CM CM CM TH CO \ rH OS rH rH rH CM \ TH O rH CO co oo 1 rH CM CO CO CO CO 1 CM CM CO oo 1 00 kO CM rH rH OS 1 0 "^ ^H OS 2g O oj rH c- OS O rH rH rH CD 0 CM T"1 CO 0 "* CO rH rH rH co CO CM rH 00 rH CO o •l> CO CM CM CO lO CM *"*^ O r-H CM OS rH O C- O O5 g Tj< CO ^* T}* rH O rH C~ T^ ^« CO O5 CO CM CM D- rH rH ?H fo 1 rH CO CO 1 OJ \ 0 O 0 1 00 1 rH 1 OS 1 ^ CO cy ^ LO ^* CO CO 10 rH 0 rH rH rH rH CM rH CO lO CM co os t~- oo O ^ 00 oo O 00 00 •<# OS Tj^ rH g co TH rH CM CM CM C- o rH o 00 O CD lO OO OS 05 CM CM TH o CO 02 » O oo CM O O co rH OS co r . ^ CM co OJ CM t- rH rH TH CO CD CO O CM CM rH OS •— LO O t^ 1 OJ 1 ^* 1 0 \ O \ ^^ 1 rH 1 O 1 t- 1 rH 1 05 O •<* co CM CO 00 CM 00 rH 00 05 CM CO rH CO 05 CM CM CO M 13 t- I—I os OO CO O 05 CO rH rH rH .. co CO rH CO CM CM rH CO rH CD 0) O> d) o> o> 0) o> be c be c be c be C be fi be c be s be c ti J C oS oS ^ §1 C 03 oS o> fi 03 C | PH C$ C oj C oS C oS « s tf S P4 § K % « § tf S tf^ «S PH § .M § -(-> 01 oS -4-> 13 c •3 cp c co O) 13 c8 1 1 'rt +3 c 0) C oS 13 !-< CU -«J CO -»J O) to CO rH > 13 £ £ Q ^ o PH P-H 0) 0 "S V o o> £ « -^ oS 01 -JJ COS 0) OT PH 0 CO rH O Q L f ». i f~"i t-0 ^J m o! *i | oS •H fl o o> fi f~\ , <» o O g 0) CO Q X 0 X o o ^ O ^ O 0 e rC ^ o> i's r« U ,C X u — C os X 0 X "be^ be ^ c3 be be -T be " be " 03 .-y be ^ "S ^ 8 C m "to ^ C ^ c ,5 C "frt r- ^ "w C c £ "tr* O O> r5 J PH O> ^_, P-^ O S'g, rJ CO co "S 01 ,5 0) a) hJ CO .S QJ Q O 4) O> rJ CO CU r5 rJ PL. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 33 and degree of sclerotization over the life span of adult mites of this genus. It may be that plates and setae only appear to increase in size because of changes in sclerotization as a result of aging. Genus Haemolaelaps Berlese Haemolaelaps Berlese, 1910, Redia, 6: 261. Atricholaelaps Ewing, 1929, Man. Ext. Parasites, p. 186. Ischnolaelaps Fonseca, 1935, Mem. Inst. Butantan, 10: 19. Type-species: Haemolaelaps marsupialis Berlese, 1910. We have obtained specimens of this genus from nearly every species of rodent we have collected, as well as from several marsupials. It appears that almost all host species have a distinct population of Haemolaelaps and fur- ther, that each host species has distinctive populations in each collecting lo- cality. Either host specificity is very marked, with little individual varia- tion in a population, or else host specificity is almost entirely lacking and variation is much greater than is known for other laelaptine genera. If the latter is true, then it may be that variation is inversely proportional to host specificity. The shape of the pilus dentilis frequently has been used as a specific character. Allred (1958) has suggested that it would be better employed to separate groups of species. This structure is delicate and is susceptible to distortion which renders it of questionable taxonomic value. We have a series in which the pilus dentilis is inflated in some specimens while in others it appears to have burst during the mounting process. In another series from Nyctomys sumichrasti (pi. 13) , the apical as well as the proximal por- tion is swollen. Some of the specimens from Metachirus nudicaudatus are similar to those from Nyctomys, while others have the pilus dentilis inflated proximally only, as figured in plate 18. Other characters too are variable within a series from a single host ani- mal. Degree of sclerotization, length and number of setae (the latter may be due in part to orientation of the mite on the slide) , and shape and size of the anal plate vary considerably. On the other hand some characters are re- markably constant, e.g., the shape of the sternal plate, the shape and length of the setae of coxa I, and the relative length of the gnathosomal and hypo- stomal setae. We are referring most of our specimens to a single species, Haemolaelaps glasgowi (Ewing), although it is evident that there are some extreme dif- ferences among our specimens. Specimens collected from Peromyscus nudipes and Reithrodontomys sumichrasti at high elevations conform more closely to the figures and description given by Strandtmann (1949) than do those collected at low elevations. Plates 12 to 23 show the range of varia- tion. Haemolaelaps glasgowi (Ewing). Plates 12-23. Laelaps glasgowi Ewing, 1925, Proc. Ent. Soc. Wash. Haemolaelaps glasgowi Strandtmann, 1949, Jour. Pa MATERIAL EXAMINED : 102 females from Peromyscus nudipes, 41 females Laelaps glasgowi Ewing, 1925, Proc. Ent. Soc. Wash., 27: 1-7 (Illinois, U.S.A.). Haemolaelaps glasgowi Strandtmann, 1949, Jour. Parasit., 35, (3), pp. 325-352. 34 ECTOPARASITES OF PANAMA from Oryzomys fulvescens, 19 females from 0. albigularis, 14 females from Reithrodontomys mexicanus, 12 females from R. sumichrasti, 17 females and 1 male from R. creper, 1 female from Scotinomys teguina, and 19 females from S. xerampelinus, from Rancho Mojica, Cerro Punta or Boquete Trail; 49 females from Sigmodon hispidus, 14 females and 1 male from Oryzomys capito, 1 female from O. fulvescens, 4 females from O. caliginosus, 6 females from O. alfaroi, 1 female from O. bombycinus, 1 female from Tylomys pana- mensis, 11 females from Nectomys alfari, 4 females from Nyctomys sumi- chrasti, 3 females from Liomys adspersus, 4 females and 1 male from Hoplomys gymnurus, 7 females from Rattus rattus, 9 females from Proe- chimys semispinosus, 3 females from Heteromys desmarestianus, 2 females from H. australis, 2 females from Zygodontomys microtinus, 26 females from Sciurus granatensis, 47 females from Metachirus nudicaudatus, 8 females from Philander opossum, and 3 females from Didelphis marsupialis, mostly from Cerro Azul (Panama) and Canal Zone. REMARKS: Specimens from Rancho Mojica and Cerro Punta are more nearly like figures and description given by Strandtmann (1949) than those from Cerro Azul and the Canal Zone. Those from Hoplomys gymnurus are the largest (dorsal plate over 800 /*, in length) and those from Sigmodon hispidus the smallest (dorsal plate 572 /* in length) . The setae of the dorsal plate are robust and uniform in size in specimens from marsupials (pi. 18) , while in specimens from oryzomine rodents (pi. 13) , the setae of the dorsal plate are smaller and not uniform in size. The pilosity of unarmed areas in specimens from Nyctomys sumichrasti (pi. 13) is sparse whereas it is much denser in specimens from Metachirus nudicaudatus. In considering these differences, especially the extremes in the range of variation, it seemed advisable to describe several new species. However, it was impossible to find a well delineated group of specimens that did not merge into the next group. The genus Haemolaelaps is badly in need of revision. For the Neotrop- ical Region it will be necessary to study long series from many localities in order to understand variation and geographical distribution. To describe new species now would only add confusion to a problem which becomes even more complex as additional material becomes available. Genus Laelaps Koch Laelaps Koch, 1836, Deutschl. Crust. Myriap. Arach., pt. 4, p. 19. Tipton, 1960, Univ. Calif. Publ. Ent., 16, (6), pp. 260-262 (generic revision). Type-species: Laelaps agilis Koch, 1836. Several South American representatives of Laelaps occur in Panama. This is not surprising since several of the host genera occur throughout Central and South America. KEY TO THE PANAMANIAN SPECIES OF LAELAPS FEMALES 1. Gnathosomal setae at least twice as long as distal seta of coxa I L. dearmasi Furman and Tipton Gnathosomal setae not twice as long as distal seta of coxa I 2 TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 35 2. Distance between fourth pair of genito-ventral setae approximately the same as distance between first pair of genito-ventral s'etae L. nuttalli Hirst Distance between fourth pair of genito-ventral setae much less than the distance between first pair of genito-ventral setae 3 3. Distance between second pair of genito-ventral setae approximately four times greater than distance between fourth pair of genito-ventral setae L. paulistanensis Fonseca Distance between second pair of genito-ventral setae not more than three times greater than distance between fourth pair of genito-ventral setae 4 4. Proximal seta of coxa I at least three times as wide as distal seta 5 Proximal seta of coxa I les's than three times as wide as distal seta . . . L. thori Fonseca 5. Medial setae of the dorsal plate about 30 /* in length L. pilifer n. sp. Medial setae of the dorsal plate about 52 /j, in length L. castroi Fonseca Laelaps castroi Fonseca. Plate 24. Laelaps castroi Fonseca, 1958, Mem. Inst. Butantan, 28: 116 (Pernambuco, Brazil). MATERIAL EXAMINED : 64 females from O. alfaroi and 30 females from O. fulvescens from Cerro Punta (Chiriqui) ; 13 females from O, fulvescens and 1 female from Peromyscus nudipes from Rancho Mojica (Bocas del Toro) , January 1960 to March 1962, collected by C. 0. Handley, C. M. Keenan and V. J. Tipton. REMARKS: L. castroi may be distinguished from other species of the genus in Panama by the following combination of characters : short, robust proximal seta and a longer piliform seta on coxa I, fairly robust setae on the dorsal plate, and gnathosomal setae almost as long as the medial hypostomal setae. Our specimens are much smaller than those from Venezuela and are not as heavily sclerotized. The length of the idiosoma as recorded by Fonseca is 670 /a, while our specimens are only 560 /*. The size and shape of the setae of coxa I vary somewhat from the figures given by Fonseca but are of the same general configuration. Laelaps dearmasi Furman and Tipton. Plates 25, 26, 30 (figs. 4, 6, 7, 9) . Laelaps dearmasi Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle, 21, (60), pp. 187-191, pis. 8, 9. MATERIAL EXAMINED: A total of 191 females and 7 males as follows: 139 females, 5 males from Zygodontomys microtinus, 20 females from Proechimys semispinosus, 6 females and 1 male from Oryzomys capita, 1 female from O. caliginosus, 5 females from Rattus rattus, 19 females and 1 male from Sigmodon hispidus and 1 female from "rat" ; all from Canal Zone (except 16 females from Cerro Azul) , July 1956 to September 1960, collected by R. M. Altman, C. M. Keenan and V. J. Tipton. REMARKS : L. dearmasi is the only Panamanian species of Laelaps that has long, robust gnathosomal setae which extend beyond the posterior mar- gin of the gnathosoma. Panamanian specimens differ little from Venezuelan material. The distal seta of coxa I is somewhat more robust in our speci- mens. 36 ECTOPARASITES OF PANAMA Laelaps nuttalli Hirst Laelaps nuttalli Hirst, 1915, Bull. Ent. Res., 6: 183 (Colombo, Ceylon). Laelaps hawaiiensis Ewing, 1924, Bull. B. P. Bishop Mus. Honolulu, 98: 118. Haemolaelaps nuttalli Turk, 1950, Parasitology, 40, (1-2), p. 67. MATERIAL EXAMINED: 11 females from Rattus norvegicus, 6 females from Proechimys semispinosus, plus hundreds of specimens in alcohol from Rattus rattus; Canal Zone, July 1959 to December 1961, collected by C. M. Keenan and V. J. Tipton. REMARKS : L. nuttalli has a short, spinif orm distal seta and a longer pili- form proximal seta on coxa I. The distance between the first pair of epigy- nial setae is about the same as the distance between the fourth pair. Speci- mens from Panama fall within the range of intra-specific variation for this species and are very similar to specimens from the United States. Laelaps paulistanensis Fonseca. Plate 27. Laelaps paulistanensis Fonseca, 1935, XII Int. Congr. Zool., p. 1610. MATERIAL EXAMINED: 6 females from Oryzomys alfaroi from Cerro Punta (Chiriqui), January and February 1960, collected by C. M. Keenan and V. J. Tipton. REMARKS : L. paulistanensis is the largest of the Panamanian species and as in L. dearmasi, its epigynial plate is greatly expanded so that the distance between the second pair of setae is at least twice as great as the distance between the fourth pair. Our specimens are smaller (936 /* compared with 1030 ^ recorded by Fonseca) , the anterior setae of the dorsal plate are longer, the anal plate is broader in outline, and the anterior margin straighter than is described and figured by Fonseca (1936). However, the measurements of the ventral plates and setae and the coxal seate conform quite closely to Fonseca's de- scription. Laelaps pilifer Tipton, new species. Plates 28, 30 (figs. 5, 11) . DIAGNOSIS : L. pilifer n. sp., belongs to a species complex which includes L. paulistanensis Fonseca, L. differens Fonseca, L. castroi Fonseca, L. oryzomydis Pratt and Lane, and L. manguinhosi Fonseca. L. paulistanensis and L. pilifer have short delicate setae on the dorsal plate, a robust proximal seta and a piliform distal seta on coxa I. L. paulistanensis is a larger species (idiosoma over 1000 /* compared with 510 /*) and the distance between the second pair of epigynial setae is four times the distance between the fourth pair, while it is approximately three times in L. pilifer. In the latter species the distal seta of coxa I is much more delicate and the proximal seta shorter than in L. paulistanensis. DESCRIPTION : Idiosoma. — 510 /* long, by 322 M wide. Dorsum. — Dorsal plate elliptical, with prominent shoulders; 477 n long by 292 /t wide; covers almost entire dorsum leaving only narrow, latero-posterior band of soft integument. Setae of dorsal plate rather delicate, not reaching to bases of setae in next row; 35 pairs; penultimate pair much smaller than medial setae; last pair longer than others; one pair of round pores adjacent to penultimate setae, an additional pair cephalad and laterad of these. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 37 Venter. — Tritosternum bifurcate, pilose. Sternal plate 108 ^ wide by 86 /* long with three pairs setae of about same length; two pairs slit-like pores. Endapodal plates rather well defined; setae about same size as sternal setae. Epigynial plate expanded, with four pairs setae; distance between first pair 80 p., second pair 145 /*, third pair 88 /x, fourth pair 52 /*; surface of plate with transverse lines. Metapodal plates discrete. Anal plate shield-like; 77 /j. wide by 58 /j. long (measured to base of postanal seta) ; adanal setae originate cephalad of posterior margin of anus ; 31 /* long. Postanal seta 40 /* long. Approximately 10 pairs setae on unarmed venter. Peritreme reaches beyond middle of coxa II ; extends posterior of stigma. Legs. — Robust, with claws and caruncles. Leg IV longest, leg III shortest. Coxae I, III with spiniform setae; proximal seta of coxa I short, spiniform; distal seta very delicate, shorter than proximal seta. Dorsal apices of femur and genu I with setae no stronger than dorsal setae. Gnathosoma. — Deutosternum with seven rows of two to five teeth per row. Gnatho- somal setae slightly more than one-half length of medial hypostomal setae. Epipharynx long, apex rounded. Hypostome bipartite, fimbriate. Chelae well developed, toothed; fixed digit with long, slightly inflated seta. Tectum single, membranous flap; apex rounded. TYPE MATERIAL : Holotype female from Oryzomys capita (field no. 7104) , Rio Seteganti (Darien), 5 February 1961, collected by C. M. Keenan, V. J. Tipton and C. E. Yunker. In the collection of the United States National Museum. Paratypes : 80 females from O. capita, 2 females from O. caligino- sus and 1 female from 0. bombycinus, Cerro Azul ; 57 females from O. capita, 9 females from Proechimys semispinosus and 1 female from Didelphis marsupialis, Camp Pina (Canal Zone) ; 10 females from O. capita, Rio Seteganti (Darien) ; 8 females from O. capita, Cerro Hoya (Los Santos) ; July 1956 to February 1962, collected by R. M. Altman, C. O. Handley, C. M. Keenan and V. J. Tipton. In the collections of the Environmental Health Branch, Canal Zone, and V. J. Tipton. Laelaps thori Fonseca. Plates 29, 30 (figs. 1, 8). Laelaps thori Fonseca, 1939b, Mem. Inst. Butantan, 12: 111 (transl., p. 133), fig. 5. MATERIAL EXAMINED: 14 females from Oryzomys albigularis and 2 fe- males from Peromyscus flavidus, Rancho Mojica (Bocas del Toro) , Septem- ber 1961, collected by V. J. Tipton; 3 females from O. albigularis, Cerro Punta (Chiriqui), February 1960, collected by C. M. Keenan and V. J. Tip- ton ; 2 females from O. capita, Cerro Azul, July 1956, collected by R. M. Alt- man and C. M. Keenan. REMARKS: L. thori has delicate dorsal setae and the proximal seta of coxa I is stout but not spiniform. A comparison of our specimens with fig- ures given by Fonseca reveals some differences which may constitute spe- cific characters. In our specimens the venter is more setose, the anal plate appears to be wider, and the dorsal setae are shorter and more delicate. In spite of these differences we are referring our specimens to this species. In specimens from Cerro Azul (pi. 29) the proximal seta is shorter and thicker than in those from Rancho Mojica, and there are two or three more setae on the venter. Specimens from Cerro Punta are closer to Fonseca's figures than those from Rancho Mojica and the Rancho Mojica specimens are closer than those from Cerro Azul. This entire series is probably con- specific and may even be an extension of the L. pilifer-castroi complex. 38 ECTOPARASITES OF PANAMA Laelaps species. Plate 30 (figs. 2, 10) . MATERIAL EXAMINED : A single female from Oryzomys bicolor, Camp Pifia (Canal Zone), 29 August 1956, collected by R. M. Altman and C. M. Keenan. REMARKS : This specimen differs markedly from L. pilifer n. sp. in that the gnathosomal setae are minute, the proximal seta of coxa I is setiform, not spinif orm and the pilus dentilis is inflated as in Haemolaelaps. Although it probably represents an undescribed species of Laelaps, we shall await fur- ther material before making a decision as to its disposition. Comments on the Genus Laelaps We are not satisfied with this arrangement of the Laelaps species. It is only tentative. We lack basic biological information — e.g., on host specific- ity, ecological and geographical patterns, and morphological changes in rela- tion to age — essential to an understanding of the systematics of a taxon. Apparently the species of this genus have a wide geographic range. Within a large group of specimens, the castroi-pilifer-thori complex, there are several more or less distinct subgroups. The characters which separate the species of the complex are not clearly defined and there is ex- treme variation within a long series from the same host specimen. Charac- ters thought to be constant in this genus, e.g., shape and size of coxal setae, plate size and shape and details of the chelicerae, are variable even on the same specimen. If one could obtain specimens from enough localities a few miles apart, it is possible that one would find clines of characters throughout Central and South America. Many species now regarded as distinct might prove to represent population differences along these clines. Genus Mysolaelaps Fonseca Mysolaelaps Fonseca, 1935, Mem. Inst. Butantan, 10: 17. Type-species: Mysolaelaps parvispinosus Fonseca, 1935. We have collected only one species of Mysolaelaps although a second is probably present. Mysolaelaps heteronychus Fonseca occurs on a species of Rhipidomys in Venezuela. We have been unable to collect this host in Panama, although one species is represented in the fauna. Mysolaelaps parvispinosus Fonseca. Plate 31. Mysolaelaps parvispinosus Fonseca, 1936, Mem. Inst. Butantan, 10: 17. (Sao Paulo, Brazil). MATERIAL EXAMINED: 2 females from Oryzomys capito, El Valle, 27 March 1957, and 3 females from O. fulvescens, Cerro Jeffe, 7 February 1958, collected by R. M. Altman and C. M. Keenan. REMARKS : Mysolaelaps parvispinosus is readily separated from the other two neotropical species of Mysolaelaps. The genito-ventral setae of M. heteronychus are minute and approximately equal in size, while in M. par- vispinosus the first pair of genito-ventral setae are much smaller than the last three pairs. The sternal setae of M. microspinosus are small and ap- TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 39 proximately equal in size whereas the first pair of sternal setae of M. par- vispinosus are small, the second and third pairs are more robust and are more than twice as long as the first pair. The first pair of epigynial setae measure 68 to 80 /j, long whereas they were only 57 //, long in specimens from Brazil and Venezuela. The fourth pair of epigynial setae are more widely separated in our specimens (169- 198 /j.; mean of five specimens, 175 /x) than in Venezuelan specimens (130 to 156 ju,; mean of five specimens, 151 p.). Although our series is small there appears to be considerable variation in this species. Genus Steptolaelaps Furman Steptolaelaps Furman, 1955b, Jour. Parasit., 41, (5), p. 519. Type-species : Neolaelaps heteromys Fox. Only one species of this genus has been collected in Panama. Steptolaelaps heteromys (Fox). Plates 32, 33. Neolaelaps heteromys Fox, 1947, Zoologica, 32, (3), pp. 117-119 (Venezuela). Steptolaelaps heteromydis Furman, 1955b, Jour. Parasit., 41, (5), p. 521. Steptolaelaps heteromys Furman and Tipton, 1961, Mem. Soc. Cienc. Nat. La Salle; 21, (60), p. 195. MATERIAL EXAMINED : 7 males and 43 females from Heteromys desmares- tianus; 23 males, 71 females and 2 nymphs from H. australis; 4 females from Liomys adspersus; 2 females from Oryzomys sp. ; 1 female from Tylomys panamensis ; from the Canal Zone, Rio Seteganti, Cerro Azul, and El Valle; March 1957, January and February 1958, January and October 1960, col- lected by R. M. Altman, C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS: S. heteromys is distinct from the only other species of the genus, S. liomydis, in having long and tapering (rather than short) gnathosomal setae and in details of the female chelicerae (Furman, 1955b) . The adanal setae in our specimens appear to be slightly longer than in those figured by Furman (1955b) . Otherwise, our specimens are remarkable for their lack of variation. These mites are most commonly associated with heteromyid rodents and have been collected most frequently at elevations below 3000 feet. We have collected 76 specimens of Heteromys desmarestianus at elevations above 3000 feet. None of them harbored S. heteromys. Genus Tur Baker and Wharton Protonyssus Turk, 1946, Ann. Mag. Nat. Hist., (11), 13:347. Tur Baker and Wharton, 1952, Introd. Acar., p. 85 (new name for Protonyssus Turk, not Protonyssus Trouessart, 1915). Furman and Tipton, 1958, Jour. Parasit., 44, (5), pp. 541-7 (redescription of genus). Type-species : by original designation and monotypy, Protonyssus uniscutatus Turk. A new species, described below, possesses characteristics of both Laelaps and Tur. This has raised serious doubts as to the validity of the genus Tur. However, we choose to recognize it for the time being. 40 ECTOPARASITES OF PANAMA Tur anomalus Tipton, new species. Plates 30 (figs. 3, 12), 34. DIAGNOSIS: Tur anomalus n. sp., runs to T. lativentralis (Fonseca) in the key provided by Furman and Tipton ( 1961 ) . It is similar to that species in that the anal plate is separate from the epigynial plate and the gnathoso- mal and hypostomal setae are reduced in both species. However, T. anoma- lus is only 525 ^ long whereas T. lativentralis is more than 1000 /x. In addi- tion, the setae of coxa I are very short and robust and the adanal setae do not reach the base of the postanal seta in T, anomalus, while the setae of coxa I are at least half the length of the coxa and the adanal setae reach beyond the base of the postanal seta in T. lativentralis. DESCRIPTION : Idiosoma. — 525 /j. long by 309 /j. wide. Dorsum. — Dorsal plate strongly elliptical, with prominent shoulders; 504 p long by 260 fj. wide; does not cover entire dorslim, with narrow postero-lateral band of soft integument. Setae of dorsal plate in definite pattern; 39 pairs setae; some reticulation adjacent to bases of setae. Venter. — Tritosternum pilose. Sternal plate 128 /* wide by 109 fj. long. Anterior margin straight, posterior margin slightly convex; three pairs sternal setae of approxi- mately same length; two pairs slit-like pores. Endapodal plates fairly well defined; setae slightly longer than sternal setae. Metapodal plates elliptical. Epigynial plate expanded; with four pairs of setae; distance between first pair of setae 62 /*, second pair 108 /*, third pair 126 /*, fourth pair 92 M; posterior margin truncate, in juxta- position with anal plate. Anal plate broadly triangular with straight anterior margin ; 92 fj. wide by 71 fj. long (measured to base of postanal seta) ; adanal setae with bases cephalad of posterior margin of anus; 24 ^ long; postanal seta 28 /JL long. Approxi- mately nine pairs of setae on unarmed portion of venter. Peritremalia posterior to stigmata broad, lying in juxtaposition with parapodal plates which partially encircle coxae IV; peritremes sinuate, reaching to anterior margin of coxae II. Legs. — Coxa I with two stout, striated, spiniform setae; proximal seta longer than distal seta; spiniform setae on coxae II and III; femur I with strong spiniform seta on ventral surface, two strong setae on dorsal surface. Legs I and II more robust than other legs; leg IV longest, leg III shortest. Gnathosoma. — Deutosternum with usual rows of teeth. Gnathosomal setae slightly coarser than medial hypostomal setae. Epipharynx tongue-like, with medial groove. Details of the chelicerae not readily discernible but long seta at base of chela; chelae toothed; pilus dentilis apparently absent. TYPE MATERIAL: Holotype female from Hoplomys gymnurus (field no. 4038), Cerro Azul (Panama), 29 January 1958, collected by R. M. Altman and C. M. Keenan. In the collection of the United States National Museum. Eleven paratype females, same data and repository as the holotype. REMARKS: T. anomalus exhibits even more characteristics of Laelaps than does T. lativentralis. The gnathosomal and hypostomal setae are re- duced. The posterior margin of the sternal plate is not deeply invaginated and the setae of the dorsal plate as well as of the venter are not robust as in other species of the genus. The broad peritremalia, the anal plate lying in juxtaposition with the epigynial plate and the details of the female chelicerae, though not clearly visible, show some affinities with other species of Tur. The discovery of this species emphasizes again (Furman and Tipton, 1961) that the differences between Tur and Laelaps are not as marked as was first supposed. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 41 Tur uniscutatus (Turk) . Plates 35, 36. Protonyssus uniscutatus Turk, 1946, Ann. Mag. Nat. Hist., (11), 13: 347. Tur uniscutatus Baker and Wharton, 1952, Introd. Acar., p. 85. Furman and Tipton, 1958, Jour. Parasit., 44 (5), pp. 541-547. MATERIAL EXAMINED: 1228 females and 43 males from Proechimys semispinosus from the Canal Zone ; 303 females from P. semispinosus from Almirante (Bocas del Toro) and 21 females and 1 male from P. semispinosus from Cerro Azul; 16 females from Didelphis marsupialis, 3 females from Marmosa robinsoni, 1 female from Philander opossum and 1 female from Heteromys desmarestianus , Camp Pina (Canal Zone) ; 13 females from Nasua nasua, Gamboa (Canal Zone) ; 6 females from Hoplomys gymnurus, Cerro Azul and Almirante ; 13 females from "rat", Barro Colorado Island ; collected from July 1956 to July 1960 by R. M. Altman, C. 0. Handley, C. M. Keenan and V. J. Tipton. REMARKS: More than 95 percent of our 1649 specimens are from Proechimys semispinosus. Although this represents a long series, the range of variation is not as great as in species of other neotropical genera. Plates 35 and 36 show the extremes in variation, mostly differences in the diameter of the gnathosomal setae, size and number of setae on the venter, and minor differences in the details of the female chelicerae. Figures of Panamanian specimens given by Furman and Tipton (1958) differ slightly from our il- lustrations in these same characters. A study of all the specimens from which the drawings were made reveals that the differences are not as great as the illustrations indicate. However, specimens from Bocas del Toro do have heavier body setae and the female holoventral plate is slightly more ex- panded than in specimens from other localities. 42 ECTOPARASITES OF PANAMA HOST-PARASITE LIST Class MAMMALIA Order MARSUPIALIA Family Didelphidae Marmosa robinsoni Tur uniscutatus (Turk) Didelphis marsupialis Gigantolaelaps inca Fonseca oudemansi Fonseca Laelaps pilifer n. sp. Tur uniscutatus (Turk) Haemolaelaps glasgowi (Ewing) Philander opossum Tur uniscutatus (Turk) Haemolaelaps glasgowi (Ewing) Metachirus nudicaudatus Haemolaelaps glasgowi (Ewing) Order RODENTIA Suborder Sciuromorpha Family Sciuridae Sciurus granatensis Haemolaelaps glasgowi (Ewing) Family Heteromyidae Heteromys australis Steptolaelaps heteromys (Fox) Haemolaelaps glasgowi (Ewing) Heteromys desmarestianus Steptolaelaps heteromys (Fox) Eubrachylaelaps jamesoni Furman Tur uniscutatus (Turk) Haemolaelaps glasgowi (Ewing) Liomys adspersus Steptolaelaps heteromys (Fox) Haemolaelaps glasgowi (Ewing) Suborder Myomorpha Family Cricetidae Oryzomys albigularis Gigantolaelaps inca Fonseca Laelaps thori Fonseca Oryzomys alfaroi Gigantolaelaps gilmorei Fonseca inca Fonseca oudemansi Fonseca Laelaps castroi Fonseca " paulistanensis Fonseca Haemolaelaps glasgowi (Ewing) Oryzomys bicolor Gigantolaelaps oudemansi Fonseca Laelaps sp. Oryzomys bombycinus Gigantolaelaps gilmorei Fonseca oudemansi Fonseca Laelaps pilifer n. sp. Haemolaelaps glasgowi (Ewing) Oryzomys caliginosus Gigantolaelaps gilmorei Fonseca oudemansi Fonseca wolffsohni (Oudemans) Laelaps dearmasi Furman and Tipton pilifer n. sp. Haemolaelaps glasgowi (Ewing) Oryzomys capito Gigantolaelaps gilmorei Fonseca oudemansi Fonseca wolffsohni (Oudemans) Haemolaelaps glasgowi (Ewing) Laelaps dearmasi Furman and Tipton pilifer n. sp. " thori Fonseca Mysolaelaps parvispinosus Fonseca Oryzomys fulvescens Gigantolaelaps oudemansi Fonseca wolffsohni (Oudemans) Eubrachylaelaps jamesoni Furman Laelaps castroi Fonseca Mysolaelaps parvispinosus Fonseca Haemolaelaps glasgowi (Ewing) Oryzomys sp. Steptolaelaps heteromys (Fox) Nectomys alfari Gigantolaelaps gilmorei Fonseca oudemansi Fonseca wolffsohni (Oudemans) Echinolaelaps lowei n. sp. Haemolaelaps glasgowi (Ewing) Tylomys panamensis Steptolaelaps heteromys (Fox) Haemolaelaps glasgowi (Ewing) Nyctomys sumichrasti Haemolaelaps glasgowi (Ewing) Reithrodontomys creper Eubrachylaelaps jamesoni Furman Haemolaelaps glasgowi (Ewing) Reithrodontomys mexicanus Haemolaelaps glasgowi (Ewing) TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES 43 Reithrodontomys sumichrasti Gigantolaelaps gilmorei Fonseca Haemolaelaps glasgowi (Ewing) Peromyscus flavidus Gigantolaelaps Inca Fonseca oudemansi Fonseca Laelaps thori Fonseca Peromyscus nudipes Eubrachylaelaps jamesoni Furman Gigantolaelaps inca Fonseca Laelaps castroi Fonseca Haemolaelaps glasgowi (Ewing) Zygodontomys microtinus Gigantolaelaps gilmorei Fonseca goyanensis Fonseca oudemansi Fonseca wolffsohni (Oudemans) Laelaps dearmasi Furman and Tipton Haemolaelaps glasgowi (Ewing) Scotinomys teguina Haemolaelaps glasgowi (Ewing) Scotinomys xerampelinus Haemolaelaps glasgowi (Ewing) Sigmodon hispidus Gigantolaelaps gilmorei Fonseca oudemansi Fonseca wolffsohni (Oudemans) Laelaps dearmasi Furman and Tipton Haemolaelaps glasgowi (Ewing) Family Muridae Rattus norvegicus Laelaps nuttalli Hirst Rattus rattus Echinolaelaps echidninus (Berlese) Laelaps dearmasi Furman and Tipton nuttalli Hirst Haemolaelaps glasgowi (Ewing) Suborder Hystricomorpha Family Echimyidae Proechimys semispinosus Tur uniscutatus (Turk) Gigantolaelaps gilmorei Fonseca Laelaps dearmasi Furman and Tipton nuttalli Hirst pilifer n. sp. Haemolaelaps glasgowi (Ewing) Hoplomys gymnurus Tur uniscutatus (Turk) anomalus n. sp. Haemolaelaps glasgowi (Ewing) Order CARNIVORA Family Procyonidae Nasua nasua Tur uniscutatus (Turk) References ALLRED, D. M. 1958. Mites found on mice of the genus Peromyscus in Utah. IV. Families Lae- laptidae and Phytoseiidae (Acarina). Pan-Pac. Ent., 34, (1): 17-32, pis. I-VI. BAKER, E. W., AND WHARTON, G. W. 1952. An introduction to acarology. New York, Macmillan Co., xiii + 465 pp., 377 figs. BERLESE, A. 1882-1902. Acari, Myriapoda et Scorpiones hucusque in Italia reperta. Portici et Padua. 1910. Lista di nuove specie e nuovi generi di Acari. Redia, 6, (1909), pp. 242-271. EWING, H. E. 1924. Ectoparasites of some Polynesian and Malaysian rats of the genus Rattus. Bull. B. P. Bishop Museum, 14: 7-11, 1 fig., 1 map. 1925. New parasitic mites of the genus Laelaps. Proc. Ent. Soc. Wash., 27: 1-7. 1929. A manual of external parasites. Springfield, Illinois. Charles C. Thomas. xv + 225 pp., 96 figs. 1933. New genera and species of parasitic mites of the super-family Parasitoidea. Proc. U. S. Nat. Mus., 82, art. 30, pp. 1-14, 4 pis. FONSECA, F. DA. 1936. Notas de Acareologia. XVIII. Generos e especies de acarianos parasitas de ratos (Acari. Laelaptidae). Mem. Inst. Butantan, 10: 17-23. ] 937. New genera and species of Acari "Laelaptidae" from Brazilian rodents. XII Int. Congr. Zool., Lisboa, pp. 1597-1615, 1 fig. 1939a. Notas de Acareologia. XXV. Os Laelaptidae gigantes, parasitas de reodores sul-americanos ; genero e especies' novos (Acari). Mem. Inst. Butantan, 12: 7-53, figs. 1-30. (English translation, pp. 55-102). 1939b. Notas de Acareologia. XXVI. Novos estudos sobre o genero Laelaps Koch, 1836 (Acari. Laelaptidae). Loc. cit., pp. 103-123, figs. 1-9. (English transla- tion, pp. 125-145). 1958. Notas de Acareologia. XLIV. Inquerito sobre a fauna acarologica de para- sitas no Nordeste do Brasil. Mem. Inst. Butantan, 28: 99-186, figs. 1-54. 1960. Notes d'Acarologie. XLV. Enquete Acarologique au Perou. Acarologia, 2, (1), pp. 1-34, figs. 1-12. Fox, I. 1947. Notes on ectoparasites from Venezuela (Siphonaptera and Acarina). Zoo- logica, 32, (3), pp. 117-119, figs. 1-2. 44 TIPTON, ALTMAN, AND KEENAN I LAELAPTINE MITES 45 FURMAN, D. P. 1955a. Revision of the genus Eubr achy lae laps (Acarina: Laelaptidae) with the descriptions of two new species from Mexico. Ann. Ent. Soc. Amer., 48: 51-59, pis. I-III. 1955b. Steptolaelaps, a new genus of mites parasitic on neotropical rodents. Jour. Parasit., 41, (5), pp. 519-525, figs. 1-12. FURMAN, D. P., AND TIPTON, V. J. 1958. Tur uniscutatus (Turk) 1946 (Acarina: Laelaptidae) from neotropical ro- dents. Jour. Parasit., 44, (5), pp. 541-547, pis. I-II. 1961. Acaros parasites laelaptine (Acarina: Laelaptidae) de Venezuela. Mem. Soc. Cienc. La Salle, 21, (60), pp. 166-212, pis. I-XII. HIRST, STANLEY 1915. On some new acarine parasites of rats. Bull. Ent. Res., 6: 183-190, figs. 1-8. KOCH, C. L. 1835-44. Deutschlands Crustaceen, Myriapoden und Arachniden. Ein Beitrag zur deutschen Fauna. Herausgegeben von Herrich-Schaffer. 40 parts. MORLAN, H. B. 1951. Notes on the genus Gigantolaelaps and description of a new species, Giganto- laelaps cricetidarum (Acarina: Laelaptidae). Jour. Parasit., 37: 273-279, pis. I-II. STRANDTMANN, R. W. 1949. The blood-sucking mites of the genus Haemolaelaps (Acarina: Laelaptidae) in the United States. Jour. Parasit., 35, (3), pp. 325-352, figs. 1-45, tab. 1. TIPTON, V. J. 1960. The genus Laelaps with a review of the Laelaptinae and a new subfamily Alphalaelaptinae (Acarina: Laelaptidae). Univ. Calif. Publ. Ent., 16, (6), pp. 233-356, pis. 22-47. TURK, F. A. 1947. A new genus and two new species of mites parasitic on Muridae. Ann. Mag. Nat. Hist., (11), 13, (1946), pp. 347-354, figs. 1-2. 1950. Studies of Acari. — VI. Parasitic mites from mammalian hosts obtained in Ceylon. Parasitology, 40, (1-2), pp. 63-76, figs. 1-15. PLATE 1. ECTOPARASITES OF PANAMA Echinolaelaps lowei Tipton, new species, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 2. Eubrachylaelaps jamesoni Furman, female. 1, dorsal view. 2, gnathosoma. 3, anal plate. 4, ventral view. PLATE 3. ECTOPARASITES OF PANAMA A.MflAZOE Coxa I and sternal plates. Gigantolaelaps gilmorei Fonesca (figs. 4, 8). G. goyanensis Fonseca (figs. 1, 6). G. oudemansi Fonseca (figs. 2, 7). G. wolffsohni (Oudemans) (figs. 3, 5,9, 10). TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 4. Gigantolaelaps gilmorei Fonseca, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 5. ECTOPARASITES OF PANAMA Gigantolaelaps goyanensis Fonseca, female. 1, dorsal view. 2, gnathosoma. 3, anal plate. 4, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 6. Gigantolaelaps inca Fonseca, female. 1, dorsal view. 2, chela. 3, gnathosoma. 4, anal plate. 5, ventral view. PLATE 7. ECTOPARASITES OF PANAMA Gigantolaelaps oudemansi Fonseca, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 8. Gigantolaelaps wolffsohni (Oudemans), female, group A, from Oryzomys caliginosus, Cerro Campana. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral PLATE 9. ECTOPARASITES OF PANAMA Gigantolaelaps wolffsohni (Oudemans), female, group B, from Oryzomys capita El Valle. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 10. Gigantolaelaps wolffsohni (Oudemans), female, group C, from Sigmodon hispidus, Al- mirante. 1, dorsal view. 2, gnathosoma. 3, chela. 4, ventral view. PLATE 11. ECTOPARASITES OF PANAMA Gigantolaelaps wolffsohni (Oudemans), female, group D, from Oryzomys fulvescens, Rancho Mojica. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN I LAELAPTINE MITES PLATE 12. Haemolaelaps glasgowi (Ewing), female, from Peromyscus nudipes, Rancho Mojica, and Reithrodontomys sumichrasti, Cerro Punta. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 13. ECTOPARASITES OF PANAMA Haemolaelaps glasgowi (Ewing), female, from Nyctomys sumichrasti, Cerro Azul. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 14. Haemolaelaps glasgowi (Ewing), female, from Liomys adspersus, Canal Zone. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 15. ECTOPARASITES OF PANAMA Haemolaelaps glasgowi (Ewing), female, from Sigmodon hispidus, Canal Zone. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 16. Haemolaelaps glasgowi (Ewing) , female, from Oryzomys fulvescens, Canal Zone. 1, dor- sal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 17. ECTOPARASITES OF PANAMA Haemolaelaps glasgowi (Ewing), female, from Hoplomys gymnunis, Cerro Azul. 1, dor- sal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 18. 50 NO be Haemolaelaps glasgowi (Ewing), female, from Philander opossum, Canal Zone. 1, dor- sal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 19. ECTOPARASITES OF PANAMA Haemolaelaps glasgowi (Ewing), female, from Sciurus granatensis, Canal Zone. 1, dor- sal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN I LAELAPTINE MITES PLATE 20. Haemolaelaps glasgowi (Ewing), female, from Oryzomys capito, Cerro Azul. 1, dorsal plate. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 21. ECTOPARASITES OF PANAMA Haemolaelaps glasgowi (Ewing), female, from Nectomys alfari, Cerro Azul. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 22. Haemolaelaps glasgowi (Ewing), female, from Metachirus nudicaudatus, Canal Zone. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 23. ECTOPARASITES OF PANAMA Haemolaelaps glasgowi (Ewing), female, from Metachirus nudicaudatus, Cerro Azul. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN I LAELAPTINE MITES PLATE 24. Laelaps castroi Fonseca, female. 1, dorsal view. 2, gnathosoma. 3, coxa I. 4, anal plate. 5, ventral view. PLATE 25. ECTOPARASITES OF PANAMA Laelaps dearmasi Furman and Tipton, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 26. Laelaps dearmasi Furman and Tipton, male. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 27. ECTOPARASITES OF PANAMA r*" Laelaps paulistanensis Fonseca, female. 1, dorsal view. 2, chela. 3, gnathosoma. 4, anal plate. 5, coxa I. 6, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 28. sonobe. Laelaps pilifer Tipton, new species, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 29. ECTOPARASITES OF PANAMA Laelaps thori Fonseca, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 30. Coxa I and anal plate. Laelaps dearmasi Furman and Tipton, female (figs. 4, 7), male (figs. 6, 9). L. pilifer Tipton, new species, female (figs. 5, 11). L. thori Fonseca, female (figs. 1, 8). Laelaps sp., female (figs. 2, 10). Tur anomalus Tipton, new species, female (figs. 3, 12). PLATE 31. ECTOPARASITES OF PANAMA Mysolaelaps parvispinosus Fonseca, female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 32. Steptolaelaps heteromys (Fox), female. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. PLATE 33. ECTOPARASITES OF PANAMA Steptolaelaps heteromys (Fox), male. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 34. Tur anomalus Tipton, new species, female. 1, dorsal view. 2, gnathosoma. 3, anal plate. 4, ventral view. 5, chela (drawn from photograph). PLATE 35. ECTOPARASITES OF PANAMA Tur uniscutatus (Turk), female, from Proechimys semispinosus, Canal Zone. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. TIPTON, ALTMAN, AND KEENAN : LAELAPTINE MITES PLATE 36. Tur uniscutatus (Turk), female, from Hoplomys gymnurus, Cerro Azul. 1, dorsal view. 2, gnathosoma. 3, chela. 4, anal plate. 5, ventral view. The Dermanyssid Mites of Panama ( Acarina : Dermanyssidae ) CONRAD E. YUNKERX AND FRANK J. RADOVSKY2 The dermanyssid mites of Panama have previously been known only from a single record of Ornithonyssus bursa (Berlese) , published by Ewing (1922). Intensive sampling of these blood-sucking parasitic mites was ac- complished by the senior author from 1960 to 1962 in conjunction with a survey of acarine-borne diseases. Collections were made from over 3000 Panamanian vertebrates, mostly mammals. Hosts were collected by shoot- ing or trapping. A large percentage of the bats were caught in mist nets. Parasites were taken either by combing carcasses that had been kept over- night in individual plastic bags, or from pans of water where they had dropped from carcasses suspended overnight by strings above the pans. Many hosts, but not all, were sampled for intranasal mites by a nasal wash- ing technique (Yunker, 1961). As a result, 41 species of Dermanyssidae were found. At least 11 of these are new species, of which only four are described here. The remaining seven, all from bats, will be described in a revision of world-wide species of bat-infesting dermanyssids (Radovsky, in ms.) . A new genus is erected for one of the new species described here, and Neoichoronyssus dentipes is transferred to another new genus. Keys to the species from Panama and a host list are provided. Of particular interest is the absence of dermanyssid mites from rice rats (Oryzomys spp.). Over 100 rice rats representing at least seven spe- cies were examined, but dermanyssid mites were never found. Neither do Strandtmann and Wharton (1958) list dermanyssids for Oryzomys. Host specificity may account for this lack of association. On the other hand, 1 Department of Health, Education, and Welfare, Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Middle America Research Unit, Balboa Heights, Canal Zone, and Rocky Mountain Laboratory, Hamilton, Montana. 2 Department of Entomology and Parasitology, University of California, Berkeley. Present address: George Williams Hooper Foundation, The University of California Medical Center, San Francisco. 83 84 ECTOPARASITES OF PANAMA infestation with dermanyssids may be precluded by the presence of species of Gigantolaelaps, the predominant mesostigmate mites on Panamanian Oryzomys. The disease survey that permitted collateral collection of most of the mites reported here was originated by Dr. James M. Brennan, Rocky Moun- tain Laboratory, Hamilton, Montana. Dr. Alexis Shelokov, Chief, Labora- tory of Tropical Virology and Dr. Henry Beye, Director, Middle America Research Unit, provided administrative support for the survey. Maj. Vernon J. Tipton and Mr. Charles M. Keenan, Environmental Health Branch, United States Army Caribbean, generously facilitated our collect- ing and on occasion provided field men, Pantaleon Sanchez, Vicente Alvarez, Victor Barria, and Wilbur Lowe, who worked long and hard and took an active interest in our program. Our own assistants, Belgica Rodriquez R. and Angel Munoz, Middle America Research Unit, enthusiastically and care- fully performed the laborious preparatory work involved in such a program. Mammals were identified by Dr. Charles O. Handley, and Panamanian birds by Dr. Alexander Wetmore, both of the United States National Museum. Reptiles were identified by Mr. Hymen Marx, Chicago Natural History Mu- seum. Dr. Edward W. Baker, United States National Museum, Dr. Gordon M. Clark, Rocky Mountain Laboratory, Mr. Pedro Galindo, Gorgas Memorial Laboratory, and Dr. Jesse S. White, Delta State Teachers' College, Cleveland, Miss., provided additional material for study. Dr. Joseph H. Camin, Uni- versity of Kansas, Lawrence, examined Draconyssus belgicae and gave help- ful advice. We are grateful to all of these persons. KEY TO THE PANAMANIAN GENERA OF DERMANYSSIDAE FEMALES 1. Dorsal shield divided; metasternal setae absent; on birds Pellonyssus Clark and Yunker Without this combination of characters 2 2. Always on bats 6 On rodents, birds, or reptiles ; never on bats 3 3. At least some coxae bearing non-setigerous ventral spurs 4 Coxae without ventral spurs or coxae I or III with a setigerous spur or pedicel 5 4. Spur of coxa I bifid and setigerous Acanthonyssus n. gen. Spur of coxa I neither setigerous nor bifid Hirstionyssus Fonseca 5. With a single dorsal shield; on rodents, marsupials or birds. . . .Ornithonyssus Sambon With an anterodorsal prosomal shield and a posterodorsal pygidial shield or cluster of platelets; intranasal parasites of lizards Draconyssus n. gen. 6. Dorsal shield divided Steatonyssus Kolenati With a single dorsal shield 7 7. Caudal setae peg-like and barbed; sternal plate with band-like posterior thicken- ing; (protonymphs — the stage most often collected — differing from all others on bats in having claws of leg II much larger than those on other legs) Ichoronyssus Kolenati (group I) All setae smooth ; sternal plate without posterior band 8 8. Anterior margin of coxa II bearing one bifid spur or two simple spurs Radfordiella Fonseca Anterior margin of coxa II with one simple spur 9 9. Legs' stout, leg I more so than leg II ; with ridge-like ventral elevations on coxae I-IV ; palpal trochanter without process New genus no. 1 YUNKER AND RADOVSKY : DERMANYSSID MITES 85 Legs moderate in thickness; leg I no stouter, usually more slender, than leg II ; with ventral elevations on coxae II-IV but not on coxa I ; palpal trochanter with ven- tral spur or ridge 10 10. Chelae edentate; palpal trochanter with ventral spur arising distally Ichoronyssus Kolenati (group II) Fixed chela with two slender, curved, ventral teeth; palpal trochanter with ridge- like process arising along most of its length. . . .Ichoronyssus Kolenati (group III) Genus Pellonyssus Clark and Yunker Pellonyssus Clark and Yunker, 1956, Proc. Helminth. Soc. Wash., 23:93. Type-species: Pellonyssus passeri Clark and Yunker, 1956. KEY TO NEW WORLD SPECIES FEMALES 1. Peritreme long, reaching to mid-level of coxa I; anteromarginal spur of coxa II absent P. marui n. sp. Peritreme short, not reaching past mid-level of coxa II; anteromarginal spur of coxa II present 2 2. First sternal setae short (<15 /j.) ; sternal plate crescentic, nine or 10 times wider than long P. passeri Clark and Yunker First sternal setae long ( >40 /*) ; sternal plate rectangular, not more than six times wider than long P. gorgasi n. sp. Pellonyssus marui, new species. Figure 1E-H. DIAGNOSIS: Similar to P. passeri (fig. 1A-D) ; differing in size (smaller), in having a relatively longer sternal plate, in lacking an antero- marginal spur on coxa II and in the longer peritreme. DESCRIPTION, HOLOTYPE FEMALE: Partially engorged; body approximately 330 n wide at stigmata, 510 /* long exclusive of gnathosoma; dark brown in life. Venter. — Tritosternum with two pilose laciniae. Sternal plate a short, wide band about 20 fi long by 165 n wide. First pair of sternal setae about 19 /u. long, second pair more than twice as long (45 fj.) and third pair nearly three times as long (58 /u). Sternal plate with two pairs of slit-like pores. Third and fourth pairs of pores inserted on small circular platelets on integument; the former near coxae II, the latter near coxae III. Two similar pairs of pores on circular platelets seen ventrally on opisthosoma. Meta- sternal s'etae absent. Epigynial plate pointed posteriorly, reaching past fourth coxae; with a single pair of setae (26 /*) . Anal plate ovoid, with flat anterior margin. Anal opening in anterior part of plate. Paired adanal setae (32 /u) arising on either side of anal opening just posterior to its midpoint. Single postanal seta shorter than adanals (20 /j.) and arising anterior to cribrum. Peritremal plate posteriorly embracing coxa IV, anteriorly continuing on venter to humeral region, proceeding dorsally for a short distance at level with coxa I. Peritreme about 230 p long, arising posteriorly in stigma at level of coxa IV and terminating anteriorly at mid-level of coxa I. Postcoxal apodeme III continuous with peritremal plate and extending into the lateral vaginal wall as spur- shaped apodeme. Opisthosoma with 13 pairs of setai that increase in size posteriorly (f^om 25 fj. at level of epigynial plate to 70 p. at end of idiosoma). Dorsum. — Dorsal shield divided, both segments large, covering most of idiosoma. Anterior (prosomal) shield about 220 n wide at posterior end, 220 ^ long, roughly tri- angular; posterior margin straight; surface reticulate, bearing nine pairs of setae: five lateral and four submedian, the former about 23 /u. long, the latter about 17 /«. long, and a single pair of anterior, lyriform pores. A pair of minute vertical setae (5 n) on integument just anterior to shield. Posterior shield slightly narrower than base of an- terior shield, about 170 /u wide at anterior end, 260 /j, long. Anterior margin straight, lateral margins tapering to a blunt point. Shield strongly reticulate, with six pairs 86 ECTOPARASITES OF PANAMA of setae: the three posterior pairs submarginal, increasing in size posteriorly (13 n- 17 /J.-26 /*) , the three anterior pairs submedian, about 15-17 /u long. Dorsal integu- ment with 27-28 pairs of setae (including verticals), those posterior and lateral heavier and longer (e.g., 48 /*) than those anterior and median (e.g., 19 /*). Gnathosoma. — Chelicerae with elongate, attenuate apical segment (210 //, long by 7-10 n wide), and shorter wider basal segment (93 /u, long by 14 // wide) ; terminating in small, shear-like chelae. Chelae with two recurved teeth on movable digit, and a membranous fringe on fixed digit. Legs. — All legs with caruncles and paired claws. Coxa II without obvious antero- marginal spur. MALE: Body approximately 270 ^ wide at stigmata, 430 //, long exclusive of gnatho- soma. Holoventral plate irregular in outline, approximately parallel-sided between coxae, widening slightly posterior to coxae IV, markedly constricted at beginning of anal plate; surface reticulate, with six to eight pairs of setae in addition to three setae on anal plate. First sternal seta3 approximately 5 /j. long, second and third approx- imately 23 /j. long; metasternal setae absent; genital setae approximately 20 /* long; two to four pairs of ventral integumental setae of size similar to that of genital setae; paired adanal setae (30 /a) arising on either side of posterior of anal opening, shorter (21 /j.) postanal seta arising anterior to cribrum. Ventral and ventrolateral aspects of unsclerotized venter with eight to ten pairs of setae, the lateral and terminal setae heavier and longer (e.g., 46 AC) than the ventral setae (e.g., 20 p) . Peritreme arising at stigma located at level of coxa IV, curving regularly to anterior level of coxa II; approximately 155 /u long. Tritosternum with two laciniae, pilose from base anteriorly. Coxa II without obvious anterodorsal spur. Dorsal shield entire, elongate and narrow (496 (j. long by 225 /* wide), tapering to blunt point, surface reticulate, with 12 pairs of setae. Dorsum with 15-17 pairs of non-plate setae, of sizes similar to corresponding setae on venter. Chelicerae shorter and stouter than in female, basal segment expanded, remainder elongate and narrow, length of apical segment, including chelae, 130 //.. Sper- matodactyl 19 /a long, bifurcate, with both members of nearly equal length, the thicker member trough-like. Fixed chela shorter and narrower than spermatodactyl. Deuto- sternum with eight teeth in single file. PROTONYMPH (unfed) : Body approximately 296 /* long by 185 n wide. Sternal plate oval, 111 ft, long by 93 /* wide; with three pairs of setae, each 25 ^ long. Anal plate oval, 67 /j. long by 56 n wide, with a pair of adanal setae (37 /u) and a single postanal seta (19 /n). Anus near anterior margin of plate. Five pairs of non-plate setae on venter. Peritreme originating in stigma at level of coxa IV and extending anterior to level of middle of coxa III, approximately 37 /a long. Coxa II without obvious antero- marginal spur. Tritosternum with two indistinctly pilose laciniae. Dorsum with four shields: one large prosomal (177 M long by 179 ^ wide), one smaller pygidial (37 /t long by 92 /j. wide) and two minute irregular platelets (18 /u, diam.) just posterior to prosomal plate. Prosomal shield with nine pairs of setae, each 10 /u long; a pair of minute verti- cal setae on integument just anterior to shield. Pygidial shield with three pairs of setae, the anteriormost pair minute (7 //. long), the other two pairs heavier and longer (48 /it and 60 ^). Chelicera functional, similar in form to that of female. Apical seg- ment, including chelae, approximately 150 /A long. TYPE MATERIAL: Holotype female (U.S.N.M. no. 66608), 4 paratype females, 1 paratype protonymph and 2 paratype males from nest of Cassidix mexicanus, Ancon (Canal Zone), 10 May 1961, collected by C. E. Yunker and N. Smith; 17 paratype females, 11 paratype protonymphs and 2 para- type males from Vireo flavoviridis, and 7 paratype females from Turdus grayi, Carrasquilla (Panama), 3 June 1961, collected by A. Munoz; 2 paratype females from Progne chalybea, Corozal ice plant (Canal Zone), 11 April 1961, collected by C. M. Keenan; 4 paratype females, 2 paratype males and 1 paratype protonymph from nest and young of Columbigallina YUNKER AND RADOVSKY : DERMANYSSID MITES 87 talpacoti, Corozal (Canal Zone) , 26 June 1961, collected by C. E. Yunker and A. Munoz. Paratype specimens to be distributed among the collections of the fol- lowing institutions : United States National Museum ; Chicago Natural His- tory Museum ; British Museum (Natural History) , London ; Rocky Mountain Laboratory, Hamilton, Montana; Institute of Acarology, Ohio State Uni- versity, Columbus; Department of Entomology and Parasitology, Univer- sity of California, Berkeley; Snow Entomological Museum, University of Kansas, Lawrence ; South African Institute for Medical Research, Johannes- burg. Fig. 1. Pellonyssus passeri Clark and Yunker. A, female sternal plate. B, female coxa II. C, female anal plate. D, male chelae. Pellonyssus marui, new species. E, female sternal plate. F, female coxa II. G, female anal plate. H, male chelae. Pellonyssus gorgasi, new species, female. I, sternal plate ; J, coxa II ; K, anal plate. ADDITIONAL MATERIAL EXAMINED : Pellonyssus marui was also seen from neotropical areas other than Panama : 23 females and 1 male from Pitangus sulfuratus, Rio de Janeiro, Brazil, 29 December 1948, collected by H. W. Krumm ; 3 females from "Gray kingbird," Little Inagua Island, British West Indies, "8-5-30," collected by H. S. Peters ; 20 females, 42 protonymphs and 15 males from Coereba portoricensis, Mayaquez, Puerto Rico, 28 March 1948, collected by Virgilio Biaggi. 88 ECTOPARASITES OF PANAMA Pellonyssus gorgasi, new species. Figure 1 I-K. DIAGNOSIS : Differing from P. passeri and P. marui in possessing elongate sternal setae I (nearly twice as long as plate) ; differing further from P. marui in having an anteromarginal spur on coxae II and a short peritreme. DESCRIPTION, HOLOTYPE FEMALE: Damaged; body approximately 336 /a wide at stig- mata, 580 fjL long exclusive of gnathosoma. Venter. — Tritosternum with two pilose laciniae. Sternal plate approximately rec- tangular, about 20 /u long by 113 n wide. First pair of sternal setae 43 /j. long, second pair 58 /j., and third pair 68 //.. Only the first pair of sternal pores evident on sternal plate of the holotype, lyriform; second sternal pores not seen. Third pair of pores cir- cular, on soft integument near coxae II; a fourth pair similar to third pair on soft integument near coxae IV. Metasternal setae absent. Epigynial plate pointed poste- riorly, reaching past fourth coxae; with a single pair of setae (35 /*). Anal plate broadly ovoid, with a truncate anterior margin. Anal opening in anterior of plate; paired adanal setae (26 /*) arising at level of posterior of anal opening. Single postanal seta (23 /n) arising anterior to cribrum. Peritremal plate posteriorly embracing coxa IV, anteriorly terminating at anterior level of coxa II. Peritreme about 154 /* long, posteriorly arising in stigma between coxae III and IV and anteriorly terminating at mid-level of coxa II. Postcoxal apodeme III continuous with peritremal plate and ex- tending into lateral vaginal wall as spur-shaped apodeme. Opisthosoma with 22-24 pairs of setae that increase in size posteriorly (from 34 n at level of epigynial plate to 50 fj. at end of idiosoma). Dorsum. — Dorsal shield divided, both segments large, covering most of idiosoma. Anterior (prosomal) shield 255 /* wide at posterior end, 280 /u long, roughly in the shape of a triangle; posterior margin straight; surface reticulate, bearing nine pairs of setae: five lateral and four submedian, the former 20-25 n long, the latter 10-15 /* long, and one pair of anterior lyriform pores. A single pair of minute (<10 M) vertical setae on integument just anterior to shield. Posterior shield slightly narrower than base of anterior shield, about 220 ^ wide at anterior end, 270 /j. long. Anterior margin con- cave. Posterior of shield terminating in blunt point; lateral margins not tapering evenly to this point, but indented near posterior end. Shield reticulate, only five pairs of setae visible (but pygidial area poorly cleared) : the two posterior pairs marginal, the terminal pair longer (16 /*) than the subterminal (10 /*) ; the three anterior pairs submedian, about 12 p long. Dorsal integument with 30-34 pairs of setae (including verticals), those posterior and lateral heavier and longer (e.g., 50 /*) than those anterior and sublateral (e.g., 18 /u). Gnathosoma. — Chelicerae with elongate, attenuate apical segment (233 /x long by 10 yu wide), and shorter, wider basal segment (18 /j. long by 20 /* wide). Chelae poorly cleared ; shear-like and apparently edentate. Legs. — All legs with caruncles and paired claws. Coxa II with an elongate antero- marginal spur. TYPE MATERIAL: Holotype female (U.S.N.M. no. 66607) from Phae- thornis guy, Cerro Punta (Chiriqui), about 5200 feet elevation, 27 April 1961, collected by C. E. Yunker. Known only from the type specimen. Genus Ornithonyssus Sambon Ornithonyssus Sambon, 1928, Ann. Trop. Med. Parasit., 22: 105. Type-species: Dermanyssus sylviarum Canestrini and Fanzago, 1878. KEY TO PANAMANIAN SPECIES FEMALES 1. Proximal seta of coxa I arising from a spur-like elevation. . . .O. wernecki (Fonseca) Proximal seta of coxa I not pedicillate 2 YUNKER AND RADOVSKY : DERMANYSSID MITES 89 2. Sternal plate with two pairs of setae 3 Sternal plate with three pairs of setae 4 3. Medial and sublateral setae of dorsal shield short, not over 25 p.; on birds O. sylviarum (Canestrini and Fanzago) All dorsal shield setae long, over 35 /*; on porcupines. . .Ornithonyssus coendou n. sp. 4. Setae of dorsal shield shorter than those on dorsal integument; on birds O. bursa ( Berlese) Setae of dorsal shield at least as long as those on dorsal integument ; on rodents .... 0. bacoti ( Hirst) Ornithonyssus sylviarum (Canestrini and Fanzago) Dermanyssus sylviarum Canestrini and Fanzago, 1878, Atti R. Istit. Veneto Sci. Lett. Arti, (5), 4:124. The northern fowl mite is a common, serious, blood-sucking parasite of passeriform birds and domestic fowl in temperate regions throughout the world. Thirteen females were taken from a toucan, Aulacorhynchus prasinus, in Cerro Punta (Chiriqui), at approximately 5200 feet altitude. This is apparently the southernmost New World record for O. sylviarum, previously not recorded south of the United States. It is noteworthy, how- ever, that it has not yet been found in the tropical lowlands. This parasite should be searched for in other areas of Panama. Ornithonyssus bacoti (Hirst) Leiognathus bacoti Hirst, 1913, Bull. Ent. Res., 4: 122. The tropical rat mite is a common blood-sucking parasite of many species of rodents throughout the world, and is capable of attacking birds and many mammals including man. In Panama, it was the most common dermanyssid encountered and was taken during all months of the year. Preferred hosts were Proechimys semispinosus and Sigmodon hispidus. In the higher ele- vations of Chiriqui Province, it was collected only from Peromyscus nudipes. Although taken on nine occasions from Liomys adspersus, a spiny pocket mouse, it was never seen on Heteromys desmarestiamis, a closely related rodent, despite extensive collection of the latter. Other infrequent hosts were Zygodontomys microtinus and Rattus rattus. Localities involved were : both Atlantic and Pacific sides of Canal Zone; Achiote (Colon) ; Las Palmi- tas and Guanico (Los Santos) ; Cerro Punta, Bambito and El Hato (Chiri- qui). Ornithonyssus bursa (Berlese) Leiognathus bursa Berlese, 1888, Boll. Soc. Ent. Italiana, 20: 208. The tropical fowl mite, common on many species of birds in warmer climates of the world, was recorded by Ewing (1922) "on common hen" in the Canal Zone. Subsequent records are lacking, and the species was not encountered in this study. Here, however, birds were not frequently ex- amined. We have seen specimens from domestic fowl in Costa Rica and the species is recorded from Colombia ; there is no reason to doubt its presence in Panama. 90 ECTOPARASITES OF PANAMA Ornithonyssus coendou, new species. Figure 2. DIAGNOSIS: Similar to O. sylviarum, differing: in size (larger), in pos- session of long, subequal setae on dorsal shield, and in having adanal setae that arise at posterior level of anal slit. Known only from the porcupine, Coendou rothschildi. DESCRIPTION, HOLOTYPE FEMALE: Body approximately 425 M wide at stigmata, 670 /x long exclusive of gnathosoma. Venter. — Tritosternum with two pilose laciniae. Sternal plate rectangular, about 36 /it long at mid-line by 92 n wide at sternal setae II ; with two pairs of setae and two pairs of lyriform pores'; surface adjacent to setae ornamented with cell-like reticulations; parts of these reticulate areas covered with punctations. Third sternal setae on integu- ment between sternal plate and epigynial opening. Metasternal setae present, adjacent to coxae III. Third sternal pores located between sternal setae III and metasternal setae. Epigynial plate narrow, pointed posteriorly, broad and fan-shaped anteriorly, with a pair of setae. A pair of circular pores on integument near genital setae. Anal plate ovoid, with the usual three setae and posterior denticulate cribrum. Adanal setae arising at a level with posterior of anus, slightly shorter than postanal seta. Stigma large (21 fj. diam.) circular, located lateral and posterior to coxa III. Peritreme wide, extending anteriorly, bending dorsally, and terminating over mid-region of coxa I. Peritremal plate fusing with endopodal apodeme to nearly encircle coxa IV. Ventral integument with about 30 pairs of non-plate setae, those anterior smallest (e.g., 33 p), those lateral and posterior longest (e.g., 47 ^) • All posteroventral setae distinctly ter- minally branched (fig. 2B). A pair of narrow elongate platelets flanking posterior end of epigynial shield. Metapodal platelets irregular, small, in lateral post-coxal area. Dorsum. — With a single, long shield having sinuous lateral margins and tapering posteriorly to a narrow, rounded point. Shield with 17 pairs of setae, anteriormost pair relatively minute, others elongate (about 40 /*) , terminally branched. Dorsal integument with about 50 pairs of non-plate setae, all well developed, terminally branched and elongate. Those lateral and posterior longest (e.g., 45 /u). Legs. — I and IV subequal in length, longer than II and III, which are also sub- equal. Each leg with a pretarsus bearing ambulacral claws and a small membranous caruncle. All segments without obvious spurs or protuberances, except coxa II, which bears an anteromarginal spur. All but distal leg setae terminally branched, those dorsal somewhat longer and thicker than those ventral, especially on legs I and II. Gnathosoma. — Internal hypostomal setae much longer than external and anterior hypostomal setae and gnathosomal base setae. With nine or 10 deutosternal teeth ar- ranged in a single file. Hypostomal processes elongate. Palpal trochanter with a small ventral protuberance. Chelicerae chelate; chelae edentate, resembling those of other species of Ornithonyssus. MALE: Body approximately 278 ^ wide at stigmata, 450 /* long, exclusive of gnatho- soma. Holoventral plate irregular in outline, with 21 setae (occasionally one member of a pair off plate). With 10 pairs of ventral non-plate setae. Stigma as in female; peritreme extending anteriorly, bending dorsally and terminating over anterior region of coxa II. Dorsal shield broad, covering most of dorsum, tapering posteriorly to a point; with 19-21 pairs of setae similar to those of female. Dorsum with nine or 10 pairs of non-plate setae. Coxa II with usual anteromarginal spur. Palpal femur with a large ventral protuberance bearing a spiniform seta (fig. 2F). Chelicera as in female except for chela, which bears a scapiform spermatodactyl (fig. 2G). PROTONYMPH : Body approximately 520 n long by 325 ^ wide. Sternal plate shield- shaped, with three pairs of setae. Anal plate oval, with three setae. Six pairs of non- plate setae on venter. Peritremalia a short, lateral crescent. Coxa II with a small, blunt anteromarginal spur. Dorsum with prosomal plate having 10 pairs of setae, a pygidial plate having three pairs of setae, and two small, bare platelets just posterior to prosomal plate. Eleven pairs of non-plate dorsal setae. YUNKER AND RADOVSKY : DERMANYSSID MITES 91 Fig. 2. Ornithonyssus coendou, new species. A, female venter. B, female dorsum. C, female chelicera. D, female sternal and epigynial area. E, female anal plate. F, male palpal trochanter. G, male chelae. H, male holoventral plate. 92 ECTOPARASITES OF PANAMA TYPE MATERIAL: Holotype female (U.S.N.M. no. 66610), 1 paratype fe- male and 2 paratype nymphs from Coendou rothschildi, near Pedro Miguel River, Paraiso (Canal Zone), 21 February 1962, by J. M. Brennan and C. E. Yunker, deposited in United States National Museum. Allotype male, and 10 paratype nymphs, same data, but 12 March 1962, in United States National Museum. One paratype female and 7 paratype nymphs, same data, but 26 February to 20 March 1962, in collection of Rocky Mountain Laboratory, Hamilton, Montana. Ornithonyssus wernecki (Fonseca) Liponyssus wernecki Fonseca, 1935a, Mem. Inst. Butantan, 9: 70, figs. 1-8. This species is a common parasite of marsupials in Panama. Hosts from which we collected it are Didelphis marsupialis, Marmosa robinsoni, Phi- lander opossum and Metachirus nudicaudatus. Localities include France Field, Fort Sherman and Fort Gulick (Atlantic side of Canal Zone) ; Cerro Campana and Cerro Azul (Panama). Collected mainly during the drier months, March and April, we also have a collection taken in November, France Field (Canal Zone), from Didelphis marsupialis. Acanthonyssus, new genus Type-species : Ichoronyssus dentipes Strandtmann and Eads, 1947. DIAGNOSIS: Macronyssinae ; all setae smooth; dorsal shield entire and broadly rounded posteriorly; all coxae bearing one or more stout ventral spurs; coxa II with one simple anterodorsal spur; outer ventral spur of coxa I bifid and bearing proximal seta at furcation ; some middle segments of legs II-IV bearing strong recurved spurs. FEMALE: sternal plate short, without surface markings other than usual sculpturing and pores; sternal setae II much closer to III than to I; epigynial plate cuneate, with well-defined scale-like, anterior sculpturing; peritremes moderately broad, terminating over coxa I; chelae simple, without setae, teeth or other processes; palpal trochanter with distal ventral spur. MALE: ventral armature entire; palpal trochanter without process. REMARKS : Furman and Radovsky (1963) synonymized Neoichoronyssus by transferring the type-species, N. ivernecki, to Ornithonyssus. In doing so, they left the generic status of N. dentipes unresolved. This new genus is proposed for N. dentipes. Acanthonyssus dentipes (Strandtmann and Eads), new combination. Ichoronyssus dentipes Strandtmann and Eads, 1947, Jour. Parasit., 33: 31, figs. 1-3. Described from Sigmodon hispidus in Texas, this species was recovered mainly from Proechimys semispinosus in the Canal Zone (Miraflores, Sum- mit, Gamboa, Coco Solo, France Field and Fort Gulick) . It was taken only during the drier months, December to May, 1960-1962. Two collections, however, from Almirante (Bocas del Toro), August 1960, by P. Galindo were from the type host, Sigmodon hispidus. Some of the material from Proechimys showed various but not obviously consistent morphological differences from specimens from the type host. YUNKER AND RADOVSKY : DERMANYSSID MITES 93 These variations were particularly noticeable in the dimensions of the ster- nal plate (range = 23 p. long by 118 //, wide to 32 p. long by 112 p. wide) , the degree of biconcavity of the lateral margins of the dorsal shield, and the number and relative size of the ventral setae. It is possible that two species are involved here. Genus Hirstionyssus Fonseca, 1948 Hirstionyssus Fonseca, 1948, Proc. Zool. Soc. London, 118:266. Type-species: Dermanyssus arcuatus Koch, 1839. Seven species of Hirstionyssus are known from Panama. They are de- scribed and keyed by Strandtmann and Yunker elsewhere in this volume. Of these, all but one are from heteromyid and cricetid rodents. The excep- tion is from squirrels. A list of the species and their hosts is included in the host list of this paper. Draconyssus, new genus Type-species : Draconyssus belgicae, new species. DIAGNOSIS: Dermanyssidae ; with two dorsal shields or with a single prosomal shield and a cluster of pygidial platelets; second cheliceral seg- ment extremely elongate, but not attenuate, at rest deeply withdrawn into idiosoma ; sternal plate with two pairs of setae ; metasternal setae absent ; epigynial setae off plate; peritreme extending to level of middle of coxa II. Male unknown. REMARKS : At this point we are unable to assign Draconyssus to a sub- family within the Dermanyssidae. We suspect it to be macronyssid and to have affinities with Ophionyssus and Sauronyssus. Cheliceral conformation — the sole basis for subfamilial placement — is not applicable here. The chelicerae of Draconyssus belgicae, n. sp., although extremely elongate and withdrawn into the idiosoma as in the Dermanyssinae, are also strongly chelate and definitely not attenuate, as in the Macronyssinae. A similar, but not as pronounced, elongation of the chelicerae is seen in Pellonyssus and Ophionyssus (Macronyssinae). Hystrichonyssus (Hystrichonyssinae) shows elongate attenuate chelicerae, but here the proximal segment is elon- gate whereas the second segment is normal. It has been obvious to us for some time that cheliceral modifications are unsatisfactory indicators of phy- logeny in this group. Until a better basis for classification is reached, Dra- conyssus must remain as an unassigned genus of Dermanyssidae. Draconyssus belgicae, new species. Figure 3. DESCRIPTION, HOLOTYPE FEMALE: Body approximately 405 /j, wide at stigmata, 668 /* long exclusive of gnathosoma. Venter. — Tritosternum with two pilose laciniae and serrate lateral membranes. Ster- nal plate roughly rectangular with irregular borders, about 54 fj. long at mid-line by 93 /j, wide at sternal setae II ; with two pairs of setae and two pairs of lyrif orm pores ; surface densely covered by coarse punctations that become less dense posterior to sternal setae II. Third sternal setae on integument between sternal plate and epigynial opening. Metastrrnal setae absent. Epigynial setae on unsclerotized integument be- tween lateral margins of epigynial plate and coxae IV. Sternal and epigynial setae 94 ECTOPARASITES OF PANAMA subequal in length. Third sternal pores circular, on integument just posterior to sternal setae III, and overlapped by hyaline anterior margin of epigynial plate. A fourth pair on unsclerotized integument just posterior of epigynial setae and three similar pairs on hysterosoma. Epigynial plate narrow, pointed posteriorly, broad and fan-shaped anteriorly, and biconcave laterally in region of coxae IV. Anal plate ovoid, with the usual three setae and posterior denticulate cribrum; adanal setae arising at a level with posterior of anus, subequal in length to postanal seta. Peritreme arising at stig- mata located lateral to and between coxae III and IV, extending anteriorly and be- coming dorsal at termination over mid-region of coxa II ; peritremal plate fusing with endopodal apodeme to nearly encircle coxa IV. Ventral integument with 15 or 16 pairs of non-plate setae; those lateral and posterior longest (about 53 /j.) ; those medial and anterior of anal plate shorter, subequal to sternal and epigynial setae (about 37 fi). All ventral setae terminally branched. A pair of narrow elongate platelets flanking posterior end of epigynial shield. Metapodal platelets irregular crescents, located in lateral post-coxal area. Dorsum. — With two sclerotized shields. Propodosomal shield biconvex ; twice longer than wide; with six pairs of setae. Pygidial shield minute, irregular in outline, with- out setae. Dorsal integument with 25 or 26 pairs of non-plate setae, including verticals. All dorsal setae well developed, terminally branched, and subequal (about 50 /j, long). Two pairs of circular pores dorsal on hysterosoma. Legs. — I and IV subequal in length, longer than II and III, which are also sub- equal. Each leg with a pretarsus bearing ambulacral claws and membranous caruncles. All segments without obvious spurs or protuberances. Leg setae as in body setae, ter- minally branched; those dorsal somewhat longer and thicker than those ventral, espe- cially on legs I and II. Gnathosoma. — Internal hypostomal setae much longer than external and anterior setae and gnathosomal base setae. With eight or nine deutosternal teeth arranged in a single file. Hypostomal processes elongate, ensheathed by lateral folds of tectum. Tectum forming an elongate tube, extending to level with base of palpal tarsi. Chelicerae elongate and whip-like. Basal segment short (21 /j.) ; distal segment sinuous and elongate (363 n). Chelae chelate and edentate, the movable digit somewhat shorter and stouter than the fixed digit; the latter with a slight, recurved tip (fig. 3C). TYPE MATERIAL: Holotype female (U.S.N.M. no. 66609) intranasal in Ameiva bifrontata, a teiid lizard, Nuevo Emperador (Canal Zone), 10 Au- gust 1961, collected by C. E. Yunker and A. Mufioz. Three paratype females bearing the same data as the holotype ; 3 paratype females intranasal in Ameiva sp., K-9 Road (Pacific side of Canal Zone), 18 October 1961, col- lected by C. E. Yunker and A. Mufioz. The holotype and a paratype will be deposited in the United States National Museum ; remaining paratypes will be distributed among the collections of Chicago Natural History Museum ; Rocky Mountain Laboratory, Hamilton, Montana; Institute of Acarology, Ohio State University, Columbus ; Snow Entomological Museum, University of Kansas, Lawrence. REMARKS: Variation was seen in the shapes of the dorsal shields and ventral plates, which were often eroded and irregular. The pygidial shield of certain specimens was represented only by a cluster of two or three ir- regular platelets. The epigynial plate varied in outline from an irregular, asymmetrical linguiform one, to one with perfectly parallel sides ending posteriorly in a glans-shaped expansion. The outline of the dorsal shield was occasionally irregular, and in one specimen it included the bases of two lateral setae typically found on the bare integument. The sternal shield was often seen to be eroded on its posterior margin. YUNKER AND RADOVSKY : DERMANYSS.ID MITES 95 Fig. 3. Draconyssus belgicae, new genus, new species, female. A, venter. B, dorsum. C, chelicera. D, sternal plate. 96 ECTOPARASITES OF PANAMA The species is named for Miss Belgica E. Rodriquez R., Middle America Research Unit, who first recovered specimens from nasal washings. Genus Steatonyssus Kolenati Steatonyssus Kolenati, 1858, Wien. Ent. Monatschr., 2:6. Type-species: Dermanyssus murinus Lucas, 1840 (— Dermanyssus periblepharus Kolenati, 1858). Steatonyssus occidentalis (Ewing) Ceratonyssus occidentalis Ewing, 1933, Proc. U.S. Nat. Mus., 82: 10, pi. 3 (fig. 5), pi. 4 (fig. 1). This is the only species of this large, widely distributed genus taken in Panama. It was recovered from an unidentified bat in El Valle (Code), 1 June 1961, collected by W. E. Woodcock. In North America, the usual host is Eptesicus fuscus; the range of this bat includes Panama. Genus Ichoronyssus Kolenati Ichoronyssus Kolenati, 1858, Wien. Ent. Monatschr., 2: 5. Type-species: Ichoronyssus scutatus Kolenati, 1858. The interpretation of Ichoronyssus followed in this paper is that of Strandtmann and Wharton (1958). It will be reinterpreted in a revision by the junior author (in manuscript) . The genus (sensu Strandtmann and Wharton) is here divided into three groups of related species. Group I KEY TO PANAMANIAN SPECIES FEMALES 1. Third pair of sternal setae on platelets joined to sternal plate by thread-like connec- tions /. robustipes ( Ewing) At most, slight constrictions between portions bearing sternal setae III and re- mainder of plate 2 2. Sternal plate without constrictions proximal to third pair of setae; anteromedial setae on dorsal shield about 40 fj. or more 7. venezolanus (Vitzthum) Sternal plate with slight constrictions proximal to third pair of setae; antero- medial setae on dorsal shield about 25 ^ or less. . . ./. haematophagus (Fonseca) PROTONYMPHS 1. Coxa I with blunt lateral spur /. venezolanus (Vitzthum) Coxa I without lateral spur 2 2. Unarmed venter with five pairs of setae lateral or anterior to anal plate /. haematophagus (Fonseca) Unarmed venter with seven pairs of setae lateral or anterior to anal plate 7. robustipes ( Ewing) Ichoronyssus robustipes (Ewing) Liponyssus robustipes Ewing, 1925, Ent. News, 36: 20. This species is a common parasite of Tadarida brasiliensis , a bat ranging over much of North and South America and the West Indies. In Panama, /. robustipes was thrice taken from this host in a cave in Cerro Punta YUNKER AND RADOVSKY : DERMANYSSID MITES 97 (Chiriqui), 3 April 1961, by C. E. Yunker. Several mites also were taken from Myotis nigricans roosting in the same cave. Ichoronyssus venezolanus (Vitzthum) Liponyssus venezolanus Vitzthum, 1932, Zeitschr. Parasitenk., 4: 9, figs. 3-13. Previously known from the type collection from Molossus nasutus in Venezuela, protonymphs of I. venezolanus were collected from M. coibensis and "molossid bats" in a church attic in Pacora (Panama), 6 June 1961, by C. M. Keenan and C. E. Yunker. Ichoronyssus haematophagus (Fonseca) Liponyssus haematophagus Fonseca, 1935b, Mem. Inst. Butantan, 10: 43, figs. 1-2. Protonymphs of this species were taken from a molossid bat, probably Molossus coibensis, in a church attic in Pacora (Panama), 6 June 1961, by C. M. Keenan and C. E. Yunker. The type collection is from southern Bra- zil, from Molossus rufus. Group II KEY TO PANAMANIAN SPECIES FEMALES Sternal satae III arising from posterior angles of sternal shield, and shield with slight, neck-like constrictions proximal to these setae; dorsal shield with 26 or 27 pairs of setae /. kochi Fonseca Sternal setae III arising slightly anterior to posterior angles of shield, not set apart by constrictions ; dorsal shield with 24 pairs of setae New species no. 1 Ichoronyssus kochi Fonseca Ichoronyssus kochi Fonseca, 1948, Proc. Zool. Soc. London, 118: 278, figs. 17-20. The type host is Artibeus sp. from Brazil, and our records indicate that Artibeus is the usual host genus. In Panama, /. kochi was taken from Artibeus toltecus, Cerro Punta (Chiriqui), April 1961, and Rio Changena (Bocas del Toro), September 1961 ; from Artibeus jamaicensis, Juan Mina (Canal Zone), June 1961, and Rio Changena (Bocas del Toro), September 1961, by C. E Yunker. Ichoronyssus (group II), new species no. 1 This species was found in Panama on Vampyrops vittatus. Collections were made from several bats at Rio Changena (Bocas del Toro) , September 1961, by V. J. Tipton and C. E. Yunker and from a single bat at Cerro Punta (Chiriqui) , February 1960, by V. J. Tipton. Group III KEY TO PANAMANIAN SPECIES FEMALES Peritreme ending over coxa I; dorsal shield with longest anterolateral setae about two or three times as long as shortest anteromedial setae I. crosbyi (Ewing and Stover) Peritreme ending over coxa II ; dorsal shield with longest anterolateral setae about nine or ten times as long as shortest anteromedial setae New species no. 1 98 ECTOPARASITES OF PANAMA Ichoronyssus crosbyi (Ewing and Stover) Liponyssus crosbyi Ewing and Stover, 1915, Ent. News, 26: 112, pi. 4, fig. 3. In Panama, 7. crosbyi was recovered from Myotis chiloensis and from a mixed lot of M. chiloensis and Myotis nigricans roosting in a cave in Cerro Punta (Chiriqui), 5 May 1961, by C. E. Yunker. At least one bat was in- fested with this species and with Ichoronyssus (group III), n. sp. no. 1. The latter parasite was far more numerous at this site. In the United States, 7. crosbyi is known from several other Myotis species. Ichoronyssus (group III), new species no. 1 Large numbers of this species were found on many individual Myotis nigricans in a cave in Cerro Punta (Chiriqui) , 5 May 1961, by C. E. Yunker. In the same cave, it was taken on a few specimens of Myotis chiloensis. A single collection was taken on M. nigricans in Cerro Punta (Chiriqui), in March 1962, by V. J. Tipton. Genus Radfordiella Fonseca Radfordiella Fonseca, 1948, Proc. Zool. Soc. London, 118: 270. Type-species: Radfordiella oudemansi Fonseca, 1948. KEY TO PANAMANIAN SPECIES FEMALES 1. Spur on anterior margin of coxa II bifid; sternal shield short (length at mid-line about 25 /t), with strongly arched posterior margin R. oudemansi Fonseca Coxa II with two simple spurs on anterior margin ; sternal shield length at mid-line over 50 /j. 2 2. Dorsal shield length over 410 /u and usually more than 430 p.; sternal shield with moderately arched posterior margin reaching slightly beyond level of tricho- pores of sternal setae III New species no. 1 Dorsal shield length under 410 M; sternal shield with weakly arched posterior margin rarely reaching beyond level of pores of sternal setae III . . New species no. 2 Radfordiella oudemansi Fonseca Radfordiella oudemansi Fonseca, 1948, loc. cit., 118: 274, figs. 45-48. R. oudemansi has not been collected in Panama, but it probably occurs there and hence is included in the key. The type material was taken on Desmodus rotundus in Brazil. Radfordiella, new species no. 1 The common vampire bat, Desmodus rotundus, appears to be the normal host. Mites were taken in Panama from a number of individuals of D. rotundus at Las Palmitas (Los Santos), January 1962, by C. 0. Handley and F. Greenwell. We have seen material from the same host in Guatemala and Trinidad. Radfordiella, new species no. 2 This species was collected in Panama as follows: from Carollia per- spicillata, Sardanillo, Summit (Canal Zone), 12 August 1961; in a mixed collection of C. perspicillata and Lonchorhina aurita, same locality and date ; YUNKER AND RADOVSKY : DERMANYSSID MITES 99 from C. perspicillata, Juan Mina (Canal Zone) , 30 June 1961 ; from Carollia castanea, Cerro Pirre (Darien), 3 February 1961, by C. E. Yunker. Only the first of these collections contained more than one specimen. New Genus no. 1 Three undescribed species belonging to to this genus were found in Panama. All of these were taken from phyllostomid bats, as follows : n. sp. no. 1 from Glossophaga soricina, Rio Changena (Bocas del Toro), 19 Sep- tember 1961, by C. E. Yunker; n. sp. no. 2 from Sturnira ludovici same data; n. sp. no. 3 from Carollia, perspicillata, Juan Mina (Canal Zone), 20 June 1961, by C. E. Yunker. We also have specimens of n. sp. no. 1 from a phyllostomid bat collected in Costa Rica by J. S. White. Species inquirendae Ten collections, consisting solely of protonymphs or males were seen. Due to the existing lack of knowledge concerning immature and male meso- stigmates they are not fully identifiable. Two of these, from rodents, are not referable to any of the species known from Panama. They are : Sub- family Dermanyssinae, 2 protonymphs from Hoplomys gymnurus, Cerro Azul (Panama), 17 March 1961; Subfamily Macronyssinae, 4 protonymphs of Ornithonyssus sp. from Dasypus novemcinctus, near Pedro Miguel River, Paraiso, (Canal Zone) , 27 February 1962. The remaining eight collections, all macronyssines from bats, are refer- able to Radfordiella, Ichoronyssus (group II), new genus no. 1, and an un- known genus. These probably all represent new species, but material is inadequate for treatment. Addenda Since this paper was set in type, Till (1964: 90, 92) has synonymized Pellonyssus passeri Clark and Yunker, the type-species of Pellonyssus Clark and Yunker, under Steatonyssus reedi Zumpt and Patterson (1952: 163, fig. 3). A summary reclassification (Radovsky, 1966a), including the bat-parasitizing Der- manyssidae referred to here, has been in press at the same time as the present paper. The subfamily Macronyssinae is treated as a separate family, Macronyssidae, which would include nearly all of the Panamanian Dermanyssidae given in this paper (excep- tion: Dermanyssinae incertae sedis off Hoplomys gymnurus). Ichoronyssus (group I) is interpreted as Chiroptonyssus Augustson. Ichoronyssus (group II) is placed in a new genus. Ichoronyssus (group III) is included in Macronyssus Kolenati. New genus No. 1 is described, with its new species No. 1 described and designated as type-species of the genus. Descriptions of the six other new species off bats, referred to here by code numbers, are included in a fuller revisionary work (Radovsky, 1966b) to be pub- lished soon. Abstract Forty-one species of Dermanyssidae were collected from Panamanian vertebrates, mostly mammals. Of these, at least 11 are new species; the remainder are new records for Panama. Described here are: Ornithonyssus coendou n. sp. from Coendou roths- childi; Draconyssus belgicae n. gen., n. sp. from Ameiva bifrontata; Pellonyssus marui n. sp. from Cassidix mexicanus; Pellonyssus gorgasi n. sp. from Phaethornis guy. The remaining seven new species, from bats, are not described here. Acanthonyssus n. gen. is erected for Neoichoronyssus dentipes Strandtmann and Eads. Keys to the genera and species and a host list are provided. 100 ECTOPARASITES OF PANAMA HOST-PARASITE LIST Class REPTILIA Order SQUAMATA Family Teiidae Ameiva bifrontata Draconyssus belgicae n. sp. Ameiva sp. Draconyssus belgicae n. sp. Class AVES Order GALLIFORMES Family Phasianidae "common hen" Ornithonyssus bursa (Berlese) Order COLUMBIFORMES Family Columbidae Columbigallina talpacoti Pellonyssus marui n. sp. Order APODIFORMES Family Trochilidae Phaethornis guy Pellonyssus gorgasi n. sp. Order PICIFORMES Family Ramphastidae Aulacorhynchus prasinus Ornithonyssus sylviarum (Canestrini and Fanzago) Order PASSERIFORMES Family Hirundinidae Progne chalybea Pellonyssus marui n. sp. Family Turdidae Turdus grayi Pellonyssus marui n. sp. Family Vireonidae Vireo flavoviridis Pellonyssus marui n. sp. Family Icteridae Cassidix mexicanus Pellonyssus marui n. sp. Class MAMMALIA Order MARSUPIALIA Family Didelphidae Marmosa robinsoni Ornithonyssus wernecki (Fonseca) Philander opossum Ornithonyssus wernecki (Fonseca) Metachirus nudicaudatus Ornithonyssus wernecki (Fonseca) Didelphis marsupialis Ornithonyssus wernecki (Fonseca) Order CHIROPTERA Family Phyllostomidae Glossophaga soricina New genus n. sp. no. 1 Carollia castanea Radfordiella n. sp. no. 2 Carollia perspicillata Radfordiella n. sp. no. 2 New genus n. sp. no. 3 Sturnira ludovici New genus n. sp. no. 2 Vampyrops vittatus Ichoronyssus (group II) n. sp. Artibeus jamaicensis Ichoronyssus kochi Fonseca Artibeus toltecus Ichoronyssus kochi Fonseca mixed collection of Carollia perspicillata and Lonchorhina aurita Radfordiella n. sp. no. 2 phyllostomid bat New genus n. sp. no. 1 YUNKER AND RADOVSKY : DERMANYSSID MITES 101 Family Desmodidae Desmodus rotundus Radfordiella n. sp. no. 1 Family Vespertilionidae Myotis chiloensis Ichoronyssus crosbyi (Ewing and Stover) (group III) n. sp. no. 1 Myotis nigricans Ichoronyssus robustipes (Ewing) (group III) n. sp. no. 1 Mixed lots of Myotis nigricans and Myotis chiloensis Ichoronyssus crosbyi (Ewing and Stover) Family Molossidae Tadarida brasiliensis Ichoronyssus robustipes (Ewing) Molossus coibensis Ichoronyssus venezolanus (Vitzthum) molossid bat Ichoronyssus haematophagus (Fonseca) Family unknown bat Steatonyssus occidentalis (Ewing) Ichoronyssus venezolanus (Vitzthum) Order RODENTIA Family Sciuridae Sciurus granatensis Hirstionyssus keenani n. sp. Sciurus variegatoides Hirstionyssus keenani n. sp. Family Heteromyidae Liomys adspersus Hirstionyssus microchelae n. sp. Ornithonyssus bacoti (Hirst) Heteromys desmarestianus Hirstionyssus heteromydis n. sp. panamensis n. sp. minutus n. sp. microchelae n. sp. lunatus n. sp. Family Cricetidae Peromyscus nudipes Hirstionyssus galindoi n. sp. Zygodontomys microtinus Ornithonyssus bacoti (Hirst) Scotinomys xerampelinus Hirstionyssus galindoi n. sp. Sigmodon hispidus Acanthonyssus dentipes ( Strandtmann and Eads) Ornithonyssus bacoti (Hirst) Family Muridae Rattus rattus Ornithonyssus bacoti (Hirst) Family Erethizontidae Coendou rothschildi Ornithonyssus coendou n. sp. Family Echimyidae Proechimys semispinosus Acanthonyssus dentipes (Strandtmann and Eads) Ornithonyssus bacoti (Hirst) Hoplomys gymnurus Ornithonyssus sp. References BERLESE, A. 1888. Acari austro-americani collegit Alloysius Balzan et illustravit Antonio Berlese. Boll. Soc. Ent. Italiana, 20: 171-222, 13 pis. CANESTRINI, G., AND FANZAGO, F. 1878. Intorno agli Acari Italiani. Atti R. Istit. Veneto Sci. Lett, ed Arti, (5), 4: 69-208. CLARK, G. M., AND YUNKER, C. E. 1956. A new genus and species of Dermanyssidae (Acarina: Mesostigmata) from the English Sparrow, with observations on its life cycle. Proc. Helminth. Soc. Wash., 23: (2), pp. 93-101, 3 pis. EWING, H. E. 1922. The dermanyssid mites of North America. Proc. U. S. Nat. Mus. 62: 1-26, pis. 1, 2. 1925. New mites of the family Dermanyssidae. Ent. News, 36: 18-22. 1933. New genera and species of parasitic mites of the superfamily Parasitoidea. Proc. U. S. Nat. Mus., 82; art. 30, pp. 1-14, 4 pis. EWING, H. E. AND STOVER, A. J. 1915. New parasitic mites (Acarina). Ent. News, 26: 109-114, 1 pi., 1 text fig. FONSECA, F. DA 1935a. Notas de Acareologia. XIII. Novas especies sul-americanas de parasites de genera Liponyssus Kolenati, 1858 (Acarina: Liponissidae). Mem. Inst. Butantan, 9: 69-98, 17 text figs. (English Translation, pp. 99-114). 1935b. Notas de Acareologia. XXII. Liponyssus haematophagus, sp. n. (Acari: Liponissidae). Ibidem, 10: 43-46, 2 text figs. 1948. A monograph of the genera and species of Macronyssidae Oudemans, 1936 (synom. : Liponissidae Vitzthum, 1931) (Acari). Proc. Roy. Zool. Soc. London, 118:249-334, 52 text figs. FURMAN, D. P., and RADOVSKY, F. J. 1963. A new species of Ornithonyssus from the white-tailed antelope squirrel, with a rediagnosis of the genus Ornithonyssus (Acarina: Dermanyssidae). Pan- Pacific Ent. 39: 89-98, 7 text figs. HIRST, S. 1913. On three new species of gamasid mites found on rats. Bull. Ent. Res., 4: 119-124, 4 text figs. KOCH, C. L. 1839. Deutschlands Crustaceen, Myriapoden und Archniden. Ein Beitrag zur deutschen Fauna. Part 24. 102 YUNKER AND RADOVSKY : DERMANYSSID MITES 103 KOLENATI, F. 1858. Synopsis prodroma der auf Chiroptern als Epizoen vorkommenden Lausmil- ben, Carida Kolenati. Wien. Ent. Monatschr., 2: 4—7. LUCAS, P. H. 1840. Histoire Naturelle des Crustaces, des Arachnides, et des Myriapodes. Pt. 1. 600 pp., 46 pis. RADOVSKY, F. J. 1966a. Revision of the macronyssid and laelapid mites of bats: Outline of classifica- tion with descriptions of new genera and new type species. Jour. Med. Ent., 3, (1) . [In press]. 1966b. The Macronyssidae and Laelapidae (Acarina: Mesostigmata) parasitic on bats. Univ. Calif. Publ. Ent. [In press]. SAMBON, W. L. 1928. The parasitic Acarians of animals and the part they play in the causation of the Eruptive Fevers and other diseases of man. Preliminary considerations based upon an ecological study of Typhus Fever. Ann. Trop. Med. Parasit., 22: 67-132, 19 text figs. STRANDTMANN, R. W., AND EADS, R. B. 1947. A new species of mite, Ichoronyssus dentipes (Acarina: Liponyssinae), from the cotton rat. Jour. Parasit., 33: 51-56, figs. 1-3. STRANDTMANN, R. W., AND WHARTON, G. W. 1958. A manual of mesostigmatid mites parasitic on vertebrates. Contr. No. 4, Institute of Acarology, Univ. Md., College Park, xi + 330 pp., 96 figs, on 69 pp. STRANDTMANN, R. W., AND YUNKER, C. E. 1964. The genus Hirstionyssus (Acarina: Dermanyssidae) in Panama. [This volume, pp. 105-124]. TILL, W. M. 1964. A revision of the genus Pellonyssus Clark and Yunker (Acari: Mesostig- mata). Jour. Linn. Soc. Lond., Zool., 45, (304), pp. 85-102. 39 text figs. VITZTHUM, H. G. 1931. Neue parasitische Fledermausmilben aus Venezuela. Zeitschr. Parasitenk., 4, (1), pp. 1-47, 28 text figs. YUNKER, C. E. 1961. A sampling technique for intranasal chiggers. (Trombiculidae) . Jour. Parasit. 47:720. ZUMPT, F., AND PATTERSON, P. M. 1952. Three new species of parasitic mites from the Ethiopian Region. Jour. Ent. Soc. S. Africa, 15, (2), pp. 159-164, figs. 1-3. The Genus Hirstionyssus Fonseca in Panama (Acarina: Dermanyssidae) RUSSELL W. STRANDTMANN 1 AND CONRAD E. YuNKER2 Seventeen species of the relatively large genus Hirstionyssus are known from the New World. With the exception of H. butantanensis (Fonseca, 1932) from Brazil, all of these records are North American. A recent col- lection from Panamanian rodents contained seven new species, described below. Six (H. heteromydis, panamensis, minutus, galindoi, lunatus, and microchelae) are from heteromyid and cricetid rodents, and one (H. keenani) is from squirrels. Three (H. heteromydis, panamensis, and mi- nutus) form a closely related group. A key to the females of Hirstionyssus of Panama is included. The authors are grateful to Lieutenant Colonel Vernon J. Tipton, Medi- cal Service Corps, United States Army, formerly Chief, Environmental Health Branch, United States Army Caribbean, and Dr. Nathan B. Gale, Division of Veterinary Medicine, Panama Canal Company, for aid in collect- ing the hosts, and to Dr. Charles 0. Handley, Division of Mammals, United States National Museum, for identifying them. Genus Hirstionyssus Fonseca Hirstionyssus Fonseca, 1948, Proc. Zool. Soc. Lond., 118: 226. Type-species: Dermanyssus arcuatus C. L. Koch, 1839. Hirstionyssus heteromydis, new species. Figures 4, 5. DIAGNOSIS: The female is 700 ^ long and one-half as wide. Its sternal plate is rectangular and not quite twice as wide as long ; its epigynial plate narrowly linguiform, and its anal plate elliptical. The movable chela is about 1 Department of Biology, Texas Technological College, Lubbock. 2 U.S. Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Middle America Research Unit, Balboa Heights, Canal Zone, and Rocky Mountain Laboratory, Hamilton, Montana. 105 106 ECTOPARASITES OF PANAMA one-half the length of the second cheliceral segment. The coxal spur formula is 0-2-2-0. Tarsus II is without apical claw-like spurs. DESCRIPTION, HOLOTYPE FEMALE : Idiosoma. — 767 fi long by 378 /* wide. Venter (fig. 4D). — With 20-25 pairs of subequal and rather delicate opisthosomal setae. Sternal plate (fig. 4F) rectangular, sides slightly concave, posterior margin straight, anterior angles projected between coxae I and II, posterior angles broadly truncate; with three pairs of setae and two pairs of circular pores; first sternal setae just off the plate; entire plate lightly stippled, with very faint reticulations. Presternal area sclerotized and reticulated. Tritosternum (fig. 4F) without hyaline margins; basal portion lightly wrinkled ; with two laciniae, weakly plumose at tips. Metasternal seta and pore present but metasternal plate absent. A narrow endopodal apodeme present be- tween coxae III and IV. Epigynial plate slender, linguiform, obtusely pointed posteriorly, surface punctate with two median, longitudinal, slightly divergent lines; with a single pair of setae; membranous anterior portion partly overlapping sternal plate. A pore each side on soft integument near genital setae. Anal plate about twice as long as wide, elliptical; with a pair of small, circular marginal pores (generally more marginal than shown) ; paired adanal setae at the anterior margin of the anus; anal setae slightly smaller than the ventral setae. All coxae with pilif orm setae ; small fimbria present on anterior peripheral margins of coxae. Coxa I with two setae. (Both coxae I of the holotype have a noticeable longitudinal furrow adjacent to the proximal seta as in fig. 4D. This is seen as a shallow depression in some paratypes and is not evident at all in others.) Coxa II with two setae; anterior marginal spur prominent; posterior margin sharply angulate; the single ventral spur low, broad and dolabrate. Coxa III with two setae, a blunt ventral spur and a slender, sharp, posterior marginal spur. Coxa IV with a single seta; without spur. Stigma ventral, appearing between femora III and IV; peritremalia narrow and curving posterior to coxa IV : Peritreme bending dorsally before coxa II and terminating at a point level with middle of coxa I ; peritremal plate narrow, widening at level of seta L2, continuing forward to anterior margin of coxa I. Dorsum (fig. 4E). — Dorsal plate with straight or slightly convex sides, tapering posteriorly to a point; with a pair of slit-like pores at anterior margin and at least eight pairs of small circular pores scattered over remainder of plate; with about 25 pairs of small, widely separated setae, those anterior about twice the size of the others. Some 27 pairs of small setae on unarmored dorsum. (Under oil-immersion magnification the posterior setae may be seen to have one or two small barbs.) Gnathosoma (figs. 4A-C). — With 16-18 rows of one or two deutosternal teeth per row; hypostomal processes drawn out into two, long indistinct lacinae; corniculi lacking or not visibly defined (as is true of all Dermanyssidae), tectum a transparent, flaccid membrane with a transverse, blunt, denticulate anterior margin; chela slender, edentate and very long, making up almost one-half of the length of the second cheliceral segment ; setae of gnathosoma slender; of the two apical setae on dorsal side of pedipalp, the inner is blunt and a bit heavier than the outer. Legs. — Setation as shown ; without unusual modifications ; femora I and II each with three setae more robust than others ; all ventral setae longer than dorsal setae and espe- cially long on tarsus where there are two ventral whip-like setae as long as the segment ; without clawlike setae or spurs at apex of tarsus II. Measurements of sample. — Ten female specimens were measured. The numbers are averages. Idiosoma, length (exclusive of gnathosoma) 700 n; width 355 /*. Dorsal shield, length 600 /*; width 300 /*. Sternal plate, length (at midline) 70 n; width (at bases of second sternal setae) 127 p. Epigynial plate, length 240 /*; width (just posterior to genital setae) 75 M. Anal plate, length (anterior border to base of postanal seta) 85 n; width 60 /*. Legs (including coxa but excluding pretarsus) : I, 410 /JL; II, 335 /*; III, 320 /i; IV, 390 M. ALLOTYPE MALE (figs. 5B-D) : Length 460 ft,; width 270 /n. Legs: I, 330 n; II, 260 M; III, 260 n; IV, 330 /*. Coxae I-III are as in female; coxa IV has a sharply pointed posterior marginal spur, and tarsus II has two apical claw-like setae ventrally. The holoventral plate is slightly more heavily sclerotized in the region of the genital pore STRANDTMANN AND YUNKER : HirstionyssUS 107 FIG. 4. Hirstionyssus heteromydis, new species, female. A, palp. B, chelicera. C, gnathosoma, ventral view. D, venter. E, dorsum. F, sternal plate and tritosternum. 108 ECTOPARASITES OF PANAMA than elsewhere. The sternal pores are circular, as in the female. The dorsal plate tapers less than in the female and covers almost all of the dorsum. DEUTONYMPH (figs. 5E-H) : Length 490 M (480-550). Legs: I, 330 /*; II, 280 M; HI, 270 ft; IV, 310 /*. The three pairs of terminal setae on the dorsal plate are progressively larger toward the end, and are faintly serrate (the terminal pair, which is twice as long as the subterminal pair, is shown greatly enlarged in fig. 5F). TYPE MATERIAL: Holotype female (U.S.N.M. no. 2817) and 10 paratype females from Heteromys desmarestianus , Pifia (Canal Zone), 20 December 1960, collected by C. E. Yunker, in the United States National Museum. Allotype male, same data but 13 December 1960, in the United States Na- tional Museum. Remainder of material, including 26 paratype females, 3 paratype males and 5 paratype nymphs, same data but 13-20 December 1960, distributed among the collections of Rocky Mountain Laboratory, Hamilton, Montana; Texas Technological College, Lubbock; Institute of Acarology, Agriculture Experiment Station, Wooster, Ohio ; Snow Entomological Mu- seum, University of Kansas, Lawrence; British Museum (Natural His- tory), London; Entomology Research Institute, Canada Department of Agriculture, Ottawa; Zoological Institute, Academy of Sciences U.S.S.R., Leningrad; Natal Museum, Pietermaritzburg ; Musee National d'Histoire Naturelle, Paris ; Institute Royal des Sciences Naturelle de Belgique, Brus- sels ; and Institute Butantan, Sao Paulo. ADDITIONAL MATERIAL EXAMINED: 19 females of H. heteromydis from Heteromys desmarestianus, from Pifia (Canal Zone), 6-21 December 1960; 10 females from same host, from Fort Gulick (Canal Zone), 1 February 1961; a single female from same host, Rio Changena (Bocas del Toro), at lower camp, approx. 22 miles WSW of Almirante, 9 September 1961; all collected by C. E. Yunker. REMARKS : Very little variation was seen in the sample. The denticulation on the distal margin of the coxae, however, was quite variable, and in addi- tion, it could not always be established clearly that coxa III had two spurs. The first sternal setae do not always appear to be off the plate, but are some- times seen to be connected to the plate by indistinct sclerotized bridges. Hirstionyssus panamensis, new species. Figure 6. DIAGNOSIS: The female is slightly less than 600 //, long and is one-half as wide. Its sternal plate is rectangular, three times wider than long, its genitoventral plate linguiform, and its anal plate oval. The movable chela is one-third the length of second cheliceral segment. The coxal spur formula is 0-2-2-1. Tarsus II is without apical claw-like spurs. DESCRIPTION, HOLOTYPE FEMALE: Idiosoma. — 537 fi long by 310 «. wide. Venter (fig. 6A). — With 18-21 pairs of short, piliform opisthosomal setae. Sternal plate rectangular, anterior margin and sides nearly straight, posterior margin concave, anterior angles acute, posterior angles rounded; nearly three times wider than long; with three pairs of setae, first pair on anterior margin of plate; with two pairs of slit- like sternal pores; anterolaterally with reticulations. Presternal area sclerotized and reticulated. Tritosternum similar to that of H. heteromydis. A. pair of metasternal setae and a circular pore present on soft integument adjacent to coxa III. Epigynial plate linguiform, not greatly constricted in middle; rounded posteriorly; surface punc- STRANDTMANN AND YUNKER I Hirstionyssus 109 A FIG. 5. Hirstionyssus heteromydis, new species, female (A), male (B-D), deutonymph (E-H). A, coxae I and II. B, venter. C, chelicera. D, tarsus II, ventral view. E, dorsum. F, terminal seta of dorsal shield. G, venter. H, chelicera. 110 ECTOPARASITES OF PANAMA tate ; membranous anterior portion slightly overlapping sternal plate. A circular pore on soft integument either side of plate. Anal plate broadly oval, about three-fourths as wide as long; paired marginal pores appearing as minute punctations; anus in anterior of plate; adanal setae arising at a level with middle of anus. Coxa I with two piliform setae; coxa II with two piliform setae, a sharp anterior marginal spur, a broad dolabrate ventral spur and a sharply angulate posterior margin; coxae III with two piliform setae, a broad sharp ventral spur, and a slender, sharp, posterior marginal spur; coxa IV with a single piliform seta and a small, ventral, posterior marginal spur. Stigma ventral, appearing between femora III and IV; peritremalia narrow and curving posterior to coxa IV. Peritreme bending dorsally in area of coxa II and terminating at a point level with middle of coxa I. Peritremal plate narrow, widening at level of seta L2, continuing forward to anterior margin of coxa I. Dorsum (fig. 6B). — Dorsal plate with straight sides, tapering posteriorly to a point, with at least 25 pairs of piliform setae, those anterior longest, those posterior slightly shorter than the 16-20 pairs of setae on adjacent soft integument. Gnathosoma (fig. 6C— E). — Similar to that of H, heteromydis except that the movable chela forms no more than one-third of the total length of the chelicera. Deutosternal teeth not seen. Legs. — Not significantly different from those of H. heteromydis except femora I and II without robust setae and tarsus II without whiplike setae (fig. 6F). Measurements of sample. — Four females were measured. The numbers are averages. Idiosoma, length (exclusive of gnathosoma) 590 /j,; width 300 /*. Dorsal shield, length 540 n; width 260 /i. Sternal plate, length (at midline) 40 /*; width (at bases of second sternal setae) 115 /*. Epigynial plate, length 200 /u; width 80 /JL. Anal plate, length (anterior border to base of postanal seta) 65 /j.; width 50 /*. Legs: I, 310 /j.; II, 240 /*; III, 225 p; IV, 290 p. TYPE MATERIAL: Holotype female (U.S.N.M. no. 2818) from Heteromys desmarestianus, Pina (Canal Zone), 20 December 1960, collected by C. E. Yunker, in the United States National Museum. Three paratype females, same data but 13 December 1960, distributed among the collections of United States National Museum ; Rocky Mountain Laboratory, Hamilton, Montana ; and Texas Technological College, Lubbock. REMARKS: H. panamensis is similar to H. heteromydis in the tectum, tritosternum, hypostomal processes, and ventral spur on coxa II. It differs in many ways from the latter. In panamensis the sternal plate is much wider and shorter, has rounded posterolateral angles, and possesses slit-like pores. In addition, its epigynial plate is broad in relation to length, and is rounded posteriorly. Its adanal setae originate at a point level with the middle of the anus. The movable chela forms less than one-third the length of the second cheliceral segment. The peritremalia is relatively wide posterior to the stigma. Tarsus II has only short setae. None of the material before us offered a distinct view of the complete peritremalia, the deutosternum or the dorsal plate. It is probable that a pair of anterior pores are present on the dorsal plate, as well as more small pores and setae than we show. Some paratypes showed small barbs on the pos- terior dorsal setae. Hirstionyssus minutus, new species. Figure 7. DIAGNOSIS: This is a small species. The female is 400 ^ long and the male 280 ^. The female sternal plate is rectangular, about twice wider than long. Its epigynial plate is linguiform and perceptibly broader in the post- STRANDTMANN AND YUNKER : HirstiomjSSUS 111 FIG. 6. Hirstionyssus panamensis, new species, female. A, venter. B, dorsum. C, gnathosoma, dorsal view. D, chelicera. E, gnathosoma, ventral view, and tritosternum. F, tarsus II, ventral view. 112 ECTOPARASITES OF PANAMA coxal area than in the intercoxal area. The movable chela is more than one- half the length of the second cheliceral segment. The coxal spur formula is 0-2-1-1. Tarsus II is without apical claw-like setae. DESCRIPTION, HOLOTYPE FEMALE : Idiosoma. — 425 n long by 242 ft. wide. Venter (fig. 7A). — With 12-17 pairs of short, piliform opisthosomal setae, that be- come progressively shorter posteriorly. Sternal shield rectangular, anterior margin straight, lateral and posterior margins concave; with three pairs of setae and two pairs of pores; first sternal setae on anterior margin of plate; surface of plate with indistinct longitudinal wrinkles, anterolateral corners and presternal area reticulate. Tritosternum as in H. heteromydis. Metasternal seta and pore present but metasternal plate absent. A narrow endopodal apodeme between coxae III and IV. Epigynial plate linguiform, relatively broad in postcoxal area; bluntly rounded caudally; anteriorly overlapping part of sternal plate; with a pair of setae; surface densely covered with minute puncta- tions. Anal plate a rounded oval, nearly as wide as long; with a pair of minute marginal pores ; paired adanal setae at anterior margin of anus. Coxa I with two piliform setae ; coxa II with two piliform setae, a sharp anterior marginal spur, a broad, dolabrate, ven- tral spur and a sharply angulate posterior margin; coxa III with two robust setae and a sharp ventral spur, coxa IV with a single piliform seta and a small, sharp posterior marginal spur. Stigma ventral, appearing between femora III and IV; peritremalia narrow and curving posterior to coxa IV; peritreme bending dorsally in region of coxa II, terminating at a point level with middle of coxa I; peritremal plate narrow. Dorsum (fig. 7B). — Dorsal plate oval, broadly rounded caudally, covering most of dorsum; with 27-30 pairs of small setae and many small circular pores; anteriormost pair and posteriormost pair of setae 6 or 7 /j. long, remainder extremely minute (less than 3 M). About eight pairs of minute setae on unsclerotized dorsum. Gnathosoma (figs. 7C-F). — Similar to that of H. heteromydis but movable chela slightly more than one-half length of second cheliceral segment, and a mediodorsal, apical, tarsal seta is markedly thick and blunt. Legs. — Similar to those of H. heteromydis: ventral setae generally longer and more robust than dorsal setae, except on femora I and II where reverse is true; tarsus II with some moderately long ventral setae. Measurements of sample. — Four females were measured. The numbers are averages. Length (exclusive of gnathosoma) 400 /*; width 230 /j.. Dorsal shield, length, 382 /*; width 205 p. Sternal plats, length (at mid-line) 45 /*.; width (at basees of second sternal setae) 95 /*. Epigynial plate, length 177 M; width (at widest point) 75 /*. Anal plate, length (to base of postanal seta) 39 M; width 46 /j.. Legs: I, 260 /*; II, 195 /*; III, 170 M; IV, 225 ^ ALLOTYPE MALE (figs. 7G-J) : Length, 296 /*; width, 180 /j.. Legs: I, 236 ,u; II, 180 /u; III, 130 /a; IV, 210 fj.; coxae as in female; tarsus II with two blunt claw-like setae ventrally. Holoventral plate expanded posterior to coxae; fused with anal plate; with nine pairs of setae plus the single postanal seta; with five pairs of pores; the first sternal pores slit- like, the rest circular. Dorsal plate similar to that of female. TYPE MATERIAL: Holotype female (U.S.N.M. no. 2819), paratype female and allotype male from Heteromys desmarestianus, Pifia (Canal Zone), 20 December 1960, collected by C. E. Yunker, in the United States National Museum. A paratype female, same data, in collection of Rocky Mountain Laboratory, Hamilton, Montana, and another, same data, in collection of Institute of Acarology, Agriculture Experiment Station, Wooster, Ohio. A paratype male and female, same data, but 7 December 1960, in collection of Texas Technological College, Lubbock. REMARKS: H. minutus resembles H. heteromydis in the tectum, trito- sternum, hypostomal processes, chelicerae and leg setation. It differs from the latter in size, by having only one spur on coxa III, by having less than STRANDTMANN AND YUNKER : HlTStlOnySSUS 113 FIG. 7. Hirstionyssus minutus, new species, female (A-F), male (G-J). A, venter. B, dorsum. C, palpal tarsus, dorsal view. D, gnathosoma, ventral view, and tritosternum. E, tectum and palp, dorsal view. F, chelicera. G, venter (one adanal seta omitted). H, gnathosoma, ventral view. I, tectum and palp, dorsal view. J, chelicera. 114 ECTOPARASITES OF PANAMA 17 pairs of opisthosomal setae, and by having many minute dorsal shield setae. In addition, shapes of the body plates are typical. The dorsal shield, epigynial plate and posterolateral angles of H. minutus are broadly rounded posteriorly ; the epigynial plate is relatively expanded posterior to coxae IV and the anal plate is almost circular. It differs from H. panamensis in chela/chelicera length-ratio, by adanal setae that arise at the anterior margin of the anus ; by a longer sternal plate, by having only one spur on coxa III, by the minute dorsal setae, and by the robust, blunt mediodorsal seta of the palpal tarsus. The dorsal setae are so small that it is difficult to distinguish between a setal base and a pore. Hirstionyssus microchelae, new species. Figure 8A-D. DIAGNOSIS : The female is 620 //, long and 440 ^ wide at its greatest width. Its sternal plate is rectangular and three times wider than long. Its epigynial plate is linguiform and truncate, and its anal plate circular. Its chelicerae are slender and the movable chelae are small, each less than one-seventh the length of the second cheliceral segment. The coxal spur formula is 0-2 (3?) -2-1. Tarsus II is without apical claw-like setae. The posterior dorsal setae are serrate. DESCRIPTION, HOLOTYPE FEMALE: Idiosoma. — 590 n long by 384 fj. wide. Venter (fig. 8A) — With about 22-25 pairs of opisthosomal setae. Sternal plate rectangular, with straight anterior and lateral margins and a slightly convex posterior margin; nearly three times wider than long; with three pairs of setae and two pairs of slit-like pores ; first sternal setae on anterior margin of plate ; anterolateral corners and presternal area reticulate. Tritosternum as in H. heteromydis. Metasternal seta and pore present, but metasternal plate absent. A narrow endopodal apodeme between coxae III and IV. Epigynial plate linguiform, not greatly widened in postcoxal area; bluntly rounded caudally; anteriorly overlapping part of sternal plate; with a pair of setae; surface densely punctate. Anal plate circular; with a pair of minute marginal pores; paired adanal setae at a level with posterior of anus. Coxa I with two piliform setae, its peripheral margins fimbriate; coxa II with two piliform setae, a large, sharp anterior marginal spur, a small blunt ventral spur, and a sharp, angulate posterior dorsal margin that may be spur-like; coxa III with two piliform setae, a blunt ventral spur, and a sharp posterior dorsal spur; coxa IV with one piliform seta and a small, sharp posterior marginal spur. Stigma ventral, appearing between femora III and IV; peritremalia narrow and curving posterior to coxa IV; peritreme bending dorsally in region of coxa II, terminating at a point level with middle of coxa I; peritremal plate visible on either side of peritreme, widening abruptly in anterior third, terminating adjacent to paired, slit-like, dorsal shield pores. Dorsum (fig. 8B). — Dorsal shield elliptical, sides slightly convex, tapering caudally to a blunt point; with 26 pairs of short setae, those posterior serrate; with 19 or 20 pairs of small circular pores and a pair of large, slit-like, anterior pores. With 40-50 pairs of setae on adjacent soft integument. Gnathosoma (fig. 8C). — Similar to that of H. panamensis, except that the chelicerae are relatively narrow and elongate. The movable chela is less than one-sixth the length of the second cheliceral segment (fig. 8D). Legs. — Not significantly different from those of H. panamensis. Measurements of sample. — Three females were measured. The numbers are averages. Idiosoma, length 572 n; width 378 p.. Dorsal shield, length 500 /JL; width 290 M- Sternal plate, length (at mid-line) 41 ft; width (at bases of second sternal setae) 122 p. Epigynial STRANDTMANN AND YUNKER : Hirstionyssus 115 FIG. 8. Hirstionyssus microchelae, new species, female. A, venter. B, dorsum and peritremalia. C, gnathosoma and tritosternum, oblique view. D, chelicerae. Hirs- tionyssus keenani, new species, female. E, chelae. F, tarsus II, ventral view. 116 ECTOPARASITES OF PANAMA plate, length 220 n; width 100 /u. Anal plate, length (to base of postanal seta) 70 /*; width 80 fj.. TYPE MATERIAL: Holotype female (U.S.N.M. no. 2820) and 4 paratype females from Heteromys desmarestianics, Pina (Canal Zone), 13 December 1960, collected by C. E. Yunker, in the United States National Museum. Six paratype females, same data, distributed among the collections of Rocky Mountain Laboratory, Hamilton, Montana; Texas Technological College, Lubbock ; Institute of Acarology, Agriculture Experiment Station, Wooster, Ohio ; and British Museum (Natural History) , London. ADDITIONAL MATERIAL EXAMINED: Two females from type host and lo- cality, but 29 December 1961 ; 1 female from Liomys adspersus, Guanico, Las Palmitas (Los Santos), 10 February 1962, collected by C. 0. Handley and F. Green well; 2 females from type host, Almirante (Bocas del Toro), 7 September 1960, collected by P. Galindo; 7 females from type host, Rio Changena (Bocas del Toro), at lower camp, 22 miles WSW of Almirante, elevation about 2800 feet, 17 September 1961, collected by C. E. Yunker. REMARKS : H. microchelae is easily distinguished from other Panamanian species by the chela/chelicera length-ratio and the circular anal plate. In addition, crushed specimens reveal a pair of circular pores on the medial aspect of the peritremal plate anterior to the stigma. These are closely as- sociated with a pair of square, cell-like depressions or muscle-scars (fig. 8B). Variation was apparent in the shape of the posterior margin of coxa II. In some specimens this margin was angulate and distinctly spur-like ; in others, including the holotype, no such modification could be seen. It is pos- sible that this difference is an artifact of mounting. The type specimens were taken from a host that was simultaneously in- fested with heteromydis, panamensis and minutus. All females of micro- chelae appeared to be engorged on blood, whereas none of the other species did. Hirstionyssus keenani, new species. Figures 8E-F, 9. DIAGNOSIS: This is a typical Hirstionyssus species, characterized by a female sternal plate that is deeply concave at its posterior margin and acute, elongate coxal spurs. The female is SOOju long ± 30 ^, and approximately one- half as wide. The coxal spur formula is 0-2-2-1, tarsus II lacks claw-like setae, and the dorsal shield setae are noticeably shorter than the ventral setae. DESCRIPTION, HOLOTYPE FEMALE : Idiosoma. — 513 fj. long by 325 IJL wide. Venter (fig. 9A). — With 20-24 pairs of setae. Sternal plate deeply emarginate posteriorly, seven times wider than long; with three pairs of approximately equal setae, and two pairs of circular pores; first sternal setae on anterior margin of plate; an- terolateral corners and presternal areas reticulate. Tritosternal base punctate; laciniae weakly ciliate. Metasternal setae and pore present, but metasternal plate absent. A narrow endopodal apodeme present between coxae III and IV. Epigynial plate lingui- form, not greatly widened posterior to coxae, rounded caudally; surface densely covered with minute punctae; with a single pair of setae. A circular pore on each side near genital setae. Anal plate oval; surface punctate; anus in anterior third of plate; paired adanal setae arising at a level with middle of anus. Coxa I with two piliform setae, the distal one longer and heavier than the proximal one; coxa II with two piliform setae, a sharp anterior marginal spur and an acute, elongate ventral spur; coxa III with two pili- STRANDTMANN AND YUNKER : Hirstionyssus 117 FIG. 9. Hirstionyssus keenani, new species, female (A-E), deutonymph (?) (F, G). A, venter. B, coxa IV. C, tritosternum. D, dorsum. E, coxae II and III. F, coxae and holoventral plate. G, dorsal shield and peritremalia. 118 ECTOPARASITES OF PANAMA form setae and two acute, elongate spurs ; coxa IV with a single pilif orm seta and a small, sharp marginal spur. Stigma ventral almost marginal, appearing between femora III and IV; peritreme bending dorsally in region of coxa II, continuing anteriorly as far as anterior margin of coxa I; peritremal plate narrow. Dorsum (fig. 9D). — Dorsal shield elliptical with straight, parallel sides; tapering caudally in a blunt point; with 26 pairs of pilif orm setae shorter than those on adjacent soft integument; with 17 pairs of pores, the anteriormost pair slit-like, the remainder circular. With 20-22 pairs of setae on adjacent soft integument, slightly shorter than ventral setae. Gnathosoma. — With 12-14 deutosternal teeth arranged in an irregular file. Hy- postomal processes drawn out into two long lacinae. Tectum a transparent, long mem- brane with a truncate, fimbriate anterior margin. Movable chela elongate, about one- third the length of the second cheliceral segment; with a transparent, triangular medial tooth, and a ciliated basal lobe. Fixed chela with a hyaline, stellate, seta-like structure arising opposite base of movable chela, adjacent to a small circular pore (fig. 8E). Legs. — Setation typical of Hirstionyssus spp. Femur I with two and femur II with one robust dorsal setae. Tarsus II with some long, whip-like ventral setae (fig. 8F). Tarsus IV with a terminal spur-like seta. Measurements of sample. — Five females were measured. The numbers are averages. Idiosoma, length (exclusive of gnathosoma) 500 /*; greatest width 320 /JL. Dorsal shield, length 450 n; greatest width 252 fj.. Sternal plate, length (at mid-line) 15 ^5 width (at bases of second sternal setae) 126 /*. Epigynial plate, length 257 /*; width (just posterior to genital setae) 93 /JL. Anal plate, length (anterior border to base of postanal setae) 63 /*; width 63 M. Legs: I, 333 M; II, 260 p; III, 245 p\ IV, 315 M- TYPE MATERIAL : Holotype female (U.S.N.M. no. 2821) and two paratype females from Sciurus variegatoides, Gamboa (Canal Zone), 4 December 1960, collected by C. E. Yunker, in the United States National Museum. Three paratype females, same data, distributed among the collections of Rocky Mountain Laboratory, Hamilton, Montana, and Texas Technological College, Lubbock. ADDITIONAL MATERIAL EXAMINED : Two females and one deutonymph from Sciurus granatensis chiriquensis, Martinz's dairy, Cerro Punta (Chiriqui), elevation about 6800 feet, 2 May 1961, collected by C. E. Yunker. One female, same type host and locality, but 12 March 1962, and one deutonymph, data same as holotype (figs. 9F, G). Coxae III and IV of the nymph lack the marginal spur seen in the female. The dorsal plate setae are similar to those of the female, except that the terminal pair is long (a characteristic of im- mature specimens of Hirstionyssus). REMARKS : The combination of arcuate sternal plate, coxal spur formula 0-2-2-1, and lack of claw-like setae at the ventral apex of tarsus II is shared by only one other species, H. neotomae Eads and Hightower, 1951. The latter, however, has a sternal plate less deeply concave (length-width ratio is 1 :4.3, as compared with 1 :7.4 for H. keenani) ; its coxal spurs are much smaller, and the anterior dorsal setae are longer. In H. neotomae, the first three rows of dorsal plate setae overlap, whereas in H. keenani none of the setae on the dorsal plate are long enough to reach the bases of those in the succeeding row. H. keenani also resembles H. isabellinus (Oudemans, 1913) , but in this latter species the sternal plate is even less arcuate than in H. neotomae and coxa IV lacks a spur. H. keenani is named for Charles M. Keenan, Chief, Vector Control Sec- STRANDTMANN AND YUNKER : HlTStionySSUS 119 tion, Environmental Health Branch, United States Army Caribbean, and Canal Zone naturalist. Hirstionyssus galindoi, new species. Figure 10. DIAGNOSIS : The female sternal plate is about four times wider than long. The chelae are one-half the length of the second cheliceral segment. The FIG. 10. Hirstionyssus galindoi, new species, female. A, venter. B, dorsum. C, chelicera. coxal spur formula is 0-1-2-1 or 0-2-2-1, coxa II sometimes having a small, rounded ventral knob that might be taken for a spur. Tarsus II is without claw-like setae. DESCRIPTION, HOLOTYPE FEMALE : Idiosoma. — 462 fj. long by 326 n wide. Venter (fig. 10A). — Sternal plate short, about four times wider than long, deeply and broadly concave posteriorly, anterior margin nearly straight; lightly reticulated at sides; the three pairs of subequal sternal setae shorter than plate. Presternal area 120 ECTOPARASITES OF PANAMA lightly reticulated. Tritosternum without basal hyaline margins; laciniae ciliate and extending nearly to the apical hypostomal setae. Metasternal setae subequal to sternals. Without metasternal plates. Epigynial plate linguiform; genital and all ventral setae subequal, longer than dorsals; one pair of ventrals on the posterior margin of the plate or apparently so. Posterior ventral setae as well as smaller marginal and dorsal setae weakly serrate on one side (fig. 10A). Anal plate broadly ovate; the three anal setae slender, subequal, and shorter than anal slit. Adanal setae inserted opposite the middle of the anal slit. With 15 or 16 pairs of ventral non-plate setae. Metapodal plates absent. Peritreme ventrolateral, becoming dorsal over coxa II, and extending to level of middle of coxa I ; surrounded by a narrow peritremal plate, which encircles coxa IV posteriorly. Dorsum (fig. 10B). — Dorsal shield with sides subparallel, tapering posteriorly to a blunt wedge; with 26 pairs of short setae which are a bit longer anteriorly and peripherally; lightly reticulated in scapular area. With 10-12 pairs of dorsal, non-plate setae. Gnathosoma. — Setation weak; deutosternal teeth in a double file, with about 14-17 denticles. Malae internae long, slender; tectum truncate, with a deeply ciliated margin. Chelicerae relatively short and heavy, the chelae (fig. IOC) forming one-half the length of the second cheliceral segment. Palpal genu with a transverse dorsal pore near base (fig. 10B). Legs. — Setae of legs slender and piliform; femora I and II each with two slightly enlarged dorsal setae; femora III and IV each with one slightly enlarged dorsal seta. Coxa I with two subequal piliform setae ; coxa II with an anteromarginal spur and a small, rounded ventral boss; coxa III with a ventral and a posteromarginal spur, both small and acute; coxa IV with one small, sharp marginal spur. Tarsus II without claw- like subapical setae. DEUTONYMPH : Length of idiosoma, 305 /JL. Sternal shield extending to level of posterior margin of coxa IV, as is usual for deutonymphs of this family. The first four pairs of setae are marginal, the fifth pair is off the margin near the posterior end. Coxa II with a slight ventral elevation; coxa III with an acute ventral spur; coxa IV without a spur but with small denticles on the posteroapical margin. Dorsal plate entire, bearing two long, weakly barbed setae at posterior tip. Peritreme extending to level of posterior margin of coxa I ; poststigmal plate lacking. MALE: Unknown. TYPE MATERIAL: Holotype female (U.S.N.M. no. 66415) and one para- type female from Scotinomys xerampelinus, Cerro Punta (Chiriqui), ele- vation about 7000 feet, 14 March 1962, in the United States National Mus- eum; three paratype females and 3 paratype nymphs from Peromyscus nudipes, same data, but 9 to 14 March 1962; all collected by C. M. Keenan. Paratypes distributed among collections of United States National Museum ; Texas Technological College, Lubbock; and Rocky Mountain Laboratory, Hamilton, Montana. REMARKS : This species is closest to H . breviseta Strandtmann and Mor- lan, 1953. The latter, however, is without ventral spurs or knobs on coxa II and its first sternal setae are extremely close set ; in addition, the spurs of coxa III are not as pronounced as in the present species. H. galindoi also re- sembles H. transiliensis Bregetova, 1956, but in the latter species there are no ventral non-plate setae so close to the epigynial plate as to appear to be touching it. The ventral idiosomal setae of H. galindoi are long enough to reach the bases of succeeding setae, while those of H. transiliensis are quite short. H. galindoi is named for Sr. Pedro Galindo V., Gorgas Memorial Labora- tory, Panama, who kindly provided certain specimens examined in this study. STRANDTMANN AND YUNKER : HirstlOmjSSUS 121 FIG. 11. Hirstionyssus lunatus, new species, female (A-C), male (D-F). A, venter. B, chelicera. C, dorsum. D, chelicera. E, dorsum. F, venter. 122 ECTOPARASITES OF PANAMA Hirstionyssus lunatus, new species. Figure 11. DIAGNOSIS : This is a small species ; the female is 350 //. long and has short, delicate setae. It is instantly recognizable by the unusual shape of the anal plate and especially the wide cribrum, which subtends the plate as a semi- circular crescent. The epigynial plate is uniquely scaly in appearance. The coxal spur formula is 0-3-3-1, and trochanters III and IV each bear two heavy apical spurs. Tarsus II is without claw-like apical setae. DESCRIPTION, HOLOTYPE FEMALE (figs. 11A-C). Idiosoma 340 fj, long by 238 fj. wide. Venter (fig. 11A). Sternal plate slightly arched, smooth or only faintly lined; the corners rounded, not noticeably projecting between the coxae. First sternal setae mesad of first sternal pores ; second sternal pores well mesad of sternal setae 2 and 3. Sternal pores small and circular. Sternal setae 2 and 3 close set. Sternal setae 1 slightly shorter than 2 and 3. Presternal area not heavily sclerotized, very faintly lined. Tritosternum weak, transparent; basal portion transversely wrinkled; laciniae finely ciliated. Epigynial plate broad and anteriorly overlapping part of sternal plate; pos- terior margin broad, slightly convex, with one pair of epigynial setae ; in addition a single pair of setae arise from the posterolateral margins of the plate; entire plate reticulate and scaly in appearance. Anal plate unusual for genus. Posterior margin broadly flared; cribrum subtending posterior margin as a crescentic band. Anal opening near anterior margin of plate. Adanal setae arising at anterior level of anal slit, and sub- equal to it in length. Postanal seta well removed from anal slit ; near cribrum ; subequal to adanals. From seven to 11 pairs of ventral non-plate setae which become shorter and heavier laterally. Dorsum (fig. 11C). — Dorsal shield covering most of dorsum, straight sided, and broadly rounded anteriorly and posteriorly. With 26 pairs of minute setae that are a bit larger posteriorly. At least posteriormost of these slightly serrate on one side (fig. 11C). Stigma dorsolateral ; peritreme mostly dorsal; enclosed in a narrow plate that attaches to dorsal shield over coxa II. Peritreme extending nearly to level of middle of coxa I. Legs. — Majority of setae short and slender. Both setae of coxa I piliform, the proximal somewhat longer. Coxa II with an anteromarginal spur, a ventral triangular spur, and the posterior margin produced into a large, acute projection, here counted as a spur; with two piliform setae. Coxa III with a ventral spur, a posteromarginal spur and two setae ; the anteromarginal seta slender and spinif orm, the posteromarginal setae piliform. Coxa IV with a marginal tooth and a single piliform seta. Anterior apical margin of trochanters III and IV each with three sharp spurs or teeth. Femora I and II with each a pair of enlarged dorsal setae. Tarsus II modified into a slight hook at apex but without claw-like setae. One pair of slender, flagellif orm medioventral setae on tarsus II. Inner margin of femora and genua III and IV slightly crenulated. Gnathosoma. — Setae small and slender. Hypostomal and gnathosomal setae small. Malae internae long and slender. Tectum extending as far as level of middle of palpal tibiae; with a ciliate margin. Chelicerae long and slender; chelae about one-fourth as long as second cheliceral segment (fig. 11B). MALE (figs. 11D-F) : Idiosoma. — 275 /* long by 188 fj. wide. Venter (fig. 11F). — Holoventral plate slightly expanded behind coxa IV, with eight pairs of setae, none of which quite reaches the base of the succeeding seta, plus three smaller anal setae; the latter slender and shorter than anal slit. Anal plate wide, with crescentic cribrum as in female. Dorsum (fig. HE). — Dorsal shield nearly covering dorsum; with 28 pairs of very small setae that are slightly longer posteriorly. Peritreme extending nearly to level of middle of coxa I. Stigma ventrolateral. Legs. — Tarsus II with two subapical clawlike setae, basad of these are two long flagelliform setae. Trochanters II and III lack the marginal spurs of the female. Coxa STRANDTMANN AND YUNKER : HirstiomjSSUS 123 IV with one or two anteromarginal teeth as well as the posteromarginal spur. The coxal spurs are relatively smaller than those of the female. Gnathosoma. — Not significantly different from that of the female. Chelicerae slender, chelae unmodified. TYPE MATERIAL: Holotype female (U.S.N.M. no. 66611), three paratype females and one allotype male from Heteromys desmarestianus, Rio Chan- gena (Bocas del Toro), lower camp, approximately 22 miles WSW of Al- mirante, elevation about 2800 feet, 27 September 1961, collected by C. E. Yunker. Two paratype females from type host, Pina (Canal Zone), 13 De- cember, 1960, collected by C. M. Keenan. Holotype female, allotype male and one paratype female deposited in the United States National Museum. One paratype female in the collection of Rocky Mountain Laboratory, Hamilton, Montana, and one paratype female in the collection of Texas Technological College, Lubbock. KEY TO THE PANAMANIAN SPECIES OF H1RSTIONYSSUS FEMALES 1. Epigynial plate with scalelike pattern and two pairs of setae; anal plate much wider than long, laterally angulate; trochanters III and IV with large distal marginal spurs H. lunatus n. sp. Without this combination of characters 2 2. Sternal plate approximately rectangular; ventral spur of coxa II broad and dolabrate, reduced, or absent, on heteromyid or cricetid rodents 3 Sternal plate arcuate, posterior border deeply emarginate; ventral spur of coxa II acute and elongate ; on Sciurus H. keenani n. sp. 3. Chelicerae slender, long; movable chela at most one-sixth the length of second cheliceral segment; anal plate circular H. microchelae n. sp. Chelicerae normal; movable chela at least one-third as long as second cheliceral segment 4 4. Dorsal shield setae normal or reduced but not minute; coxae III with two spurs 5 Dorsal shield setae extremely minute; coxa III with one spur; a small species about 400 /j. long ; with less than 18 pairs of ventral opisthosomal setae H. minutus n. sp. 5. Sternal plate at least three times wider than long; sternal pores slitlike; coxa IV with one spur 6 Sternal plate not quite twice wider than long; sternal pores circular; coxa IV without spurs H. heteromydis n. sp. 6. Sternal plate three times wider than long; movable chela one-third length of second cheliceral segment H . panamensis n. sp. Sternal plate four times wider than long; movable chela one-half length of second cheliceral segment H . galindoi n. sp. References BREGATOVA, N. G. 1956. Gamasoidea. Tabl. anal. Faune U.S.S.R., no. 61. 247 pp., 563 figs. (In Russian.) FONSECA, F. DA 1932. Notas de Acareologia II. Ichoronyssus butantanensis sp. n. (Acarina: Der- manyssidae). Mem. Inst. Butantan, 7: 135-138, 1 fig. 1948. A monograph of the genera and species of Macronyssidae Oudemans, 1936 (synom. : Liponyssidae Vitzthum, 1931). (Acari). Proc. Zool. Soc. Lond., 118: 249-334, 52 figs. KOCH, C. L. 1839. Deutschlands Crustaceen, Myriapoden und Arachniden. Ein Beitrag zur deutschen Fauna. Pt. 24. OUDEMANS, A. C. 1913. Acarologische Aanteekeningen. xlviii. Ent. Ber., 3: 384-387. STRANDTMANN, R. W., AND MORLAN, H. B. 1953. New species of Hirstionyssus and a key to the known species of the world. Texas Rep. Biol. Med., 11: 627-637, pis. 1-3. 124 The Spinturnicid Mites of Panama1 (Acarina: Spinturnicidae) DEANE P. FURMAN 2 In recent years, particularly extensive collections have been made of ecto- parasites from Central American mammals and birds. These have chiefly been obtained in the course of epidemiological investigations of arthropod- borne diseases that are transmissible to man, or as a prerequisite to such in- vestigations, because knowledge of the systematics and biology of ectopara- ites is basic to understanding potential pathogen-vector relationships. The present report on the Spinturnicidae, mites which are exclusively parasitic on bats, deals with the Panamanian collections and, with few ex- ceptions, those Trinidadian collections pertaining to new species that occur in both Panama and Trinidad. The rich Panamanian collections were made available to the author by Lt. Col. Vernon J. Tipton, then Chief, Environ- mental Health Branch, Preventive Medicine Division, United States Army, Caribbean. Concurrently, the author studied extensive collections of Trini- dadian bat mites lent through the courtesy of Dr. Thomas H.G. Aitken of the Trinidad Regional Virus Laboratory of the Rockefeller Foundation at Port of Spain. Trinidad collections will be dealt with in a separate paper. All collecting localities mentioned in this paper are in Panama, unless otherwise stated. The author acknowledges with appreciation the loan of collections by Lt. Col. Tipton and Dr. Aitken and staff of the Trinidad Regional Virus Labo- ratory. Gratitude is expressed to Dr. Marc Andre for making available a specimen of Periglischrus caligus identified by Kolenati, to E. P. Catts for inking the drawings and to the several graduate students in parasitology who helped with various phases of the work. To Mr. Arthur M. Greenhall 1 This investigation was supported in part by Research Grant E-1509 from the Na- tional Institute of Allergy and Infectious Diseases of the National Institutes of Health, Public Health Service. 2 Department of Entomology and Parasitology, University of California, Berkeley. 125 126 ECTOPARASITES OF PANAMA and Dr. Charles 0. Handley, grateful acknowledgment is expressed for aid in problems of bat identification, but the author accepts responsibility for accuracy of host names recorded here. An excellent revision of the family Spinturnicidae was given by Rudnick (1960) . The reader is referred to this publication for complete synonymical lists, for details characteristic of the family and for a key to the genera of Spinturnicidae. Representatives of three genera from Panama are recorded here : Peri- glischrus, Spinturnix, and Paraspinturnix. All Panamanian species key out to the proper genus in Rudnick's (1960) key, except for a new species of Periglischrus, the male of which has peritremes, vestigial to apparently absent anterior to coxae III. Genus Periglischrus Kolenati Periglischrus Kolenati, 1857, Wien. Ent. Monatschr., 1, (2), p. 60. Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 195 (complete bibliographical synonymy). Type-species: Periglischrus caligus Kolenati, 1857, by subsequent designation (Oude- mans, 1903, Tidschr. Ent., 45:135). DIAGNOSIS (based on adults and modified from Rudnick, 1960) : Dorsum. — Two dorsal plates closely approximated, occupying greater part of podosoma in female, and most of idiosoma of male; anterior plate much larger than posterior plate. Five pairs setae bordering anterior dorsal shield anterior to stigmata. One pair setae slightly posteromedial to stigmata. Peritreme completely dorsal, usually extending from level of coxa IV to level of coxa I. Venter. — Lacking tritosternum. Female sternal plate with three pairs marginal setae which may be set off plate. Male sternal plate with five pairs setae. Epigynial plate reduced to narrow sclerotization with pair of small genital setae close to or on lateral margins of plate near posterior tip. Opisthosoma of non-teneral females usually greatly expanded to relatively flat, broad, fan-shaped appearance, with characteristically shaped areas of heavy sclerotization. Opisthosoma of male reduced, scarcely projecting posterior to level of coxae IV ; six to eight pairs setae, exclusive of adanal pair, ventrally posterior to sternal plate. Anal plate in female small, subterminal, narrow, with pair adanal setae and dorsal postanal seta ; anus terminal or dorsal. Male anal plate usually large, oc- cupying much of area between coxae IV, with minute, dorsoterminal postanal seta. Legs. — Large caruncles and claws on all legs. Dorsal and lateral setae short to long. Ventral setae usually short with exception of long, subterminal, tarsal trichobothria. Tarsus I of males with two long, bluntly tipped dorsal sensory setae located respectively at basal one-third and apical positions. REMARKS : The previously known species of the genus Periglischrus were recorded only from bats of the family Phyllostomidae. The major part of the records of Periglischrus reported here are from the Phyllostomidae, but several collections are from the Desmodidae of the same superfamily and several are recorded from bats of the family Natalidae of the Vespertili- onoidea. A single collection of several specimens is also recorded from a bat of the family Noctilionidae, superfamily Emballonuroidea. Various instars of Periglischrus species commonly are encountered on their hosts. In common with other members of the family, species of this genus pass the larval stage of development in the body of the female. The first active stage seen is the protonymph which is characterized as follows : three pairs sternal setae ; metasternal setae absent ; four pairs dorsal propo- dosomal setae; peritreme short, extending barely, if at all, anterior to level FURMAN : SPINTURNICID MITES 127 of posterior margin of coxa II. Female deutonymphs, as far as known, have a pair of metasternal setae, five pairs dorsal propodosomal setae and a long, dorsal peritreme extending to or near level of coxa I. Chelicerae as in adult female. Male deutonymphs resemble female deutonymphs but differ in hav- ing fewer setae ventrally between coxae IV, and these are arranged in the adult male pattern. KEY TO THE SPECIES OF PERIGLISCHRUS OF THE WORLD FEMALES 1. Peritreme of normal size over coxa III, narrow and thread-like from coxa III to I ; from Natahis mexicanus natali n. sp. Peritreme of normal size throughout 2 2. Sternal plate about twice or more as wide as long; seven pairs of dorsal opistho- somal setae 3 Sternal plate longer or approximately as long as wide; less than seven pairs of dorsal opisthosomal setae 4 3. Sternal plate with broad, antero-median projection bearing first pair of setae; anterior end of adanal plate flanked by pair of shell-like sclerotized areas ; four pairs strong, prominently plumose setae on unsclerotized venter in addition to normal setae; all legs with ventral broad, flat setae prominently fringed; com- mon on Pteronotus parnellii fuscus elongatus n. sp. Anterior margin of sternal plate slightly concave; adanal plate not flanked by shell-like sclerotizations; opisthosoma lacking prominently plumose setae ven- trally; ventral leg setae of normal shape and at most narrowly fringed; from Mormoops spp strandtmanni Tibbetts 4. Several pairs of ventral body setae with grossly expanded bases and acuminate tips; first and second pairs of dorsal propodosomal setae minute; from Phyl- lostomus hastatus inflatiseta n. sp. Ventral body setae simple, not with grossly expanded bases; first pair dorsal propodosomal setae may or may not be reduced ; second pair not reduced 5 5. Femur, patella and tibia of legs III and IV each with an inflated, straight, blade- like postero-ventral seta 6 Femur, patella and tibia of legs III and IV lacking straight, blade-like postero- ventral setae ; leg IV with three apically recurved postero-ventral setae 7 6. First pair of dorsal propodosomal setae minute, usually embedded on margins of anterior dorsal plate; ratio of distance between first pair of propodosomal setae to that between first and second pairs less than 3:1; from numerous genera of phyllostomid hosts, particularly Artibeus iheringi Oudemans First pair of dorsal propodosomal setae well developed, inserted on unarmed cuticula; ratio of distance between first pair of propodosomal setae to that be- tween first and second pairs greater than 4:1; common hosts Sturnira spp. aitkeni n. sp. 7. Dorsal propodosomal setae relatively long, the longest measuring over 60 n; tibia and tarsus of legs I and II lacking inflated, recurved postero-ventral setae 8 Dorsal propodosomal setae relatively short, the longest measuring not over 50 /x; tibia and tarsus of leg I and patella and tibia of II each with an inflated, re- curved, postero-ventral seta, superficially appearing blunt 10 8. Femur II with only one of dorsal setae tiny; ratio of distance between first pair of propodosomal setae to that between first and second pairs greater than 4:1; common on Desmodus desmodi n. sp. Femur II with two of dorsal setae tiny; ratio of distance between first pair of propodosomal setae to that between first and second pairs less than 3:1 9 9. Palpal tibia with strong medio-distal lobe; leg IV with large, broadly inflated scimitar-like postero-ventral setae; dorsal opisthosoma bearing four small setae; from Glossophaga soricina caligus Kolenati 128 ECTOPARASITES OF PANAMA Palpal tibia lacking medio-distal lobe; leg IV with elongate, setaceous, curved postero-ventral setae; dorsal opisthosoma bearing six medium to large setae; hosts: Anoura, Lonchoglossa, Trachops vargasi Hoffmann 10. Sternal plate broadly jug-shaped with short, narrow neck; anterior dorsal plate longer than broad (320 // by 281 /it) ; coxa III with relatively large posterior seta; anterior legs, exclusive of ambulacrum, over 500 /j, long; hosts : common on Phyllostomus, Trachops tiptoni n. sp. Sternal plate narrowly pear-shaped with eroded margins and a long neck ; anterior dorsal plate about as broad as long measured on the mid-line (243 /a) ; coxa III with very small posterior seta; anterior legs, exclusive of ambulacrum, less than 400 /* long; common on Micronycteris micronycteridis n. sp. MALES 1. Peritreme constricted and thread-like anterior to mid-level of coxa III. . .natali n. sp. Peritreme of normal size throughout 2 2. Unarmed ventral cuticula of idiosoma with numerous minute, thornlike mam- millations 3 Unarmed ventral cuticula of idiosoma lacking such mammillations 4 3. Coxa II with two long, subequal setae; distal seta of coxa I simple, setiform elongatus n. sp. Coxa II with posterior seta much larger than anterior seta; distal seta of coxa I expanded, blade-like strandtmanni Tibbetts 4. Legs I and II with several blunt, fusiform setae; some ventral setae between coxae IV inflated inflatiseta n. sp. Not as above 5 5. Sternal plate setae short; anterior pair extending about one-half distance to level of second pair 6 Sternal plate setae relatively long; anterior pair extending about four-fifths or more of distance to level of second pair 7 6. A small mite with idiosoma less than 400 fj. long; posterior seta of coxa II about 50 n long; dorsal propodosomal setae short, not over 45 fj. long. . . .caligus Kolenati Idiosoma over 500 /* long; posterior seta of coxa II about 130 /* long; longest dorsal propodosomal setae over 65 /* long vargasi Hoffmann 7. Tarsi III and IV with coarsely barbed dorsal setae; ratio of length of posterior seta to that of anterior seta of coxa II less than 2:1 tiptoni n. sp. Tarsi III and IV superficially nude; ratio of length of posterior seta to that of anterior seta of coxa II over 2:1 8 8. Anterior dorsal propodosomal setae relatively close together; ratio of distance between bases of first pair of setae to that between bases of first and second pairs about 2:1 iheringi Oudemans Ratio of distance between bases of first pair of setae to that between bases of first and second pairs about, or more than 4:1 9 9. Three pairs setae on unarmed dorsal opisthosoma ; ventral pair setae behind sternal plate about three-fourths length of posterior setae of sternal plate, .desmodi n. sp. One pair setae on unarmed dorsal opisthosoma; ventral pair setae behind sternal plate minute 10 10. Relatively large mite with anterior legs, exclusive of ambulacra, over 500 /j. long; spermatophoral process less than 100 /j. long, shaped as a shepherd's crook aitkeni n. sp. Relatively small mite, anterior legs about 400 ^ long exclusive of ambulacra; spermatophoral process over 150 n long, extensively recurved . micronycteridis n. sp. Periglischrus natali, new species. Plate 37. Both sexes of this mite differ from other known species of the genus in the sharp constriction of the peritremes to a thread-like appearance anterior to the level of coxae III. FURMAN : SPINTURNICID MITES 129 DESCRIPTION, FEMALE (pi. 37, figs. 1, 2) : A small mite for the genus, broadly rounded anteriorly and posteriorly and with lateral margins constricted at level of anterior opistho- soma. Idiosoma 486-540 /j. long by 280-380 /j. wide. Legs short, stout. Dorsum. — Overall outline of two dorsal plates ovate. Plates separated only by lateral invaginations opposite stigmata, extending inward one-fourth width of plate, and by a suture in mid-region. Anterior plate 209 /* long on mid-line by 209 /u. wide ; slightly angulate shoulders at level of second pair propodosomal setae; with 10 pairs distinct alveoli. Posterior plate 92 fj. long on mid-line by 148 /j. wide; with nine pairs alveoli; pair minute setae in posterior pair alveoli. Anterior pair propodosomal setae 25 ^ long, others progressively smaller proceeding posteriorly; ratio of distance between bases of first pair propodosomal setae to that between first and second pairs 0.7:1. Peritremes of normal length for genus but abruptly constricted and thread-like from mid-level of coxae III to level of anterior margins of coxae II, of normal width at anterior tip and posteriorly. Unarmed opisthosoma with six pairs minute setae. Posterior tip of anal plate extending dorsally, with minute postanal seta dorsal. Venter. — Sternal plate broadly jug-shaped with short, anterior, "neck" ending bluntly ; 101 fj. long by 91 n wide; three pairs fine short (7 /j.) setae just off plate; two pairs pores on plate. Pair metasternal setae subequal to sternals, located postero-lateral to plate. Epigynial plate small, fan-shaped anteriorly, constricted centrally and slightly expanded at posterior tip; pair genital setae subequal to sternals, inserted off plate opposite con- striction. Pattern of opisthosomal sclerotized areas basically as in Periglischrus tiptoni, but not visible in some specimens. Opisthosomal setae minute, slightly longer posteriorly ; 11 pairs excluding adanal pair; adanal pair longer, 18 /j., and set anterior to anal opening on subterminal, pyriform anal plate. Legs. — Short, stout; leg I 286 /j. long exclusive of ambulacrum. All coxal setae deli- cate, minute except for posterior one of coxa II which is strong and about 73 fj. long. Setae of other leg segments relatively small for the genus, otherwise not strikingly modified. Numbers of minute dorsal setae on femora I to IV respectively are 0, 1, 2, 2. Posterior margin of femur, patella and tibia IV each with quite long, narrow, sickle-shaped seta. Gnathosoma. — Inner and outer hypostomal setae absent; distal hypostomal and gnathosomal setae subequal, simple, about 11 /x long. Palpal tibia with slight medio- distal lobe. MALE (pi. 37, figs. 3, 4) : A relatively small, elliptical to ovoid mite with setation less robust than is typical of the genus. Idiosoma 324 /JL long by 243 fj. wide. Dorsum. — Dorsal plates similar to those of female but fusion of anterior and pos- terior plates more complete; combined plates cover all but narrow lateral margins of idiosoma; anterior plate 211 /x long on mid-line by 178 /u wide; posterior plate 86 /JL long by 119 /j. wide. Propodosomal setae relatively short, anterior two pairs about 24 /* long, others slightly shorter; ratio of distance between bases of first pair to that between bases of first and second pairs about 0.55:1. Unarmed opisthosoma bearing one pair of small setae postero-laterally in addition to subequal postanal seta terminally. Peritreme more reduced than in female, only the posterior extremity to the mid-level of coxa III of normal width; constricted portion reduced to vestigial state and all but invisible even in stained specimens. Venter. — Sternal plate longer than wide, modified cordate with relatively long antero- median projection and prominent antero-lateral shoulders; four pairs marginal and one pair submedian well developed, acicular setae; posterior pairs noticeably shorter than anterior pair. Two pairs pores on plate. Pair of small setae just behind sternal plate about one-third size of posterior sternal setae. Anal plate longer than broad with ir- regular margins, extending from mid-level of coxae IV to posterior body margin, widest at mid-level. Acicular setae on and around anal plate slightly smaller than posterior sternal setae; one pair anterior to anal plate, one at level of anterior one-fifth of plate and one pair opposite constriction of posterior one-third of plate; three pairs setae on plate anterior to adanal pair of setae; latter smaller than other setae and situated just anterior to anus. Anus terminal. Pair small adanal platelets flanking postero-lateral margins of anal plate appearing typically as sinuous, longitudinally oriented bars. 130 ECTOPARASITES OF PANAMA Legs. — Leg I of allotype 297 /* long exclusive of ambulacrum. Proximal seta of coxa I and anterior seta of coxa III very small, others relatively long; posterior seta of coxa II longest, 56 p., over twice as long as next longest coxal seta. Ventral leg setae small, acicular. Dorsal leg setae acicular, nude or very minutely and sparsely barbed; few long setae, majority small to minute; one minute seta each on femora II, III and IV, none on femur I. Gnathosoma. — Pair gnathosomal setae 12 /u, long, slightly longer than distal hypo- stomal pair. Inner and outer hypostomal setae absent. Hypostomal processes elongate with membranous expansion. Usual pair of blunt, sinuous setae on tip of tibia very strongly developed. Chelicerae normal, each with spermatophoral process tubular, re- curved, about 96 n long. Tectum a simple lobe with narrowly rounded apex and broad base with slight proximal constriction. MALE DEUTONYMPH : Characteristic of the genus except that peritreme is abruptly constricted over anterior one-third of coxa III and continued anteriorly as thread-like, sinuous, dorsal line to posterior level of coxa I. TYPE MATERIAL: Holotype female and allotype male, together with 2 female, 2 male and 1 deutonymphal paratypes (host no. 6729) collected by C. M. Keenan and V. J. Tipton from Natalus stramineus mexicanus at San Lorenzo Caves, Fort Sherman (Canal Zone), 15 March 1961. Five para- types including 3 females, 1 male and 1 protonymph with same collection data as type but taken as a separate collection. Holotype, allotype and 2 paratypes in the United States National Museum; 2 paratypes, male and female, Chicago Natural History Museum; 1 male paratype, Trinidad Regional Virus Laboratory ; remaining paratypes in collection of author. ADDITIONAL MATERIAL EXAMINED : 1 female, same data as the type, and 4 males from Natalus tumidirostris haymani, Mount Tamana Caves, Trini- dad, by T. H. G. Aitken, 20 November 1957. REMARKS : This is the only spinturnicid collected from bats of the genus Natalus. Goodwin and Greenhall (1961) recorded Natalus tumidirostris haymani in Trinidad roosting in caves in association with several other species of bats, which are recorded in the present paper as hosts of five other distinct species of Periglischrus. In view of the apparent opportunity for transfer of mites from host to host under such conditions, failure to find any evidence of such transfer indicates a rather strict host specificity of P. natali to bats of the genus Natalus. Periglischrus elongatus, new species. Plate 38. Females of Periglischrus elongatus are characterized by a sternal plate which is broader than long and with the first pair of setae on a broad anterior projection of the plate ; the legs bear numerous ventral, prominent, flattened, bipectinate setae. Both sexes, in common with Periglischrus strandtmanni, have numerous minute, ventral, thorn-like mammillations. Males differ from the latter species in having a long, typically setiform, distal seta on coxa I and in having two long, subequal setae on coxa II. DESCRIPTION, FEMALE (pi. 38, figs. 1, 2) : An elongate mite with relatively short, stout legs. Idiosoma 1000 fj. long by 405 //, wide at level of coxae IV. Dorsum. — Plates relatively small ; larger anterior one 250 /j. long by 232 p wide, with- out pronounced shoulders of Periglischrus strandtmanni; with 11 pairs circular alveoli, at least two pairs with minute setae. Small posterior plate 69 p. long by 110 n wide, with seven pairs alveoli, posterior pair bearing minute setae. First pair propodosomal setae FURMAN : SPINTURNICID MITES 131 about one-half length of remaining four pairs which measure up to 55 /u long; ratio of distance between bases of first pair to that between first and second pairs 1.8 :1. Opistho- soma with seven pairs well developed setae. Peritreme normal. Anal opening dorsal, subterminal; sclerotized rim of anus subcircular; postanal sclerotization anterior to anus in form of inverted U with slightly serrated inner rim ; minute postanal seta centered on inner rim of postanal sclerotization. Venter. — Sternal plate shorter than broad, 61 /* long on mid-line by 128 n wide, with broad anteromedian projection bearing first pair setae; posterior margin slightly con- cave in middle; three pairs sternal setae subequal, approximately 28 /JL long; second and third pairs located on posterior margin of plate; two pairs rounded pores on plate. Pair metasternal setae about one-half length of sternal setae, located midway between sternal and epigynial plates. Genital opening a small transverse slit in mid-line at level of pos- terior margins of coxae III. Epigynial plate a narrow longitudinal strip 67 /JL long by 9 /j. wide bearing two prominent, lightly pilose, long setae, 61 /JL long, on posterior margin. Adanal pair setae similar to genital setae but somewhat shorter, arising from posterior margin of ventral platelet which is a narrow, elongate bar, slightly enlarged posteriorly and widely separated from dorsally situated anus ; anterior end of platelet in a cavity bordered by two lateral shell-like sclerotized areas. Opisthosomal cuticula minutely striated, bearing nine pairs setae; pair larger setae anterior to adanal setae and two pairs on postero-lateral margins prominently bipectinate; other setae smaller and most arising from cup-shaped alveoli as illustrated. Podosomal cuticula in region of sternal plate furnished with numerous minute, thorn-like mammillations. Legs. — Relatively short and stout; legs I 383 LL long exclusive of ambulacrum; pos- terior seta of coxa II modified to appearance reminiscent of palmate hairs of Anopheles mosquito larvae, with enlarged, flattened basal section gradually tapering toward apex and bipectinate throughout; distal seta coxa I slightly shorter than posterior seta of II, typically setiform, sparsely and minutely barbed; anterior setae of II and III similar but smaller; other coxal setae simple, short. Coxa II with antero-dorsal marginal spur and coxa III with posterior marginal spur. Prominent palmate setae similar to that of coxa II present on ventral side of most other leg segments distal to coxae, in addition to unmodi- fied setae, several of which are slightly pectinate. Dorsal setae well developed, simple; long setae very minutely and sparsely barbed ; no minute setae on femora. Gnathosoma. — Pair distal hypostomal setae 16 M long, somewhat longer than gnatho- somal pair. Inner hypostomal setae absent; outer hypostomal pair represented only by pair of small alveoli. Palpi unmodified except for sclerotized ventro-medial, bidentate to tridentate flange on each trochanter, flanking mouthparts. Chelicerae normal. MALE (pi. 38, figs. 3-5) : A relatively long-legged mite with ovoid body 350 /JL long by 285 n wide in allotype. Other specimens up to 430 /* long. Dorsum. — Dorsal plates closely approximated and covering all but narrow margin of idiosoma. Anterior plate 226 /JL long by 235 /JL wide, with narrowly rounded anterior margin, moderate shoulders over coxae I and widest at level between coxae II-III; posterior margin truncate and angularly concave; surface with 12 pairs alveoli. Pos- terior plate 75 /JL long by 104 n wide; subtriangular ; surface with seven pairs alveoli, posterior pair bearing minute setae. Propodosomal setae all of moderate length, up to 34 fj. long; ratio of distance between bases of first pair to that between first and second pairs 1.7-2.4:1 based on four specimens. Peritremes normal for genus except that they bend laterad markedly between coxae II and III and dip to ventral surface at this point on some specimens. Unarmed opisthosoma lacking setae. Venter. — Sternal plate broadly flask-shaped with long anterior neck, 151 /JL long on mid-line by 140 /JL wide at level of second pair setae; with four pairs marginal and one pair post-centrally located setae; anterior pair 33 /* long, others subequal or slightly shorter; surface with pattern of transverse lines. Pair submedial setae posterior to sternal plate three-fourths length of posterior sternal setae. Five pairs simple setae, subequal to posterior sternals, located between coxae IV on unarmed cuticula; adanal setae stouter and longer and located on small anal plate just anterior to terminal anus; anal plate small with ill-defined margins. Postanal seta minute and dorsal. Ventral cuticula of idiosoma furnished with minute thorn-like mammillations as illustrated. 132 ECTOPARASITES OF PANAMA Legs. — Relatively long ; leg I 470 /u long exclusive of ambulacrum. Coxal setae simple, well developed; anterior and posterior setae of coxae II subequal, other segments ven- trally with setae normal, relatively short. Coxa II with antero-dorsal marginal spur and coxa III with posterior marginal spur. Dorsal setation normal for genus with all setae essentially nude. Gnathosoma. — Gnathosomal and distal hypostomal setae simple, small, subequal. Outer hypostomal setae vestigial, set in relatively prominent alveoli. Inner hypostomal setae absent. Hypostomal processes long, elliptical, each with membranous, blade-like expansion. Palpi relatively long, thin. Chelicerae each with short, recurved spermato- phoral process about 55 fi long. FEMALE DEUTONYMPH : Idiosoma shape similar to that of male, 432 fj. long by 356 ^ wide, but legs relatively stout in relation to length; leg I 443 /u long exclusive of ambu- lacrum. Peritreme extending only to level of posterior one-fifth of coxa II, posterolaterad of dorsal propodosomal seta IV. Dorsal plates as in male. Venter with angulate mam- millations as in male. Sternal plate subcircular with anterior margin slightly projected; surface with transverse lines; three pairs setae on plate. Two pairs setae in space be- tween sternal plate and level of anterior margin of coxae IV. Unarmed cuticula between coxae IV and anterior to anal plate with seven pairs simple setae. Three pairs minute peg-like setae lateral to anal plate. Anal plate ovate, broader posteriorly, terminal, with pair well developed adanal setae. MALE DEUTONYMPH : Idiosomal shape as in female deutonymph but legs not as stout in relation to length. Dorsum as in female deutonymph; typical specimen appears to have narrower margin of body unprotected by dorsal plates. Sternal plate with antero- median lobe more pronounced than in female deutonymph; three pairs setae on plate and three pairs immediately posterior to it. Unarmed cuticula between coxae IV and anterior to anal plate with five pairs simple setae; lacking three pairs peg-like setae lateral to anal plate. Angulate mammillations and other ventral features as in female deutonymph. PROTONYMPH : A broadly elliptical mite with short, stout legs bearing rather delicate setae. Idiosoma 395 /* long by 319 /JL wide. Leg I 405 ,u long exclusive of ambulacrum. Dorsal plates as in male. Only four pairs propodosomal setae similar in size to those of male. Peritreme dorsal, extending anteriorly only to space between coxae II and III, ending on lateral margin of body or even ventro-laterally. Sternal plate margins in- distinct, with three pairs well developed, simple setae and two pairs pores ; surface trans- versely sculptured. Ventral area between coxae IV and anterior to anal plate with four pairs simple setae. Leg setae simple, nude. Gnathosomal and distal hypostomal setae relatively longer than in male. Outer hypostomal setae small but distinct, much larger than in male. TYPE MATERIAL: Holotype female (colln. no. T275) together with para- type female collected by W. G. Downs from "Chilonycteris rubiginosa fusca" at Mt. Tamana Caves, Trinidad, 12 June 1956. Allotype male and one para- type male (colln. no. T269) with same collection data as type. Other para- types include 2 protonymphs with same data as type; 13 males, 1 female deutonymph and 1 protonymph from same host and locality taken by T. H. G. Aitken, 20 November 1957 ; 2 males from same host at Heights of Guanapo, Trinidad, by T. H. G. Aitken, 19 May 1957, and 1 female, 5 males and 3 proto- nymphs with same data but collected 1 October 1957 ; 6 females, 31 males, 3 female deutonymphs, 4 male deutonymphs and 8 protonymphs from same host species taken at Paraiso (Canal Zone), by C. M. Keenan and V. J. Tipton, 24 July 1959. Holotype, allotype and 9 paratypes, including 4 males, 1 female, 2 proto- nymphs and 2 deutonymphs (male and female), in the United States Na- tional Museum ; 8 paratypes, including all stages, Chicago Natural History FURMAN : SPINTURNICID MITES 133 Museum; 4 paratypes (males and females), Trinidad Regional Virus Labo- ratory ; remaining material in the collection of the author. ADDITIONAL MATERIAL EXAMINED: From Pteronotus parnellii fuscus: Paraiso (Canal Zone) , 16 September 1959 ; Madden Air Field (Canal Zone) , 23 May and 3 October 1961; Chilibrillo Caves (Panama), 2 August 1960; Bocas del Toro Province, 1 February 1960; Cerro Hoya (Los Santos), 18 February 1962. From Pteronotus suapurensis, a single collection of 3 males, Chilibrillo Caves (Panama), collected by C. M. Keenan and V. J. Tipton, 8 March 1960. REMARKS : Periglischrus elongatus is most closely related to Periglischrus strandtmanni Tibbetts, which also occurs on chilonycterine bats, but of the genus Mormoops. The question of degree of intraspecific variation in Periglischrus elon- gatus is still open. Two abnormal female specimens are at hand from Pteronotus parnellii fuscus taken in Peninsula de Azuero (Los Santos Prov- ince), 24 February 1962. In these the characteristics of the species are for the most part greatly exaggerated. The legs are stouter and more heavily sclerotized, and more of the ventral setae are modified into heavily fringed "palmate" setae. More striking is the presence of a strong, deep camerostome bordered posteriorly by the sternal plate and laterally by heavily sclerotized margins. The dorsal propodosomal setae are modified into short, stout, blunt spines. Whether these two specimens represent genetic freaks or are actually representative of a distinct population is un- known. The single male at hand from this area is apparently normal. Periglischrus strandtmanni Tibbetts Periglischrus strandtmanni Tibbetts, 1957, Jour. Kansas Ent. Soc., 30, (1), p. 14, pis. 1-3 — United States National Museum, Washington (Frio Cave, Uvalde County, Texas, from Mormoops megalophylla senicula). Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 199. This mite has been recorded previously only from the type collection, from Texas, and while it has not been collected in Panama, a recent collection from Trinidad indicates its probable occurrence in Panama. Comparison of a female specimen from Trinidad with Tibbetts' descrip- tion and figures demonstrates close agreement. However, the "Y"-shaped anal plate with two long posterior setae described by Tibbetts as dorsal in po- sition is actually ventral just as in the new species, P. elongatus, described in the present publication; the two long setae are adanal setae. Tibbetts' species lacks the pair of adanal, shell-like sclerotized platelets flanking the anal plate of P. elongatus. The anal opening and minute postanal seta are dorsal in position and appear just as described for P. elongatus. Many of the ventral leg setae are sparsely bipectinate, but none show the inflated "palmate" appearance characteristic of P. elongatus. Other differential characters are given in the discussion of the latter species. The specimen recorded here is a female taken from Mormoops mega- lophylla tumidiceps Miller, at Mount Tamana Caves, Trinidad, by T. H. G. Aitken, 20 November 1957. 134 ECTOPARASITES OF PANAMA Periglischrus inflatiseta, new species. Plate 39. The female of Periglischrus inflatiseta differs from all other members of the genus in possessing characteristically inflated setae on the venter. The male is very similar to Periglischrus tiptoni from which it differs in its smaller size, its much shorter and broader, blunt, ventral setae on legs I and II, shorter anterior legs, and in the absence of strongly serrated setae on tarsus III. DESCRIPTION, FEMALE (pi. 39, figs. 1, 2) : A relatively small mite for the genus with idiosoma of gravid specimens broadly rounded anteriorly and opisthosoma moderately flared in fan shape. Idiosoma of type specimen 760 fj, long by 454 p wide. Dorsum. — Overall outline of two dorsal plates ovate with broad end anterior. An- terior plate 230 fj, long on mid-line by 244 jj, wide; broadest at level of coxae II; with 10 pairs of obvious alveoli, some bearing microsetae; a longitudinal median slit seen in all mounted specimens may represent merely a zone of weakness resulting in an artifact; posterior margin truncate and bridged to posterior plate by two small, submedian lobes. Posterior plate approximately triangular with posterior apex rounded; 83 fj. long by 151 fj. wide; seven pairs alveoli of which subterminal pair bear microsetae. Propodosomal setae minute, especially first two pairs; ratio distance between bases first pair setae to that between first and second pairs approximately 2:1. Pair of minute metapodosomal setae mesad of stigmata. Four pair minute setae on opisthosoma. Minute postanal seta subterminal. Peritremes extend almost to level of second pair propodosomal setae. Venter. — Sternal plate longer (100 /j.) than wide (98 /j.) , pentagonal with rounded corners, concave antero-lateral margins, and straight to concave posterior margin ; three pairs short simple setae on margins, first pair 18 fj. long, others 26 n long. Pair meta- sternal setae lateral to posterior corners sternal plate, strongly sclerotized, flattened and broadly fusiform basally, abruptly acuminate distally, 39 n long by 15 /u. wide. Pair epigynial setae similar to metasternals but smaller, arising from small, poorly sclerotized epigynial platelet behind transverse genital slit; small, fan-shaped pattern of striae on cuticula anterior to genital slit. Opisthosomal pattern of sclerotization similar to that of P. Iheringi. Opisthosoma with 10 pairs setae in addition to adanal pair: pair small, normal, submedian setae on anterior of opisthosoma followed in diverging rows posteriorly by two pairs prominent inflated setae of same type as metasternals but larger; smaller inflated pair submedian setae at mid-opisthosomal level; remaining setae on posterior half of opisthosoma normal, slightly fusiform, arranged as in P. tiptoni. Anal plate as described for P. tiptoni. Legs. — Short, stout; leg I 405 /j. long exclusive of ambulacrum. Coxal setae well developed: I with two acicular setae, basal slightly longer than distal; anterior seta of II fusiform, posterior one longer than others (72 /*), minutely barbed; anterior seta of III small, normal, posterior one large, 34 ^ long, broadly fusiform, similar to metasternals ; seta of IV well developed, normally setiform with slightly fusiform base. Ventrally trochanter, femur and tarsus I and segments of II exclusive of coxa, each with one or more prominent, short, stout, blunt, flattened setae with tendency toward slight barbs on one edge; III and IV with several strong postero-ventral setae, fusiform basally, acutely tipped. Dorsally setal vestiture normal for genus with long setae minutely barbed; femora I and II each with minute antero-basal seta; III and IV each with two small basal setae. Gnathosoma. — Gnathosomal setae small, acicular ; distal hypostomal pair similar, but twice as long; outer hypostomal pair vestigial; inner pair absent. Palpal tibia with very slight medio-distal lobe; trochanter with blunt, ventral, flap-like emargination distally. Chelicerae normal for genus. MALE (pi. 39, figs. 3, 4) : Idiosoma broadly ovoid, widest at level of coxae II-III; 405 fj. long by 356 /j, wide. Opisthosoma rudimentary. Dorsum. — Dorsal plates as in Periglischrus tiptoni but smaller; anterior plate 246 n long by 246 n. wide; posterior plate 120 /j. long by 148 /j. wide. Propodosomal setae short, FURMAN : SPINTURNICID MITES 135 subequal, 16-20 fj, long; ratio of distance between first pair of setal bases to that between first and second pairs approximately 2:1. Unarmed opisthosoma bearing one pair small setae. Venter. — As described for Periglischrus tiptoni with following exception: first and third pair of large setae in space between coxae IV basally inflated. Legs. — Leg I 440 ^ long exclusive of ambulacrum. Coxae with relatively long setae ; posterior seta of II 59 p. long, that of III 48 ^ long. Ventrally trochanter and femur I each with two prominent, blunt, broadly fusiform, short setae; similar though less prom- inent setae on patella and tibia; tarsus I with short, broad fusiform seta proximal to mid-level. Ventrally leg II with short, broad fusiform setae on all segments but coxa. Ventral setae of III and IV tend to be basally inflated and terminate in acute tips. Tarsi III and IV ventrally each with one proximal and a pair of short, subapical, bluntly con- ical, stout setae in addition to others. Dorsally femur I with one to two minute setae, femur II with one, femur III with two, and femur IV with one minute setae. Tarsus IV with pair strongly serrate dorsobasal setae; tarsus III lacking serrate setae. Gnathosoma. — Gnathosomal and distal hypostomal setae subequal, 17 fj. long; outer hypostomal setae minute; inner hypostomals absent. Similar in general appearance to P. tiptoni but lateral seta of palpal femur not prominently serrated, and two stout apical setae of tibia are particularly well developed in P. inflatiseta. TYPE MATERIAL : Holotype female, allotype male, and 5 paratypes includ- ing 3 females, 1 male and 1 female deutonymph (host no. 43065) collected by V. J. Tipton from Phyllostomus hastatus panamensis at Panama City (Panama) , June 1961. Paratype series also includes the following : 1 female from Phyllostomus hastatus panamensis, Chepo Road (Panama), C. M. Keenan and V. J. Tipton, 8 October 1959. From Phyllostomus hastatus hastatus, 1 female and 2 males, Heights of Guanapo, Trinidad, T. H. G. Aitken, 19 May 1958 ; a female, male and protonymph, at Heights of Guanapo, Trinidad, W. G. Downs, 7 November 1956 ; 3 females and 1 male from same location, T. H. G. Aitken, 11 August 1957. Holotype, allotype and 2 paratypes, male and female, in the United States National Museum ; 2 paratypes (male and female) , Chicago Natural History Museum and Trinidad Regional Virus Laboratory; remaining material in the collection of the author. OTHER MATERIAL EXAMINED: In addition to the paratypes listed above, 1 male and 2 females were taken from (2) Phyllostomus hastatus by K. von Sneidern at La Macarena, Rio Guapaya, Colombia, 14 March 1957. REMARKS : Periglischrus inflatiseta is usually found in association with Periglischrus tiptoni on subspecies of Phyllostomus hastatus, but the more common tiptoni apparently occurs on a wider range of host species. Periglischrus iheringi Oudemans Periglischrus iheringi Oudemans, 1902, Ent. Ber., 1, (6), p. 38 — Rijksmuseum fur Naturlijke Historic, Leiden (Sao Paulo, Brazil, from Vampyrops lineatus). Rud- nick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 197, pi. 30. This species was adequately redescribed by Rudnick (1960) . Specimens identified as this species in the current study agree closely with Rudnick's description. Slight intraspecific differences appear in some characters such as the length of the small antero-basal seta on the dorsal aspect of femur II of the female. For example, in specimens from Artibeus lituratus pal- marum this seta is consistently larger than in specimens from Artibeus j. 136 ECTOPARASITES OF PANAMA jamaicensis; in Rudnick's (1960) figure of this species he has omitted the small, dorsal, antero-basal seta of femur II. On some females three pairs of ventral alveoli can be seen, arranged in two diverging rows between and posterior to coxae IV ; minute setae are visible in the alveoli on some well- mounted specimens. The female may be distinguished from other species by the following combination of characters : A large mite with expansile, fan-shaped opis- thosoma. Palpal tibia lacking prominent subapical lobe. Anterior pair dorsal propodosomal setae tiny, usually inserted on dorsal plate; other propodosomal setae long (up to 67 ^} . Ratio of distance between bases of first pair propodosomal setae to that between bases of first and second pairs less than 3 :1. Posterior seta coxa II much longer than other coxal setae, minutely barbed ; posterior seta coxa III approximately 55 ^ long, inflated, blade-like; postero-ventral margins legs III and IV with several straight, blade-like setae. Proximal dorsal seta of femur, genu and tibia II long. The male of Periglischrus iheringi resembles that of the new species Periglischrus aitkeni. It may be differentiated by the longer, dorsal, pro- podosomal setae (up to 73 //. long), by the longer spermatophoral processes (198 //, long) and by the position of the anterior pair of dorsal propodosomal setae which are more closely approximated ; the ratio of the distance between the bases of the first pair of propodosomal setae to that between the first and second pair ranges from 1.7 :1 to 2.2 :1 (19 specimens measured) ; this com- pares to a ratio of 5.5 :1 for P. aitkeni. From the closely related Periglischrus vargasi, males of P. iheringi may be distinguished by the presence of, at most, one small to medium-sized dorsal seta on femur II, in contrast to two minute to small setae in this position on P. vargasi. Periglischrus iheringi is the most commonly encountered spinturnicid in collections seen from Panama. Not only does it occur in considerable num- bers on its hosts, but it infests an unusually large number of bat genera of the family Phyllostomidae, and even extends to the Desmodidae. MATERIAL EXAMINED : The following new distribution records represent collections made by V. J. Tipton alone or with C. M. Keenan. Each collection was from a single bat. From Uroderma b. bilobatum: Summit Gardens (Canal Zone) , 3 collections of 1 female, of 1 male, and of 4 females, 2 males and 1 nymph, 11 September 1959; Bocas del Toro Province, 3 females, 5 February 1960; Escobal (Colon), 1 female, 28 September 1960; Rodman Dispensary (Canal Zone), 5 females, 2 males and 3 nymphs, 27 April 1961. From Vampyrops vittatus: Cerro Punta (Chiriqui), 2 females, 2 males, 2 nymphs, 5 February 1960; Rio Changena (Bocas del Toro), 3 females, 4 males, 1 nymph, 24 September 1961, and 2 males, 1 nymph, 20 September 1961. From Vampyrops helleri: Fort Sherman (Canal Zone), male and nymph, 2 December 1959; Almirante (Bocas del Toro), 2 collections of 2 males and of a female, male, 2 nymphs, 23 January 1960, and of a female and 2 males, 24 January 1960; Cerro Hoya (Los Santos), 1 male, 10 February 1962, 1 male on 13 February 1962 and 1 female on 24 February 1962. From Vampyressa pusilla: from Cerro Hoya, two collections of 1 female and of 1 nymph on 10 February 1962, 3 females and 1 male on 22 February 1962 and 2 females on 24 February 1962. From Vampyrodes major: Bocas del FURMAN : SPINTURNICID MITES 137 Toro Province, 1 male, 25 January 1960; Pena Point (Darien), two collec- tions comprised of 3 females, 3 males, 2 nymphs, 24 March 1960 ; Rio Sete- ganti (Darien), 1 female and 1 male on 1 February 1961, 1 male on 23 Sep- tember 1961 and 2 females, 1 male and 2 nymphs on 24 September 1961. From Chiroderma salvini: Rio Changena (Bocas del Toro), 1 male, 22 Sep- tember 1961. From Artibeus toltecus: Casa Lewis, Cerro Punta (Chiriqui) , two collections of 1 female and 1 nymph and of 1 female on 5 February 1960, and of 1 male on 3 February 1960 ; Rio Changena (Bocas del Toro) , two col- lections of 1 female each, 20 September 1961. From Artibeus j. jamaicensis: Fort Kobbe (Canal Zone), numerous specimens, 13 October 1960, and 1 male and a nymph on 24 July 1959 ; Bocas del Toro Province, 4 collections of 1 male, of 6 females, of 1 female and of 2 females, 1 male and 1 nymph on 23 January 1960 ; other collections from the same area are 3 collections of several specimens on 24 January 1960, two collections totaling 5 specimens on 25 January 1960, 1 female on 26 January 1960, one to several specimens from single collections on 27, 28 and 30 January 1960 and 1 February 1960 ; Pena Point (Darien), female and nymph, 24 March 1960; Fort Clayton (Canal Zone), 3 females, 19 October 1960; Rio Seteganti (Darien), 8 col- lections of from 1 to 5 specimens each from 1 to 5 February 1961, and 2 collections of several specimens each on 17 and 20 September 1961. From Artibeus j, jamaicensis: Rio Changena (Bocas del Toro), female, 2 males and nymph on 21 September 1961, 1 male and nymph on 22 September 1961 and 1 female on 23 September 1961. From Artibeus lituratus palmarum: Paraiso (Canal Zone) , male, 5 November 1959 ; Fort Sherman (Canal Zone) , male and nymph, 4 December 1959; Fort Clayton (Canal Zone), several, 19 October 1960; Bocas del Toro Province, male, 24 January 1960; Rio Seteganti (Darien) , 7 collections from 1 to 5 February 1961, each containing from one to several mites ; Juan Mina (Canal Zone) , female, 29 June 1961 ; from Cerro Hoya, one female, 21 February 1962. From Artibeus cinereus: Rio Changena (Bocas del Toro) , 4 collections, 18 to 24 September 1961, each containing from one to several mites ; from Cerro Hoya, one female, 12 Feb- ruary 1962. From Artibeus aztecus: Chiriqui Province, 2 collections of 1 female and 2 females, 2 March 1962. From Artibeus species : Rio Changena (Bocas del Toro), several specimens, 24 September 1961. From Enchis- thenes harti: Cerro Hoya (Los Santos), 3 females, 5 males, 8 March 1962; from same locality, 4 collections of 1 to 3 mites each, 9 to 18 February 1962. From Desmodus rotundus murinus: Bocas del Toro Province, female and male, 23 January 1960, and 2 males, 25 January 1960; Cerro Punta (Chi- riqui), 2 females, 5 February 1960. Periglischrus aitkeni, new species. Plate 40. Periglischrus aitkeni resembles Periglischrus iheringi, from which fe- males may be distinguished in having the first pair of dorsal propodosomal setae well developed and inserted very near the bases of the second pair of propodosomal setae. Males differ in having a relatively short spermato- phoral process, less than 100 /x long, and in having the anterior, dorsal pro- podosomal setae displaced far laterad. 138 ECTOPARASITES OF PANAMA DESCRIPTION, FEMALE (pi. 40, figs. 1, 2) : A large, typical number of the genus with idiosoma broadly rounded anteriorly and opisthosoma moderately flared. Idiosoma 1134 /* long by 594 /u, wide. Dorsum. — Anterior plate approximately 310 /j long on mid-line by 275 fj. wide, with broadly rounded anteromedial projection, and pronounced shoulders; sides slightly con- cave anteriorly and slightly convex posteriorly; posterior margin truncate, slightly con- cave and joined to posterior plate by two submedian lobules; 11 pairs alveoli in pattern similar to that in Periglischrus iheringi. Posterior plate bluntly triangular, 130 ^ long by 190 /j. wide, widest anteriorly and terminating in broadly rounded posterior margin; bearing five pairs prominent alveoli, the posterior pair bearing microsetae. Propodosomal setae all moderately developed, second pair 45-50 /* long, others obviously shorter; ratio of distance between bases of first pair to that between first and second pairs approxi- mately 5:1. Unarmed opisthosoma with four pairs small setae. Postanal seta minute, subterminal. Venter. — Sternal plate as broad as long, 146 n, outline that of broadly rounded penta- gon with narrowest angle anterior; three pairs small sternal setae located just off margins of plate; two pairs pores on plate, in some specimens serving as foci for invaginations of eroded plate margins. Epigynial platelet a small, narrow, longitudinal sclerite with membranous posterior enlargement; pair small genital setae located off plate anterior to posterior lobe. Pattern of sclerotized opisthosomal areas and opisthosomal setation as in P. tiptoni and P. iheringi, but anterior three pairs setae vestigial or absent and represented only by alveoli; other setae small. Anal plate as described for P. tiptoni, but adanal setae 33 n long, over twice as long as other opisthosomal setae. Legs. — Well developed, stout; leg I 545 fj, long exclusive of ambulacrum. Posterior seta coxa II 195 n long, minutely barbed; posterior seta coxa III 39 /j. long, broadly blade- like; other coxal setae much smaller, acicular. Other ventral leg segments with mostly short, simple setae, usually with inapparent, minute barbs on longer setae; several pos- terior setae of legs III and IV expanded, blade-like. Longer dorsal setae minutely barbed. Dorsal surfaces of femora with a minute seta only on legs II and III. Gnathosoma. — Distal hypostomal setae 22 /* long, about twice as long as gnathosomal pair. Inner and outer hypostomal setae apparently absent. Palpal tibia lacking pro- nounced medio-distal lobe; trochanter with blunt, ventral, flap-like emargination distally. Chelicerae normal for genus. MALE (pi. 40, figs. 3-5) : Idiosoma ovoid, widest at level of coxae II-III; 600-620 M long by 488-513 /j. wide. Legs stout, of moderate length. Dorsum. — Shape of dorsal plates and alveolar pattern similar to those of Periglis- chrus iheringi; some of alveoli bearing microsetae as illustrated. Propodosomal setae well developed, setiform, measuring 49-55 n long; ratio distance between bases of first pair to that between first and second pairs approximately 5.5:1. Unarmed opisthosoma with pair of short (18 /j.) setae subterminally. Postanal seta minute, dorsoterminal. Venter. — Sternal plate longer than wide, modified cordate with pronounced medial anterior projection and prominent shoulders; with five pairs well developed acicular setae, anterior-most of which extend to level of imaginary line projected between bases of second pair; pair of minute setae behind sternal plate. Anal plate large, elongate, occupying most of space between coxae IV, truncate anteriorly with sides constricted slightly just anterior to adanal setae; adanal setae inserted just anterior to anus; three other pairs subequal setae anteriorly on anal plate and three pairs bordering plate. Pair of subterminal accessory platelets bordering posterior end of anal plate. Legs. — Well-developed, stout; leg I approximately 540 /* long exclusive of ambula- crum. Coxal setae all well developed, typically setiform; posterior seta of coxa II over twice as long as other coxal setae, approximately 165 /j. long; anterior seta of coxa III shortest, 30 /j. long. Other leg segments ventrally with short but strong simple setae; ventro-lateral setae of legs III and IV longer and minutely barbed; pair of short sub- terminal setae of tarsus II spine-like. Long dorsal setae minutely barbed and notched apically. Femur II with only one of dorsal setae minute; no minute setae on other femora. FURMAN : SPINTURNICID MITES 139 Gnathosoma. — Gnathosomal and distal hypostomal setae subequal, well developed, 28 PL long. Inner hypostomal setae absent ; outer hypostomals vestigial but alveoli easily observed. Palpi normal for genus. Spermatophoral process a strong tubular structure recurved in shallow hook distally, short for the genus, measuring approximately 80 /j, long in allotype. TYPE MATERIAL: Holotype female and allotype male (colln. no. T60) to- gether with 3 paratype females collected by T. H. G. Aitken from Sturnira lilium lilium, at the 15% mile mark on Churchill-Roosevelt Highway, Trini- dad, 17 June 1958. Other paratypes are as follows : 3 females from Sturnira lilium parvidens, at the Rio Mandinga (San Bias), V. J. Tipton, 27 May 1957 ; 5 females from Sturnira lilium, at Rancho Grande, Venezuela, C. O. Handley, 30 March 1960 ; 3 females and 1 protonymph from the same host, Los Santos Province, V. J. Tipton, 24 February 1962; one female from Sturnira ludovici, Cerro Punta (Chiriqui), Republic of Panama, C. M. Keenan and V. J. Tipton, 30 April 1961. Holotype, allotype and 3 female paratypes, in the United States National Museum; 2 paratypes (male and female) , Chicago Natural History Museum and Trinidad Virus Laboratory ; remaining material in the collection of the author. OTHER MATERIAL EXAMINED : From Sturnira lilium, Bocas del Toro Prov- ince, 1 female, C. M. Keenan and V. J. Tipton, 23 January 1960; from Sturnira ludovici, Cerro Punta (Chiriqui) , C. M. Keenan and V. J. Tipton, 1 male and 1 female deutonymph on 2 February 1960, 1 female, 1 male and 1 male deutonymph on 3 February 1960, 2 females on 3 February 1960, 1 male deutonymph on 3 February 1960, 1 male and 1 protonymph on 15 Feb- ruary 1960, 1 female, 1 May 1960, and 1 male on 1 May 1960 ; from the same host, Chiriqui Province, a male, male deutonymph and a protonymph, V. J. Tipton, 6 March 1962 ; from Sturnira species in Los Santos, 3 females and 2 protonymphs on 1 host specimen and 2 males on another specimen, V. J. Tipton, 6 March 1962; from Noctilio leporinus at Guanico (Los Santos), 5 females and 2 males, V. J. Tipton, 24 January 1962. REMARKS: Periglischrus aitkeni is a rather common parasite of phyl- lostomid bats of the genus Sturnira in Central America. In view of the usually rather restricted host range of Periglischrus, it was surprising to find several typical specimens of P. aitkeni from a single Noctilio leporinus, a fish-eating bat of the superfamily Emballonuroidea. This species is name in honor of Dr. T. H. G. Aitken, entomologist at the Trinidad Regional Virus Laboratory of the Rockefeller Foundation, who collected the type specimens and has provided many of the other collections upon which this study is based. Periglischrus desmodi, new species. Plate 41. Periglischrus desmodi is closely related to P. vargasi. Characters dis- tinguishing the female include the very widely spaced first pair of dorsal propodosomal setae, the characteristic shape of the sternal plate and the presence of only one minute basal seta on the dorsum of femur II instead of two. Characters distinguishing the male include long sternal plate setae, three pairs of dorsal opisthosomal setae and a pair of well-developed setae behind the sternal plate. 140 ECTOPARASITES OF PANAMA DESCRIPTION, FEMALE (pi. 41, figs. 1, 2) : Idiosoma broadly rounded anteriorly, with opisthosoma slightly to moderately expanded; approximately 920 /u. long by 486 n wide; a moderately robust species with long setae. Dorsum. — Anterior dorsal plate large, 297 /u long on mid-line, by 292 /j, wide; rounded apex projecting from broad anterior margin ; broad, relatively straight posterior margin at level of coxae IV, with two submedian rounded protuberances received by anterior margin of posterior plate; bearing nine pairs of prominent alveoli, three pairs of which bear minute setae. Posterior plate small, with anterior margin contacting anterior plate with two submedian emarginations to receive corresponding projections from anterior plate; broader than long with posterior margin broadly rounded; with eight pairs of alveoli, of which two pairs bear minute setae. Propodosomal margin with five pairs long, subequal setae of approximately 85 //; ratio of distance between first pair to that be- tween first and second pairs varies from 6.6:1 to 8.4:1. Opisthosoma with six pairs setae, of which the longest are anterior. Posterior tip of anus normally dorsal, bearing minute postanal seta. Venter. — Sternal plate a modified pentagon longer than wide, with broad base, rounded sides and narrowly tapering anterior apex; three pairs of short setae may be off plate although on type first and third pair are included in plate; two pairs of pores on plate. Pair of short metasternal setae. Epigynial plate reduced, slender, long, slightly expanded anteriorly; pair of short genital setae located off plate, insertions spaced 10 n apart. Opisthosoma bearing three pairs small setae in two diverging rows behind epi- gynial plate; three pairs on median sclerotized area anterior to anal plate, two pairs anterior to this sclerotization, two pairs postero-lateral and one pair lateral to the sclero- tization. Sclerotized areas of opisthosoma as illustrated, similar to to those of P. iheringi. Anal plate ventro -terminal, elongate, narrow with pair adanal setae subequal to adjacent opisthosomal setae; canal-like structures extending anteriorly from adanal setal bases. Minute postanal seta dorsal. Legs. — Relatively short, stout; leg I 476 ^ long exclusive of ambulacrum; coxal setae, with exception of posterior seta of II, short, nude and unmodified; posterior seta of II elongate and minutely barbed; most other ventral leg setae normal with larger setae minutely barbed ; legs I and particularly II with row of inflated, leaf-like setae very prom- inently serrated; femur, patella and tibia IV each with a slightly inflated, scimitar- shaped, postero- ventral seta; dorsal setae of legs strong, many elongate; of two basal setae on femur II the posterior one is of medium size and anterior one is small. Gnathosoma. — Inner and outer hypostomal setae absent; pair distal hypostomal setae about twice as long as short gnathosomal pair; long paired hypostomal processes present. Palpi with five movable segments; tibia with moderately developed medial distal lobe. Chelicerae normal for genus. MALE (pi. 41, figs. 3-5) : Idiosoma ovoid, widest at level of coxae II— III; 485 fj. long by 350 n wide. Opisthosoma rudimentary. Dorsum. — Anterior and posterior dorsal plates similar to those of female in size, shape, surface markings and setation, although the posterior plate tends to have concave postero-lateral margins and the plates appear compressed together, rendering the line of separation indistinct; the two plates cover most of idiosoma. Propodosomal setae as in female but approximately 60-67 p long ; ratio of distance between alveoli of first pair to that between first and second pairs approximately 6:1. Three pairs small setae on narrow cuticular margin bordering postero-lateral edges of posterior dorsal plate; post- anal seta minute, terminal. Venter. — Sternal plate modified cordate with elongate antero-median projection ter- minating at male genital opening; longer than wide; bearing five pairs well-developed setae ranging from 63 p. long anteriorly to 43 /j. for posterior pair. Pair of medium-sized setae 30 /* long posterior to plate. Five and occasionally six pairs setae up to 36 ft long located in space between coxae IV and anterior to anal plate. One pair setae lateral to anal plate and mesad of pair of small adanal platelets. Anal plate small, elongate, with indistinct anterior margins; pair of adanal setae about 20 n long anterior to anal open- ing, which is terminal. FURMAN : SPINTURNICID MITES 141 Legs. — Leg I 450 M long exclusive of caruncle. With exception of posterior seta of coxa II, coxal setae longer than in female although of same type; other ventral leg setae simple, acicular, small ; dorsal setae similar to those of female, but tarsus I bearing two long, specialized, blunt setae. Gnathosoma. — Pair gnathosomal setae 18 //, long, acicular; outer hypostomal pair minute; distal hypostomal setae subequal to gnathosomal pair. Palpal tibia lacking medial lobe of female ; trochanter with distal flange-like lobe. Chelicerae with fixed digit well developed, bearing numerous sharp subterminal "teeth" on surface opposed to mov- able digit; latter slightly longer than fixed digit and bearing blunt-tipped teeth along inner surface; spermatophoral process a long tubular, recurved structure over 150 /j. in length. MALE DEUTONYMPH : Idiosoma shape similar to that of male, 497 /j. long by 362 /* wide. Dorsum. — Similar to that of male. Venter. — Sternal plate approximately diamond-shaped with corners rounded, bear- ing three pairs well-developed acicular setae; pair setae bordering postero-lateral margins of plate. Anal plate and remaining setae of venter as in male. Legs. — As in male but lacking long, blunt-tipped setae on tarsus I. Gnathosoma. — Similar to that of male, but chelicerae as in female. PROTONYMPH : A broadly elliptical mite with relatively short, very robust legs. Dorsum. — Anterior and posterior plates similar in shape and sculpturing to those of female, but covering most of idiosoma. Propodosomal margin bearing four pairs long setae. Peritremes short, entirely dorsal, extending from level of posterior margin of coxae III, terminating posterior to insertion of fourth pair propodosomal setae. Pair of long metapodosomal setae inserted just mesad of stigmata as in adults. Venter. — Sternal plate very lightly sclerotized, roughly diamond-shaped with rounded corners, bearing three pairs well-developed setae. Four pairs well-developed setae be- tween sternal plate and anal plate. Anal plate elongate, narrow with rounded anterior margin; pair of well-developed adanal setae on plate anterior to anal opening; postanal seta minute. Legs. — Setation similar to that of male deutonymph. Gnathosoma. — Similar to that of male deutonymph but palpi relatively short and stout. TYPE MATERIAL: Holotype female (colln. no. T279) collected by T. H. G. Aitken from Desmodus rotundus rotundus at St. Patrick's Estate, Arima Valley, Trinidad, 16 May 1957. Allotype male (host. no. 9182) collected by V. J. Tipton from Desmodus rotundus at Guanico (Los Santos) , 27 January 1962. Four paratypes (1 female, 1 deutonymph and 2 protonymphs) same data as male allotype ; 1 paratype protonymph same data as holotype female ; 4 paratype females from (3) Desmodus rotundus, Chiriqui Province, V. J. Tipton, 6, 7, 11 March 1962; 1 paratype female from Desmodus rotundus murinus, Casa Tilley, Cerro Punta (Chiriqui), C. M. Keenan and V. J. Tip- ton, 6 February 1960; 14 paratypes (10 females, 3 males, 1 deutonymph) from Desmodus r. rotundus at Antilles (Kern Trinidad Oilfields Ltd.), La Brea, Trinidad, T. H. G. Aitken, 14 April 1958. Holotype, allotype, 3 male and 2 female as well as proto- and deutonymph paratypes in the United States National Museum ; 1 male, 2 female para- types, Chicago Natural History Museum ; 2 male, 2 female paratypes, Trini- dad Regional Virus Laboratory ; remaining material in the collection of the author. Many other specimens of this species from Trinidad are in the author's collection, all taken from Desmodus r. rotundus. 142 ECTOPARASITES OF PANAMA Periglischrus caligus Kolenati. Plate 42. Periglischrus caligus Kolenati, 1857, Wien. Ent. Monatschr., 1, (2), p. 60 — Type deposition unknown (Brazil and Surinam, from Glossophaga soricina). Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 196. Periglischrus caligus bears a general resemblance to P. vargasi from which females may be separated by the presence of a strong medio-distal lobe on the palpal tibia ; females differ from all species of the genus in having broadly inflated, scimitar-shaped setae on the postero-ventral margins of legs IV. Males are characterized by their small size, short legs, simple leg setae, small sternal plate setae and position of the dorsal propodosomal setae. As pointed out by Rudnick ( 1960 ) , this species has been known only from Kolenati's (1857) original figures and description, which do not present valid characters for a specific diagnosis. Through the courtesy of Dr. Marc Andre, a female specimen was examined which had been identified as this species by Kolenati and deposited in the Museum National d'Histoire Natu- relle, Paris. This proved to be the same as numerous specimens collected recently from Glossophaga soricina leachii from Panama, and Glossophaga s. soricina from Trinidad. Since the location of the types of Periglischrus caligus is unknown, the female is redescribed and illustrated here and the male is described for the first time. DESCRIPTION, FEMALE (pi. 42, figs. 1, 2) : A large, broadly rounded mite with inflated opisthosoma and relatively short, stout, strongly setose legs. Idiosoma 890-1080 /* long by 640 /j. wide. Dorsum. — Anterior plate approximately as wide as long, widest at level of posterior one-third, with broadly rounded anterior margin, slight shoulders and convex sides; posterior margin truncate, slightly concave and joined to posterior plate by two sub- median lobules; with 11 pairs alveoli arranged as illustrated. Posterior plate bluntly triangular, 98 n long by 146 n wide, bearing seven pairs of alveoli, the posterior pair con- taining microsetae. Propodosomal setae all long, measuring up to 70 /*; ratio of distance between bases of first setal pair to that between first and second pairs ranges from 1.2:1 to 1.4:1 based on five specimens. Unarmed opisthosoma with four small setae. Postanal seta minute, subterminal. Venter. — Sternal plate longer (110 /*) than broad (82 /*), roughly pentagonal with tapering anterior margin terminating in narrowly rounded tip ; bearing two pairs pores ; three pairs small setae about 11 /u, long located off margin of plate; subequal pair meta- sternal setae postero-lateral to plate. Epigynial plate a small, narrowly longitudinal sclerotization between coxae IV, slightly constricted opposite insertion of very small pair of genital setae which are off the plate. Pattern of sclerotized opisthosomal areas, in- cluding anal plate, and opisthosomal setation as described for Periglischrus tiptoni ex- cept that the three pairs of minute anterior setae arranged in diverging rows are re- duced to two pairs of alveoli with no evident setae. Adanal pair of setae 18 /* long. Legs. — Relatively short, stout, strongly setose. Leg I 350-370 ^ long exclusive of ambulacrum. Posterior seta of coxa II 91 fj, long, minutely barbed; other coxal setae all minute, simple and very fine. In addition to usual ventral setation legs I and II with strongly serrated, leaf-like setae on posterior margins ; similar setae on anterior margins of legs III and IV; femur, patella and tibia IV each with large, inflated, scimitar-like posterior seta. Dorsally long setae superficially appear nude but minute barbs present; femur and patella II each with two minute to small basal setae plus two long setae. Gnathosoma. — Distal hypostomal setae about one-third longer than short setiform gnathosomal pair; inner hypostomal setae absent, outer pair vestigial with only alveoli FURMAN : SPINTURNICID MITES 143 visible. Palpal tibia with prominent medio-distal lobe; trochanter with flap-like emar- gination extending from ventral medial sclerotization which is normally adpressed to other mouthparts. Chelicerae normal. MALE (pi. 42, figs. 3, 4) : A relatively small, ovoid mite with short, stout legs radially arranged. Idiosoma 377 /u long by 297 /u. wide. Dorsum. — Shape of dorsal plates and alveolar pattern similar to that of Periglischrus vargasi. Anterior plate 198 n long by 230 /* wide. Posterior plate 104 /j. long by 135 /u. wide. Propodosomal setae all well developed, setiform, measuring up to 38 /* long; ratio distance between bases of first pair to that between bases of first and second pairs ap- proximately 2.8:1. Unarmed opisthosoma with pair of short setae subterminally. Post- anal seta minute, dorso-terminal. Venter. — Sternal plate longer (159 M) than broad (147 M) . general shape as in P. var- gasi but postero-lateral margins concave ; with usual five pairs setae of which the longer ones, anteriorly, measure approximately 24 /j. and reach about halfway to imaginary line drawn between second pair. Pair of minute setae behind sternal plate. Anal plate, accessory posterior platelets and opisthosomal setae between coxae IV as in P. vargasi. Legs. — Short, stout, radially arranged; leg I approximately 380 n long exclusive of ambulacrum. Coxal setae normal, setiform; posterior seta of coxa II longest (50 M), about one and one-half times as long as anterior seta of II ; proximal seta of coxa I shortest of coxal setae (12 /*). Other leg segments ventrally with short, simple setae except for usual long tarsal trichobothria. Dorsal setae simple, setiform, superficially nude, but longer ones with minute barbs; in addition to other setae femora I and II bear two small setae, III and IV each bear one minute and one small seta. Gnathosoma. — Gnathosomal and distal hypostomal setae subequal, acicular, about as long as distance between bases of the hypostomal pair; inner hypostomal setae absent, outer hypostomal pair of setae minute. Pair of hypostomal processes long, slightly sinuous, stylet-like structures with membranous inner borders. Chelicerae normal, with spermatophoral process recurved, approximately 150 /JL long. MATERIAL EXAMINED: Plesiotype female and male (host no. 9840) were collected from Glossophaga soricina at Los Santos Province, by V. J. Tipton, 10 February 1962 ; 2 additional males, same collection as plesiotype. Two females collected in the Canal Zone by C. M. Keenan and V. J. Tipton from Glossophaga soricina leachii at Empire Range, 30 September 1959, and at Coco Solo, 20 October 1959, respectively. In addition several collections identified as this species are recorded here from Trinidad, taken from Glossophaga soricina soricina. The plesiotype male and female are in the United States National Museum; 2 females, Chicago Natural History Mu- seum ; 1 female, Trinidad Regional Virus Laboratory ; remaining specimens in the collection of the author. Periglischrus vargasi Hoffmann Periglischrus vargasi Hoffmann, 1944, Rev. Salub. y Enferm. Trop., Mexico, 5, (2), p. 91 — Institute de Salubridad y Enfermedades Tropicales de Mexico, Mexico, D.F. (Yerbabuena, Guerrero, Mexico from Leptonycteris nivalis yerbabuenae) . Rud- nick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 199, pi. 31. Specimens identified as this species from Panama agree closely with the redescription given by Rudnick (1960). All females seen, including forms from the type host from Mexico, differ from figures given by both Rudnick (loc. cit.) and Hoffmann (1944) in one respect: femur, genu and tibia IV each has a relatively long (45 ^), stout, postero-ventral seta with an attenu- ated, recurved tip ; these setae differ from comparable setae in Periglischrus caligus in that they are not inflated. 144 ECTOPARASITES OF PANAMA The following characters suffice to diagnose Periglischrus vargasi fe- males : Large, stout mites approximately 1 mm. long of typical appearance for the genus. Palpal tibia lacking medio-distal lobe. All dorsal propodo- somal setae well developed, the longest from 70-80 /* long. Ratio of distance between bases of first pair of dorsal propodosomal setae to that between bases of first and second pairs ranges from 1.1-1.5 :1. Six pairs medium to large dorsal opisthosomal setae. Sternal plate longer than wide, broadly jug-shaped. Posterior seta of coxa III small. Two minute dorsal setae on femur II. The male of P. vargasi resembles closely that of Periglischrus iheringi from which it differs in possessing two minute dorsal setae on femur II and in the relatively small size of the posterior seta of coxa III, which is only 1.7 times as large as the anterior seta. The spermatophoral process, approximately 110 n long, is considerably shorter than those measured for P. iheringi. MATERIAL EXAMINED i From Trachops cirrhosus taken at Los Santos Prov- ince, 1 female collected by V. J. Tipton, 14 February 1962. From Anoura geoffroyi, Cerro Punta (Chiriqui), C. M. Keenan and V. J. Tipton, 1 male, 1 February 1960, and 1 female, 1 male and a protonymph, 3 February 1960 ; from 2 specimens, the same host, Cerro Hoya (Los Santos) , V. J. Tipton, 11 February 1960, 7 females, 2 males and 1 protonymph. From Anoura cul- trata, Chiriqui, 1 male, V. J. Tipton, 12 March 1962 ; Rio Changena Camp, 1 male and 1 female, V. J. Tipton, 27 September 1961. Previously recorded collections of Periglischrus vargasi have been made from Texas on the north to Guatemala on the south. In addition to the Pana- manian collections recorded here, numerous specimens have been taken from Trinidad bats, to be reported in detail in a subsequent paper, and several specimens have been taken from Venezuela. For purposes of record the latter are included here. From (2) Anoura cultrata (?), Rancho Grande, Venezuela, 2 females, 2 males and 1 protonymph, C. 0. Handley, 30 March 1960 ; from Anoura caudifera at the same locality, 1 female, C. O. Handley, 20 March 1960. Periglischrus tiptoni, new species. Plates 43, 44. DIAGNOSIS : The female is distinguishable from other species of the genus by the following combination of characters: broadly jug-shaped sternal plate; five well-developed, although small, dorsal propodosomal setae, the first pair of which are very widely spaced ; palpal tibia with a pronounced apical, median lobe ; leg II with only one minute dorsal seta on femur in addi- tion to large setae. It is closely related to Periglischrus micronycteridis n. sp., from which it is distinguished by the shape of the sternal plate, longer anterior legs and well-developed posterior seta on coxa III, as well as by host association. Males are characterized by coarsely barbed dorsal seta on tarsi III and IV and by a pair of subapical, flattened, peg-like setae ventrally on tarsi III and IV. They are similar to males of P. inflatiseta but lack blunt, fusiform setae ventrally on legs I and II, and no ventral setae between coxae IV are inflated. DESCRIPTION; FEMALE (pi. 43, figs. 1, 2) : A robust typical member of the genus with FURMAN : SPINTURNICID MITES 145 idiosoma broadly rounded anteriorly and opisthosoma broadly flared in fan shape. Idio- soma 1080 ^ long by 810 p wide. An occasional unfed, teneral female has a much reduced, unexpanded opisthosoma. Dorsum. — Overall outline of two dorsal plates ovate with anterior three-fourths con- sisting of the broad anterior plate; anterior plate 324 /u long by 275 /* wide; plates con- nected by two submedian lobes. Nine pairs alveoli on anterior plate, eight pairs on posterior plate ; pair of minute setae arising from penultimate alveoli of posterior plate. Five pairs of propodosomal setae all well developed although relatively short, the longest about 45 fj. long; insertions of first pair very close to those of second pair: ratio of dis- tance between bases of first pair to that between first and second pair varies from 4.4:1 to 7:1. Peritremes extend just anterior to level of second pair propodosomal setae. Un- armed opisthosoma with four pairs of small setae. Posterior tip of anal plate extending dorsally and bearing minute postanal seta. Venter. — Sternal plate longer (159 //,) than wide (120 /*) , broadly jug-shaped with short, narrow, anterior neck and broad posterior base ; with two pairs pores ; three pairs short (20 /j.) sternal setae just off margins of plate. Pair of metasternal setae subequal to sternals. Epigynial platelet a small longitudinal structure partially divided on antero- median line, bearing two minute setae subterminally on slightly inflated posterior lobe of platelet. Pattern of sclerotized opisthosomal areas similar to those of Periglischrus iheringi. Opisthosomal setae small : three pairs minute setae arranged in diverging rows behind epigynial platelet, followed by a pair anterolateral and two pairs lateral to postero- median, ventro-anal sclerotization, two pairs on anterior part of the latter plate and two pairs lateral to anal plate proper; anal plate proper, ventroterminal, narrowly elongate, and apparently tenuously connected to more anteriorly located, posteromedian sclerotiza- tion; pair of short adanal setae terminal on opisthosoma. Adanal setae and two pairs setae lateral to anal plate with canal-like structures1 extending anteriorly from alveoli. Legs. — Well developed, stout; leg I 512 /* long exclusive of ambulacrum. Coxal setae, with exception of long, minutely barbed, posterior seta of II, acicular, nude; posterior seta of III relatively large. Ventral setation of other segments characteristic of species; posterior margins of legs I, II and IV and anterior margins of legs III and IV with long, minutely barbed setae, none markedly inflated ; tibia and tarsus of leg I and patella and tibia of II each with posterodistal, inflated, recurved seta superficially appearing as a blunt, ragged cone. Dorsally most long setae inserted near distal margin of segments, minutely barbed; femoral setae normal with only anterobasal one on femur II minute and two on femur III small. Gnathosoma. — Inner and outer hypostomal setae absent; distal hypostomal pair slightly longer than very small pair of gnathosomal setae. Palpal tibia with pronounced medio-distal lobe; trochanter with blunt, ventral, flap-like emargination distally. Cheli- cerae normal for genus. MALE (pi. 43, figs. 3, 4) : Idiosoma ovoid, widest at level of coxae II-III; 530 /* long by 405 /j. wide. Opisthosoma rudimentary. Legs long. Dorsum. — Shape of dorsal plates similar to those of Periglischrus vargasi; anterior plate as long on mid-line as broad, 351 /*; posterior plate 162 //, long by 222 ^ wide; anterior plate with 12 pairs of alveoli; posterior plate with nine pairs alveoli; microsetae visible in two pairs alveoli of posterior plate and possibly present in some of those on anterior plate. Propodosomal setae relatively short, measuring 39 /* long or less ; ratio of distance between first pair of setal bases to that between first and second pairs approximately 3.5:1. Unarmed opisthosoma bearing one pair small posterior setae in addition to sub- equal postanal seta. Venter. — Sternal plate longer than wide, modified cordate with pronounced medial anterior projection and prominent shoulders; with five pairs long, strong setae and two pairs submedian pores; two additional pairs of lateral marginal pits. Pair of minute setae behind sternal plate. Six pairs well-developed setae in addition to slightly smaller adanal pair in space between coxae IV; three pairs bordering anal plate (two pairs of these may occur on plate), one pair between anal plate and pair of small adanal plate- lets, two pairs on anterior half of anal plate; adanal setae border anterior margin anal 146 ECTOPARASITES OF PANAMA opening. Anal plate elongate, broadly rounded anteriorly, broadest at mid-level and with concave posterolateral margins, ending bluntly at anterior margin of anal opening. Legs. — Leg I of allotype 600 fj, long exclusive of ambulacrum, up to 670 /j. in some paratypes. Coxae with relatively long setae; posterior seta of coxa II 81 p long, less than one and one-half times as long as posterior seta of coxa III. Ventral leg setae tend to be flattened and tooth-like; tarsus I and tibia and tarsus II each with a broad, serrated, blunt seta; tarsi III and IV each with pair of subapical flattened peg-like setae. Dorsal setae strong, with longer setae coarsely barbed and several of shorter setae serrate ; one minute seta on femur II, III and IV; tarsi III and IV with strong, coarsely barbed setae. Gnathosoma. — Gnathosomal and distal hypostomal setae subequal, 27 /u, long; outer hypostomal setae minute; inner hypostomals absent. Palpal tibia without medio-distal lobe; femur with prominent serrated lateral seta. Chelicerae normal for genus, each with very elongate (over 170 n) , recurved, tubular, spermatophoral process terminating in two minute fimbriae. FEMALE DEUTONYMPH (pi. 44, figs. 1, 2) : Very similar to adult male, from which it differs as follows : Idiosoma 650 fj. long. Unarmed dorsal opisthosoma with five pairs small (approximately 10 /u, long) marginal setae in addition to small postanal seta. Sternal plate a rounded diamond shape, longer than wide, bearing three pairs well- developed setae and two pairs pores; pair of setae subequal to sternals located off pos- terolateral margins of plate; slightly smaller pair just posterior to plate, followed by pair of very small setae. Eleven additional pairs well-developed setae on unarmed ventral integument between coxae IV. Anal plate terminal, pear-shaped with broad end pos- terior ; adanal setae well developed, anterior to anal opening which is on posterior tip of body. Chelicerae as in female. Tarsi of legs I lack the two long, blunt-tipped sensory setae observed in adult males. MALE DEUTONYMPH (pi. 44, figs. 3, 4) : Similar to female deutonymph, from which it differs in having only one pair small setae on the unarmed margin of dorsal posterior opisthosoma, and in posssessing only six pairs of setae on the unarmed ventral integu- ment between coxae IV. PROTONYMPH (pi. 44, figs. 5, 6) : Similar in general appearance, size and body shape to deutonymphs, from which it is immediately distinguishable by short dorsal peritremes extending from level between coxae III-IV to just beyond posterior margin of coxa II. Propodosoma possessing only four pairs relatively short marginal setae, ranging from 24-30 n long. In other respects dorsum as in male deutonymph. Venter as in male deutonymph from which it differs as follows : Lacking pair of well- developed setae posterolateral to sternal plate and pair just posterior to plate; small pair of setae behind sternal plate somewhat larger than in deutonymph and located in region between coxae IV; total of four pairs setae arising from unarmed cuticula in area between coxae IV. Gnathosoma and legs as in deutonymphs. TYPE MATERIAL: Holotyps female and allotype male (host no. 43065) collected by V. J. Tipton from Phyllostomus hastatus panamensis at Panama City (Panama), June 1961. Paratype series, all, unless noted, collected by C. M. Keenan and V. J. Tipton : female, female deutonymph and male from Phyllostomus h. panamensis at Chepo Road (Panama), 8 October 1959; male deutonymph, same host, Fort Kobbe (Canal Zone), 9 October 1959; female, same host, Chilibrillo Caves (Panama), 28 October 1959; 3 females, same collection data as preceding, but 17 July 1959; 1 female, same host, Chilibrillo River (Panama), 27 August 1957; 8 females and 1 male same host, Fort Sherman (Canal Zone), 30 July 1959; 1 male, same host, Bocas del Toro, 22 January 1960 ; 4 males, 1 female, 2 deutonymphs and 3 proto- nymphs, from Phyllostomus h. hastatus, Heights of Guanapo, Trinidad, T. H. G. Aitken, 11 July 1957 ; 2 females, 1 male, 1 deutonymph and 2 proto- nymphs, same host and locality, W. G. Downs, 7 November 1956. FURMAN : SPINTURNICID MITES 147 Holotype, allotype, 3 male, 2 female, 1 protonymph, 1 male and 1 female deutonymph paratypes in the United States National Museum ; 2 paratypes (male and female) each in Chicago Natural History Museum and Trinidad Regional Virus Laboratory; remaining material in the collection of the author. ADDITIONAL MATERIAL EXAMINED : Several collections from Phyllostomus d. discolor taken in Trinidad. From Trachops cirrhosus taken in Panama : Fort Sherman (Canal Zone), 1 female, C. M. Keenan and V. J. Tipton, 23 November 1959; Los Santos, V. J. Tipton, 2 females, 1 male, on 10 February 1962, 1 male from collection of 2 bats on 11 February 1962, 4 females, 2 males and 4 protonymphs from 7 bats on 14 February 1962, 6 fe- males, 7 males, 5 male deutonymphs, 3 protonymphs from 4 bats on 21 Feb- ruary 1962. A single collection of 1 female from Trachops cirrhosus in Trinidad. A single collection of 1 female and a protonymph from Myotis chiloensis, Chiriqui Province, V. J. Tipton, 6 March 1962. Two collections of this mite have been seen from Colombia, from Phyllostomus hastatus (CNHM no. 88066) taken at La Macarena, Rio Guapaya by Kjell von Snei- dern, 14 March 1957, and from Phyllostomus elongatus (CNHM no. 88063) at Los Micos, San Juan de Arama, 21 February 1957, by the same collector. REMARKS: Periglischrus tiptoni is a common parasite of bats of the genera Phyllostomus and Trachops in Panama and Trinidad, where it is often found in association with another new species, Periglischrus inflatiseta. It is named in honor of Lt. Col. Vernon J. Tipton, United States Army, who collected many of the spinturnicids recorded in this work. Periglischrus micronycteridis, new species. Plate 45. Periglischrus micronycteridis is closely related to P. tiptoni, both species occurring on genera of Phyllostominae, although never encountered on the same genera. P. micronycteridis is a smaller species with stubby legs; female with the posterior seta of coxa III very small, the dorsal anterobasal seta of femur I very small to minute, and the sternal plate of engorged speci- mens roughly tongued-shaped. Males of P. micronycteridis differ from P. tiptoni in smaller size, short legs and lack of coarsely barbed, dorsal leg setae. DESCRIPTION, FEMALE (pi. 45, figs. 1, 2) : Gravid specimens broadly rounded anteriorly and posteriorly, with opisthosoma expanded; idiosoma approximately 970 /JL long by 756 /a wide. Legs relatively short and stout. Dorsum. — Dorsal plate similar to that of Periglischrus tiptoni, but anterior plate as broad as long (245 /*) ; posterior plate 108 n long by 155 n wide. Propodosomal setae 19-24 p. long, second pair longer than first pair; ratio of distance between bases of first pair to that between first and second pair varies from 4.6:1-7.8:1. Peritremes extend just anterior to level of second pair propodosomal setae. Unarmed opisthosoma with four pairs small setae. Posterior tip of anal plate extends dorsally, with minute post- anal seta. Venter. — Sternal plate longer (126 /x) than wide (87 n) ; in engorged type specimen, plate narrowly tongue-like with eroded margins; three pairs small setae approximately 12 /j. long located just off sclerotized margins of plate; two pairs circular pores on plate; very faint hyaline border extending beyond sclerotized margins includes bases first two pairs setae; in unengorged specimen sternal plate appears uniformly dense throughout area encompassed by hyaline border and sclerotized central area of engorged specimens. Pair metasternal setae subequal to sternals, posterolateral to sternal plate. Epigynial 148 ECTOPARASITES OF PANAMA platelet small, elongate, between coxae IV; preceded anteriorly by small, fan-shaped cuticular pattern; two minute setae lateral to platelet. Pattern of sclerotized opistho- somal areas similar to that of Periglischrus tiptoni on fed specimens; not visible in un- engorged specimen. Opisthosomal setae as in P. tiptoni. Anal plate ventroterminal, elongate, widest posteriorly; adanal setae subequal to nearby ventral setae, 21 n long, subterminal, arising anterior to anal opening which is terminal. Canal-like structures from adanal setae as in P. tiptoni. Legs. — Short, stout; leg I exclusive of caruncles 350 /* long on type, ranging from 340-370 ju long on six representative specimens ; posterior seta coxa II strong, minutely barbed, 110 /u long; others delicate and small; of latter, anterior seta of coxa III usually longest, 24 //; ventral and dorsal setation of other segments similar to that in P. tiptoni, but with somewhat shorter setae and with dorsal, anterobasal seta of femur I very small OM). Gnathosoma. — As described for P. tiptoni but with medial lobe of palpal tibia less pronounced. MALE (pi. 45, figs. 3, 4) : A small mite for the genus, with stubby legs. Idiosoma ovoid, widest at level of coxae II-III; 421 /* long by 335 /* wide. Dorsum. — Shape and alveolar pattern of dorsal plates as in Periglischrus tiptoni; anterior plate 237 /n long on mid-line by 255 /* wide; posterior plate 128 /* long by 160 ^ wide ; plates covering most of idiosoma. Propodosomal setae relatively short, measuring 38 /JL or slightly less in allotype; ratio of distance between first pair setal bases to that between first and second pairs approximately 4.7:1. Unarmed opisthosoma bearing one pair small posterolateral setae in addition to smaller postanal seta. Peritremes normal for genus, extending almost to bases of second pair propodosomal setae. Venter. — Sternal plate similar to that of P. tiptoni but setae relatively shorter, not overlapping bases of more posteriorly located setae of plate. Pair of reduced but not minute setae, 12 /* long, behind sternal plate. Six larger pairs setae in addition to sub- equal adanal pair in space between coxae IV ; anal plate and setal pattern as in P. tiptoni. Legs. — Relatively short; leg I 394 fj. long exclusive of ambulacrum. Posterior seta of coxa II long (110 /*), minutely fimbriated, three times as long as next longest coxal setae, which are acicular. Ventral setae of other segments mostly relatively short and acicular; tarsi I and II each with one and III and IV each with three short, sharp, spini- form setae. Dorsal setae of legs with longer setae very minutely barbed, superficially appearing nude; shorter setae nude, or at most some minutely barbed; femora II, III and IV each with one minute seta in addition to larger setae. Gnathosoma. — As described for P. tiptoni except that gnathosomal and distal hypo- stomal setae lengths are 16 ^ and the lateral seta of palpal tibia is nude. FEMALE DEUTONYMPH : Very similar to female deutonymph of P. tiptoni from which it differs as follows: a smaller mite with idiosoma approximately 430 ^ long; legs short, stubby; leg I 430 n long exclusive of ambulacrum; leg setae essentially nude, some with very minute barbs; posterior seta of coxa II relatively long (100 /j.). TYPE MATERIAL : Holotype female (host no. 7959) and 5 paratype females collected by R. L. Wenzel and C. M. Keenan from Micronycteris megalotis microtis near Borinquen Highway (Canal Zone) , 24 October 1961. Allotype male (host no. 10010) with 2 male, 1 female and 1 female deutonymph para- types collected by V. J. Tipton from Micronycteris minuta, Guanico (Los Santos) , 24 February 1962. Other paratypes from Micronycteris megalotis microtis: 5 females, Borinquen Highway (Canal Zone), 24 October 1961 and 2 females, Cocoli (Canal Zone) , 24 October 1961, R. L. Wenzel and C. M. Keenan ; 4 females, Chiriqui Province, 6 March 1962, V. J. Tipton ; 6 females, same data but 2 March 1962. Paratypes from Trinidad from Micronycteris m. megalotis: 7 females, Cocorite, West Port-of-Spain, 24 June 1958, and 9 females, Quinam Road, Siparia, 13 March 1959, T. H. G. Aitken. FURMAN : SPINTURNICID MITES 149 Holotype, allotype, and 4 female paratypes in the United States National Museum ; 2 female paratypes, Chicago Natural History Museum ; 3 female paratypes, Trinidad Regional Virus Laboratory ; remaining material in the collection of the author. A single male from Micronycteris megalotis microtis, Barro Colorado Island (Canal Zone) , collected by C. M. Keenan and V. J. Tipton, 12 January 1960, is doubtfully identified as P. micronycteridis. It appears abnormal in several minor characteristics. REMARKS : P. micronycteridis specimens from different hosts and of dif- ferent degrees of engorgement exhibit minor differences in morphology which are interpreted as intraspecific variation. Specimens from Trinidad bats have a short fine seta on the anterior margin of coxa III as well as II, while on all other specimens these are represented by longer, delicate setae. The single female associated with males is an unfed specimen with unex- panded idiosoma. On it, the characteristic shape of the sternal plate seen in engorged females appears quite different, with convex lateral margins in- stead of the roughly tongue-shaped structure illustrated here. However, in engorged specimens an almost transparent marginal area of the plate ap- pears to represent the actual margins as seen in the unfed specimen. Periglischrus species A single female specimen designated species "D" represents a possible new species related to Periglischrus desmodi. It was collected by C. M. Keenan and V. J. Tipton from Pteronotus parnellii fuscus, Bocas del Toro Province, 1 February 1960. According to Goodwin and Greenhall (1961), this host is known to roost in caves with Desmodus rotundus, the common host of Periglischrus desmodi. It seems probable that the single specimen of species "D" recorded here may represent a slightly atypical specimen of Periglischrus desmodi which strayed from its normal host. It differs from typical P. desmodi in having shorter dorsal propodosomal setae measuring up to 51 IJL long ; in having shorter dorsal opisthosomal setae, and in having the scimitar-shaped posterior setae of leg IV inflated as in Periglischrus caligus. A collection of two females, three males and three nymphs is designated as an undetermined Periglischrus species "K." It was taken from Loncho- phylla robusta at Chilibrillo Caves (Panama) by C. M. Keenan and V. J. Tipton, 20 August 1959. Species "K" probably represents a new species closely related to P. desmodi, but it is left undescribed here pending collec- tion of additional specimens to settle the question of intraspecific variation. Females differ from P. desmodi in having a more angular sternal plate, shorter dorsal propodosomal setae and inflated, scimitar-like posteroventral setae on leg IV. Males possess only one pair of dorsal opisthosomal setae, shorter sternal plate setae and a small idiosoma about 378 ^ long. A single female designated species "G" was taken from Macrophyllum macrophyllum at natural bridge, Madden Dam (Canal Zone), by C. M. Keenan and V. J. Tipton, 31 July 1959. This undoubtedly will prove to be a new species, but since there is only a single, damaged specimen at hand, its 150 ECTOPARASITES OF PANAMA description awaits further collection. In the key it comes out at couplet 5, but fits neither half of the couplet. Its outstanding characteristic is the possession of an expanded, pilidiform seta on the anterior margin of coxa III, a character possessed by no described species of the genus. Genus Spinturnix von Heyden Spinturnix von Heyden, 1826, Isis (Oken), 18, (6), p. 612. Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2) p. 200. Type-species: Pteroptus myoti Kolenati, 1856, designated by Opinion 128 of the International Commission on Zoological Nomenclature (1936). Spinturnix species differ from all other genera of the family in that the peritremes are short, dorsal over coxae III, with the anterior end bending ventrad, usually reaching the ventral surface between coxae II and III. They have a single dorsal plate. The tritosternum may be present or absent. Legs I and claws of female are not unusually enlarged ; caruncles are large. All males seen by the author lack the two long, bluntly-tipped setae of tarsus I characteristic of Periglischrus. The various instars of immature Spinturnix species seen by the author may be determined as follows : The female deutonymph lacks the epigynial plate but has dorsal opisthosomal setation similar to that of the adult female. The male deutonymph resembles the female deutonymph but has dorsal opisthosomal setation similar to that of the adult male. The protonymph has stigmata smaller in diameter than the width of the peritremes, in contrast to subsequent instars, and has fewer ventral setae between the sternal and anal plates than in deutonymphs. Spinturnix americanus (Banks) Pteroptus americanus Banks, 1902, Can. Ent., 34, (7), p. 173, fig. 6 — Museum of Comparative Zoology, Harvard (Type locality a cave in Indiana, from "bat," probably Myotis lucifugus lucifugus). Spinturnix americanus Banks, 1915, Kept. U.S. Dept. Agric., no. 108, p. 72, figs. 137, 138. Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 218, pis. 39, 40. Spinturnix carloshoffmanni Hoffmann, 1944, Ann. Inst. Biol. Univ. Nac. Mexico, 15, (1), p. 185, figs. 1-5 — United States National Museum, Washington (Cerro del Xitle, Tlalpan, Mexico, D.F., from Natalus mexicanus mexicanus). Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 222. New synonymy. DIAGNOSIS : General appearance typical of genus. Tritosternum present, but small. Unarmed opisthosoma of female with 10-25 long dorsal, dorso- terminal and ventroterminal setae, those near posterior body margin longer than others. Legs of both sexes with ventral and ventrolateral setae mostly short ; the pair of proximal dorsal setae of femora I and II are tiny and the proximal dorsal seta on each of femur III and IV is tiny. Males with two pairs of long opisthosomal setae on unarmed cuticula near posterior apex of dorsal shield. PANAMANIAN MATERIAL EXAMINED : From Myotis n. nigricans the follow- ing collections were made by C. M. Keenan and V. J. Tipton. Fort Davis (Canal Zone), 1 female, 7 January 1960; Fort Clayton (Canal Zone), 1 fe- male, 30 January 1960; and 6 females, 13 September 1960; Gamboa (Canal FURMAN : SPINTURNICID MITES 151 Zone), 1 female, 1 female deutonymph and 2 protonymphs, 23 September 1960; Frijoles (Canal Zone), 2 females, 2 males, 28 March 1960; Barro Colorado Island (Canal Zone), 28 females, 20 males, 2 female deutonymphs, 2 male deutonymphs, 2 protonymphs, 12 June 1960; Bocas del Toro Prov- ince, 1 female, 23 January 1960; Juan Mina (Canal Zone), 11 females, 2 males, 1 female deutonymph, 1 male deutonymph, 2 protonymphs, 28 July 1960; cave at Finca Lara (Chiriqui), two collections of 1 male and of 1 female deutonymph, 3 May 1961. From Myotis n. nigricans or Myotis chiloensis at cave, Finca Lara (Chiriqui), 35 male, 2 female deutonymphs, 6 male deutonymphs, 28 protonymphs, C. M. Keenan and V. J. Tipton, 5 May 1961. From Myotis n. nigricans in Chiriqui Province, 1 male and 2 proto- nymphs, V. J. Tipton, 7 March 1962. From Myotis albescens in Bocas del Toro Province, 1 male, C. M. Keenan and V. J. Tipton, 25 February 1960. From Myotis chiloensis, Chiriqui Province, 1 male, 2 male deutonymphs, 6 protonymphs, V. J. Tipton, 7 March 1962. From Myotis simus at Cerro Punta (Chiriqui), 1 male, V. J. Tipton, 3 May 1960; 1 male, C. M. Keenan and V. J. Tipton, 3 May 1961 ; and 1 male, C. M. Keenan and V. J. Tipton, 5 May 1961. REMARKS : Panamanian specimens of Spinturnix americanus show con- siderable variation in characters previously used to differentiate S. ameri- canus and S. carloshoffmanni. The majority, which I designate as popula- tion "B", lacks a long posterolateral seta on tibia III and IV but has such a seta on patella III and IV. Associated with this is the presence of 10-12 long, subterminal and dorsal setae on the opisthosoma. Others, designated as population "A", have a long posterolateral seta on patella and tibia IV and on patella III associated with the presence of 18-24 long, subterminal and dorsal setae on the opisthosoma. Populations of both kinds occur on the same host species and in one collection both kinds were found on the same host specimen. Rudnick (1960) also noted variation in the above characteristics. In view of the demonstrated variability of the criteria used for distinction of S. americanus and S. carloshoffmanni and the presence of such variation in series from a single host the latter species is considered a synonym. Spinturnix subacuminatus, new species. Plate 46. This species belongs to group III of Rudnick (1960), characterized by long lateroventral leg setae and lack of tiny dorsal setae on femora. It is re- lated to Spinturnix acuminatus (C. L. Koch), from which females differ in possessing 28-33 dorsal opisthosomal setae, of which the posterior 6-9 are much the largest, by a broadly pentagonal tritosternum and by relatively well-developed ventral idiosomal setae. Males differ from S. acuminatus in possessing only one pair of dorsal opisthosomal setae, a tritosternum with a straight posterior border and well rounded anterior margin, and ventral idiosomal setae relatively well developed. DESCRIPTION, FEMALE (pi. 46, figs. 1, 2) : Idiosoma ovoid, approximately 1000 n long by 730 n wide. Dorsum. — Dorsal plate ovoid, 570 n long by 405 /u. wide; with 11 pairs dorsal alveoli, 152 ECTOPARASITES OF PANAMA some bearing minute setae ; several irregular pore-like structures near posterior margin. Five pairs moderately long propodosomal setae surrounding dorsal plate anterior to peritremes, and spaced increasingly far apart proceeding postariorly. Peritremes dorsal over coxae III, bending ventrad between coxae II and III. Metapodosomal setae sub- equal to propodosomals, inserted medial to posterior borders of stigmata. Twenty-eight to 33 opisthosomal setae of which subterminal six to nine are much larger ; others sub- equal to propodosomal setae. Venter. — Tritosternum roughly pentagonal, broader than long, with posterior margin broadly V-shaped. Sternal plate broadly jug-shaped, about as broad as long, broadly rounded posteriorly, with lateral margins converging anterior to second pair of setae; anterior margin bluntly rounded; three pairs setae approximately 36 n long, partially imbedded on plate margins ; surface of plate lightly reticulated ; two pairs of pores. Pair metasternal setae on unarmed cuticula posterolaterad of sternal plate, subequal to sternals. Epigynial plate small; broadly rounded subcircular anterior portion lightly sclerotized; posterior lobe narrowly rectangular, with pair genital setae on posterior margin subequal to sternals. Pair small sclerotized bars lateral to epigynial plate. Twenty to 22 small ventral setae between epigynial and anal plates; submedian anterior pair minute. Anal plate small, subterminal, incompletely ovoid, longer than wide, with anterior heavily sclerotized arc bearing pair adanal setae at lateral ends of arc ; postanal seta 18 fj. long, subequal to adanals. Legs. — Lateroventral setae mostly long; other ventral setae mostly medium length with few short setae. Dorsal setae mostly very long. No minute dorsal setae on femora. Posterior seta of coxa II 215 n long, over two and one-half times longer than other coxal setae. Gnathosoma. — Tectum a short rounded lobe. Gnathosomal setae slightly larger than distal hypostomal pair; other hypostomal setae lacking. Palpal tarsus with inner basal, blunt, stout, prominent spine. Chelicerae normal for genus. MALE (pi. 46, figs. 3, 4) : Idiosoma ovoid, 853 /* long by 641 /* wide. Dorsum. — Similar to female with following differences : metapodosomal setae well posterior to stigmata, and only one pair posteriorly placed opisthosomal setae on unarmed integument; dorsal plate 730 /JL long by 459 ^ wide, lacking irregular pore-like structures near posterior margin. Venter. — Tritosternum a lightly sclerotized small platelet with straight posterior margin and broadly rounded anterolateral margins, broader than long. Sternal plate lightly sclerotized, with reticular surface pattern; longer than wide, widest at level of second pair setae, tapering to width of genital opening anteriorly, and to rounded ex- tremity posteriorly, with three pairs small marginal setae. Metasternal setae laterad of posterior tip of sternal plate. Pair small sclerotized, submedian bars posterior to sternal plate. Five pairs small, simple setae between posterior tip of sternal plate and anal plate. Anal plate similar to that of female. Legs. — As in female. Gnathosoma. — As described for female but with spermatophoral process a stout, re- curved tubular structure approximately 96 ^ long, tapered to blunt tip; basal spine of palpal tarsus very blunt and broad throughout. TYPE MATERIAL : Holotype female (host no. 5536) fromRhogeessatumida, taken in Bocas del Toro Province, by C. M. Keenan and V. J. Tipton, 9 Feb- ruary 1960. Allotype male (host no. 3925) from R. tumida, Fort Kobbe Beach (Canal Zone), C. M. Keenan and V. J. Tipton, 27 July 1959. Para- types as follows: 9 females, 6 males, 3 deutonymphs and 2 protonymphs, same data as allotype ; 3 females, 4 males and 1 deutonymph, same data as allotype but collected 16 November 1959. Holotype, allotype, and 2 para- types, male and female, in the United States National Museum ; 2 paratypes, male and female, Chicago Natural History Museum ; remaining material in the collection of the author. FURMAN : SPINTURNICID MITES 153 Spinturnix species A single female specimen from Myotis n. nigricans taken by C. M. Keenan and V. J. Tipton at Building 519, Fort Clayton (Canal Zone) , 22 May 1961. It closely resembles Spinturnix subacuminatus , but is distinguished by a broadly rounded sternal plate which is wider than long, by the apparent ab- sence of a tritosternum, and by other minor differences. It probably repre- sents a new species, but its description awaits collection of further material. Genus Paraspinturnix Rudnick Paraspinturnix Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2), p. 231. Type-species: Paraspinturnix globosus Rudnick, 1960. The description of this monotypical genus as given by Rudnick (1960) fits the specimens recorded from Panama with the exception that the idio- soma is typically spinturnicid-shaped instead of globose. Paraspinturnix globosus Rudnick Paraspinturnix globosus Rudnick, 1960, Univ. Calif. Publ. Ent., 17, (2) , p. 231, pi. 48, figs. 1, 2 — United States National Museum, Washington (Nickajack Cave, Marion County, Tennessee, from Myotis sodalis). Female specimens from Panama identified as this species agree in all but a few minor respects with the description and figures given by Rudnick (1960). The idiosoma has the characteristic Spinturnix shape attributed by Rudnick (loc. cit.) to newly emerged, non-gravid females. No circular areas of heavy sclerotization appear on the shoulders of the dorsal plate. The metapodosomal pair of dorsal setae arise just medial to the stigmata. A single collection of three females is designated as this species from a bat identified as Myotis n. nigricans or Myotis chiloensis, from a cave at Finca Lara (Chiriqui), C. M. Keenan and V. J. Tipton, 5 May 1961. From the same bat were collected numerous specimens of Spinturnix americanus. Abstract Thirteen species in three genera of spinturnicid mites are recorded for the first time from Panama. Descriptions are given of seven new species of Periglischrus recorded from both Panama and Trinidad, and one new species of Spinturnix recorded from Panama. The new species and typs hosts are P. natali from Natalus stramineus mexi- canus, P. elongatus from Pteronotus parnellii fuscus, P. inflatiseta from Phyllostomus hastatus, P. aitkeni from Sturnira lilium lilium, P. desmodi from Desmodus rotundus rotundus, P. tiptoni from Phyllostomus hastatus, P. micronycteridis from Micronycteris megalotis, and Spinturnix subacuminatus from Rhogeessa tumida. Periglischrus caligus Kolenati is redescribed. Spinturnix carloshoffmanni Hoffman is synonymized under Spinturnix americanus (Banks). The spinturnicid genus Periglischrus occurs primarily on bats of the family Phyl- lostomidae. Periglischrus species are not uncommon on members of the related family Desmodidae, and one atypical species occurs only on bats of the family Natalidae of a different superfamily from the other bat hosts. With few exceptions, the primary hosts of a given Periglischrus species are limited to members of a single bat genus or to mem- bers of closely related genera. One exceptional species, P. iheringi, occurs on seven genera of Stenoderminae of the Phyllostomidae as well as on a genus of Desmodidae. Spinturnix species and Paraspinturnix are primarily limited to bats of the family Ves- pertilionidae. The primary hosts of each species of these two genera recorded from Panama are limited in each instance to host species of a single genus. 154 ECTOPARASITES OF PANAMA HOST-PARASITE LIST Order CHIROPTERA Superfamily Emballonuroidea Family Noctilionidae Noctilio leporinus Periglischrus aitkeni n. sp. Superfamily Phyllostomoidea Family Phyllostomidae Subfamily Chilonycterinae Pteronotus parnellii Periglischrus species "D" elongatus n. sp. Pteronotus suapurensis Periglischrus elongatus n. sp. Subfamily Phyllostominae Micronycteris megalotis Periglischrus micronycteridis n. sp. Micronycteris minuta Periglischrus micronycteridis n. sp. Macrophyllum macrophyllum Periglischrus species "G" Phyllostomus hastatus Periglischrus tiptoni n. sp. inftatiseta n. sp. Trachops cirrhosus Periglischrus tiptoni n. sp. vargasi Hoffmann Subfamily Glossophaginae Glossophaga soricina Periglischrus caligus Kolenati Lonchophylla robusta Periglischrus species "K" Anoura cultrata Periglischrus vargasi Hoffmann Anoura geoffroyi Periglischrus vargasi Hoffmann Subfamily Carolliinae Carollia perspicillata Periglischrus sp. Subfamily Sturnirinae Sturnira lilium Periglischrus aitkeni n. sp. Sturnira ludovici Periglischrus aitkeni n. sp. Sturnira sp. Periglischrus aitkeni n. sp. Subfamily Stenoderminae Uroderma bilobatum Periglischrus iheringi Oudemans Vampyrops helleri Periglischrus iheringi Oudemans Vampyrops vittatus Periglischrus iheringi Oudemans Vampyrodes caraccioli Periglischrus iheringi Oudemans Vampyressa pusilla Periglischrus iheringi Oudemans Chiroderma salvini Periglischrus iheringi Oudemans Artibeus cinereus Periglischrus iheringi Oudemans Artibeus jamaicensis Periglischrus iheringi Oudemans Artibeus lituratus Periglischrus iheringi Oudemans Enchisthenes hartii Periglischrus iheringi Oudemans Family Desmodidae Desmodus rotundus Periglischrus iheringi Oudemans desmodi n. sp. Superfamily Vespertilionoidea Family Natalidae Natalus stramineus Periglischrus natali n. sp. Family Vespertilionidae Subfamily Vespertilioninae Myotis albescens Spinturnix americanus (Banks) Myotis chiloensis Periglischrus tiptoni n. sp. Spinturnix americanus (Banks) Myotis nigricans Spinturnix americanus (Banks) sp. Myotis nigricans or Myotis chiloensis Spinturnix americanus (Banks) Paraspinturnix globosus Rudnick Myotis simus Spinturnix americanus (Banks) Rhogeessa tumida Spinturnix subacuminatus n. sp. Family Molossidae Tadarida brasiliensis Spinturnix sp. References BANKS, N. 1902. New genera and species of Acariens. Can. Ent., 34, (7), pp. 171-176, fig. 6. 1915. The Acarina or mites, a review of the group for the use of economic entomol- ogists. Kept. U. S. Dept. of Agric., no. 108. 153 pp., 294 figs. GOODWIN, G. G., AND GREENHALL, A. M. 1961. A review of the bats of Trinidad and Tobago. Bull. Amer. Mus. Nat. Hist., 122, (3), pp. 187-302, pis. 7-46. HEYDEN, C. H. G. VON 1826. Versuch einer systematischen Eintheilung der Acariden. Isis (Oken), 18, (6), pp. 608-613. HOFFMANN, A. M. 1944a. Periglischrus vargasi n. sp. (Acarina: Parasitidae). Rev. Inst. Salub. Enferm. Trop., Mexico, 5, (2), pp. 91-96, 2 text figs. 1944b. Un nuevo acaro parasite de murcielagos. An. Inst. Biol., Univ. Nac. Mexico, 15, (1), pp. 185-189, 5 text figs. KOLENATI, F. A. 1857. Synopsis prodroma der Flughaut-Milben (Pteroptida) der Fledermause. Wien. Ent. Monatschr., 1, (2), pp. 59-61. OUDEMANS, A. C. 1902. Acarologische aanteekeningen. Ent. Ber., 1, (6), pp. 36-39. 1903. Notes on Acari. Fifth series. Tijdschr. Ent., 45: 123-150, pis. 10-12. RUDNICK, A. 1960. A revision of the mites of the family Spinturnicidae (Acarina). Univ. Calif. Publ. Ent., 17, (2), pp. 157-284, pis. 18-48. TlBBETTS, T. 1957. Description of a new Periglischrus from a bat, Mormoops megalophylla seni- cula Rehn, together with a key to the species of Periglischrus (Acarina, Spinturni- cidae). Jour. Kansas Ent. Soc., 30, (1), pp. 13-19. 155 PLATE 37. ECTOPARASITES OF PANAMA FURMAN : SPINTURNICID MITES PLATE 38. U5 c S 3 CN PLATE 39. ECTOPARASITES OF PANAMA CN •C e FURMAN : SPINTURNICID MITES PLATE 40. -a C OS h I PLATE 41. ECTOPARASITES OF PANAMA CN ID •a FURMAN : SPINTURNICID MITES PLATE 42. I •s PLATE 43. ECTOPARASITES OF PANAMA — c 08 I •C C FURMAN : SPINTURNICID MITES PLATE 44. PLATE 45. ECTOPARASITES OF PANAMA CO •a rt ti I I — a h I s I .«» £ FURMAN : SPINTURNICID MITES PLATE 46. '3 -c CN -s I 166 ECTOPARASITES OF PANAMA Addendum Since submission of the manuscript for this paper in 1962, a series of spinturnicids have been described by Machado-Allison. In 1965 (Acta Biologica Venezuelica, 4, (10), pp. 243-258) he described a new genus and species, Cameronieta thomasi from Chil- onycteris rubiginosa fusca. His figure and description of the female agree with the speci- mens I have described as heteromorphic females of Periglischrus elongatus. The male he describes is indistinguishable from typical males of P. elongatus. His female deutonymph appears to be the typical adult female of P. elongatus. His male deutonymph appears to be an adult male, and his protonymph, probably a male. I conclude that Cameronieta is a synonym of Periglischrus, and C. thomasi becomes Periglischrus thomasi (Machado- Allison). This has close relationship with P. elongatus Furman and P. strandtmanni Tibbets. If further research demonstrates that my hypothesis for existence of heter- omorphic females in P. elongatus is correct, this species will become a synonym of P. thomasi. In 1965 Machado-Allison (Acta Biologica Venez. 4, (11), pp. 259-288) refers to his pa- per in press describing 4 new species of Periglischrus. In his Acta Biologica paper (loc. cit.) he keys out the 4 species of Periglischrus referred to in his unpublished paper and includes rather inadequate photomicrographs. From these I conclude P. tiptoni Furman is a synonym of P. acutisternus Machado-Allison. P. aitkeni Furman is a synonym of P. ojastii Machado-Allison; the latter has erroneously described and photographed a teneral adult female of this species as a female deutonymph. P. micronycteridis Furman may be a synonym of P. parvus Machado-Allison. Both occur on Micronycteris species, but the photomicrographs and key characters given for P. parvus are inadequate for certain identification. I consider P. setosus Machado-Allison to be a synonym of P. caligus Kolenati. P. inflatiseta Furman is a synonym of P. torrealbai Machado-Allison. P. squamosus Machado-Allison is a synonym of P. vargasi Hoffman. P. desmodi Furman is a synonym of P. herrerai Machado-Allison. The Ticks of Panama (Acarina: Ixodoidea) GRAHAM B. FAiRCHiLD,1 GLEN M. KOHLS,2 AND VERNON J. TIPTON 3 Information on the ticks that occur in Panama is scattered and scanty. Most of the information hitherto available is due to the efforts of L. H. Dunn, who worked on the Isthmus for about twenty-five years. Beginning in 1915, he published a series of papers on the life histories, disease transmission potential, hosts and distribution of Panamanian ticks. Dunn's papers are listed in the bibliography. In 1941, Ernesto Osorno-Mesa published an exten- sive paper, with keys to genera and species, on the ticks of Colombia. In this are included a number of early records from Panama when this country was part of Colombia as well as later records assembled from the literature. Fairchild (1943) briefly summarized the records of ticks in a list which formed part of a general summary of the biting arthropods of Panama. Determinations in Osorno-Mesa's paper were largely checked by Joseph Bequaert ; those in Fairchild's in part by R. A. Cooley. The Argasidae of Panama were reviewed by Cooley and Kohls (1944) in their monograph of the species occurring in North America, Central America and Cuba. The present work is based on fairly extensive collections made in recent years by personnel of the Environmental Health Branch, Division of Pre- ventive Medicine, Office of the Surgeon, United States Army Caribbean, especially by Charles M. Keenan, under the successive commands of Major Gordon Field, Major Robert M. Altman and Major Vernon J. Tipton; of collections made by Conrad E. Yunker and James M. Brennan of the Rocky Mountain Laboratory (RML) attached to the Middle America Research 1 Gorgas Memorial Laboratory, Panama, Panama. 2 United States Department of Health, Education, and Welfare, Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Rocky Mountain Laboratory, Hamilton, Montana. 3 Lieutenant Colonel, Medical Service Corps, United States Army. The authors are listed in alphabetical order. G. B. Fairchild and V. J. Tipton assembled the material. G. M. Kohls is responsible for the taxonomy. 167 168 ECTOPARASITES OF PANAMA Unit, United States Public Health Service, in connection with their studies of Panamanian Trombiculidae ; and of collections made by personnel of the Gorgas Memorial Laboratory (GML), especially Lawrence H. Dunn, Pedro Galindo V., Eustorgio Mendez and G. B. Fairchild. These collections have been largely from wild animals trapped or shot in connection with other studies, or from animals brought in alive by country people for possible sale as pets or experimental animals. In some cases, engorged nymphs or larvae allowed to detach from hosts have been held alive until they molted to the next stage, and eggs and young larvae have been secured in a few cases by holding engorged females in the laboratory, but in general our records are based on adult ticks. A total of forty-seven species are known to occur in Panama, twelve of which are here recorded for the first time for this country. Three species, Ixodes fuscipes Koch, /. minor Neumann and Amblyomma americanum (Linnaeus) , previously reported for Panama, are not included. Nuttall and Warburton (1911) recorded and figured as /. fuscipes a female from Panama from Felis pardalis. Cooley and Kohls (1945) suggested that this speci- men may be /. boliviensis Neumann, but since the palpi and hypostome are missing, certainty is impossible. /. fuscipes is recorded from Brazil from Dasyprocta aguti and Cuniculus paca and the Rocky Mountain Laboratory has a female of this species from "agouti" from Puno (Sandia) , Peru. The female reported by Fairchild (1943) as /. minor, from Peromyscus nudipes or Oryzomys devius, Chiriqui Province, is in fact an Ixodes nymph which is not further determinable. As for A. americanum, Dunn (1923) reported that specimens had been taken by Dr. S. T. Darling on dogs and domestic hogs on San Miguel Island, one of the Pearl Islands in Panama Bay. We have not seen these specimens and have not made collections on San Miguel. No other specimens of this species have been reported from Panama and we seriously doubt its occurrence there. It appears to be restricted to parts of the United States and Mexico. Much remains to be learned about the ticks of Panama, and it seems advisable to point out here some of the more obvious lacunae in our knowl- edge. Many species appear to utilize several hosts at different stages, and the larvae and/or nymphs of many of them remain unknown or at least undescribed. The species of Amblyomma infesting, as adults, the sloths and anteaters, are a case in point. The larvae and nymphs of these species are undescribed and their hosts are unknown, since pre-adult stages are seldom taken on the adult's hosts. Life history studies of only a few of the commoner species have been made and these have been mostly based on laboratory rearings rather than field studies. The limiting factors of temperature, humidity, etc. affecting ticks in this area remain to be studied. The lack of coincidence between the ranges of the ticks and those of their preferred hosts is in need of investigation. Although little detailed information has been collected concerning the ef- fect of environmental factors on Panamanian ticks, certain generalizations may be made. In table 4 (which excludes Argasidae, Amblyomma crassum and A. pictum), we have tabulated the occurrence of various species accord- FAIRCHILD, KOHLS AND TIPTON : TICKS 169 ing to altitude (below 1000 feet, from 1000 to 5000 feet, and above 5000 feet) and according to climate (either wet or dry) . As can be seen, only a few species occur in all three altitudinal zones. In some cases, the range of favored hosts may limit tick distribution, but in others, the range of the hosts is known to greatly exceed that of the ticks. TABLE 4. DISTRIBUTION OF PANAMANIAN TICKS IN RELATION TO ALTITUDE AND CLIMATE (See text for detailed explanation.) Approximate elevation (in feet) Climatic conditions over 5000 1000-5000 under 1000 wet dry Dermacentor halli 4- 4. imitans + 4- 4- + latus 4- 4- 4- Anocentor nitens -j_ _j_ _j_ Boophilus microplus -(-once -f- + Amblyomma auricularium 4-4-4- " cajennense 4-once 4-4-4-4- calcaratum + + + + coelebs + + + dissimile -(-rare 4-4-4- geayi 4- 4- + " longirostre 4-4-4-4- naponense -f- 4- 4- nodosum 4-4-4- oblong oguttatum + 4-4-4- ovale 4-4-4-4- pacae -f twice 4-4-4-4- parvum 4- 4- pecarium 4-4-4- sabanerae 4- rare 4- 4- 4- tapirellum 4-4-4-4" " varium 4-4-4- Haemaphysalis juxtakochi + + + + " leporispalustris 4-4-4-4-4- Rhipicephalus sanguineus 4- 4- 4- 4- 4- Ixodes affinis 4- 4- 4- + boliviensis 4-4- 4- brunneus 4- 4- " lasallei 4-4-4- " loricatus 4- 4- luciae 4-4-4- " pomerantzi 4- 4- rubidus 4- 4- tapirus 4- 4- " tiptoni 4- 4- " venezuelensis 4- 4- In determining the effects of climate, it is not always possible to separate the effects of temperature and humidity. In Panama, for instance, heavier and more continuous rainfall usually occurs at higher altitudes. However, it may be generally assumed that the principal factor in limiting tick dis- tribution in areas of high elevation will be temperature, and in areas of low or intermediate elevation, humidity. 170 ECTOPARASITES OF PANAMA In general, the species of Dermacentor and Ixodes apparently prefer higher altitudes and areas of heavier rainfall. Of the three species of Dermacentor, all have been taken above 5000 feet, and only one below 1000 feet, in an area of very heavy and continuous rainfall. Among the eleven species of Ixodes, five have been taken only above 5000 feet and only three below 1000 feet, while only one species has been taken in a dry area. In marked contrast are the seventeen species of Amblyomma, of which only two, A. cajennense and A. pacae, have been taken above 5000 feet, one species once, the other twice. Preference for dry or wet areas at low eleva- tions is quite marked in a number of species, but only one, A. coelebs, has not been taken in a dry area. These facts suggest that micro-climatic factors affecting the free-living periods of some ticks may be as important in de- termining their distribution as are suitable hosts. Considerable preliminary work on the disease-transmitting potential of Panamanian ticks was done by Dunn. However, complete epidemiologies have not yet been worked out, even for such obvious diseases as relapsing fever, piroplasmosis, and Rocky Mountain spotted fever. The interrelation- ships of ticks with wild animals and their infections, and their bearing upon general problems of parasitology and the epidemiology of diseases of man and domestic animals, comprise an area of investigation as yet almost totally unexplored. Family Argasidae The Argasidae or soft ticks differ from other ticks in lacking a hard sclerotized dorsal plate, or scutum, in all stages. In nearly all cases the life history includes a six-legged larval stage and several nymphal stages, as well as the adult. In most instances, only the larvae attach themselves strongly to the host ; the nymphs and adults contact the host only for brief feedings, though there are some exceptions to this generalization. The nymphs and adults are thus generally to be found free in the habitat of the hosts, while the larvae are usually found attached to the host. Three genera of Argasidae are known from Panama. The genus Otobius Banks is also included in the subjoined key, as it is possible that it may eventually turn up in the drier parts of the Republic. KEY TO PANAMANIAN GENERA ADULTS AND LARGE NYMPHS; AFTER COOLEY AND KOHLS, 1944 1. With a definite sutural line separating the dorsal and ventral surfaces of the body Argas (persicus) Sutural line absent 2 2. Nymphal integument beset with spines ; hypostome well developed. Integument of adults granular; hypostome vestigial Otobius (megnini) Integument of adults and nymphs essentially alike, mammillated or tuberculated and lacking spines; hypostome of various forms in adults and nymphs but not vestigial 3 3. Hypostome broad at the base, scoop-like. Associated with bats Antricola (mexicanus) Hypostome of various forms but never scoop-like. Associated with various animals including bats Ornithodoros (7 species) FAIRCHILD, KOHLS AND TIPTON : TICKS 171 KEY TO PANAMANIAN GENERA AND SPECIES LARVAE 1. Claws absent, pulvilli greatly enlarged. Parasite of bats Antricola (mexicanus) Claws present, pulvilli not greatly enlarged. Parasites of various hosts including bats 2 2. Eyes present Otobius (megnini) Eyes absent 3 3. Palpal segment 4 as long or longer than other palpal segments ; dorsum with 26-30 pairs of dorsal setae. Parasite of domestic fowl Argas (persicus) Not as above Ornithodoros 4 4. Parasitic on bats 5 Parasitic on hosts other than bats 8 5. Basis capituli with a knob on each side and with a pair of pointed cornua-like ex- tensions ventrally viguerasi Not as above 6 6. Hypostome on a conical base much shorter than the hypostome itself hasei Base of hypostome about as long as the hypostome 7 7. Basal teeth of hypostome crowded and deformed azteci Basal teeth of hypostome not crowded and deformed brodyi 8. Hypostome short, approximately 0.112 to 0.130 mm. long, and bluntly rounded. . .rudis Hypostome long and pointed 9 9. Hypostome extremely long and slender, approximately 0.244 to 0.257 mm. long, 0.038 to 0.045 mm. wide puertoricensis Hypostome approximately 0.165 to 0.177 mm. long, 0.047 to 0.065 mm. wide talaje Genus Argas Latreille, 1795 Argas persicus (Oken) Rhynchoprion persicum Oken, 1818, Isis, 3: 1568, figs. Argas persicus (Oken), Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 92, as Argas miniatus Koch; 1933, Amer. Jour. Trop. Med., 13, (5), p. 482. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 583. Cooley and Kohls, 1944, Amer. Midi. Nat. Monog., no. 1, pp. 17-20, figs. This species, the common fowl tick and the vector of fowl spirochaetosis, is the only species of the genus known from Panama. It is world-wide in distribution and is mainly a parasite of chickens; it rarely attacks man. Dunn (1923) and Fairchild (1943) report it as common throughout Panama, but our only definite record based on specimens is a large lot of nymphs and adults taken from fowl cages in the Panama City market by Fairchild a number of years ago. Genus Ornithodoros Koch, 1844 KEY TO PANAMANIAN SPECIES ADULTS AND LARGE NYMPHS 1. Hypostome pointed. Parasites of bats 2 Hypostome truncated or notched at apex. Parasites of various animals including bats 3 2. Hypostome long, slender ; denticles fine and limited to apical portion. Body ventrally without sclerotized plates or transverse band of columnar mammillae just pos- terior to coxae IV. . . .azteci 172 ECTOPARASITES OF PANAMA Hypostome short, flattened, in shape of an inverted V; denticles not evident when examined in situ. Body ventrally with sclerotized plates and with a transverse band of columnar mammillae just posterior to coxa IV viguerasi 3. Discs large, conspicuous, and occupying much of the dorsal surface 4 Discs small, superficial, inconspicuous, not occupying much of the dorsal surface. . . .5 4. Small species, adults usually less than 4 mm. long. Discs as elevated shining areas. Mammillae only slightly elevated and difficult to distinguish from the discs in the posterior areas near the margin on both the dorsal and ventral surfaces. Para- site of bats hasei Larger species, adults usually over 4 mm. long. Discs as large depressed areas. Mammillae conical, well elevated and readily distinguished from the discs. Para- sites of animals other than bats puertoricensis and talaje 5. Dorsoventral groove present. Legs short, leg IV not extending to posterior margin of body. Parasite of animals other than bats; frequently found in native houses rudis Dorsoventral groove absent. Legs long, leg IV extending beyond posterior margin of body. Parasite of bats brodyi Ornithodoros azteci Matheson Ornithodoros azteci Matheson, 1935, Jour. Parasit., 21: 349-351, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 583. Cooley and Kohls, 1944, Amer. Midi. Nat. Monog., no. 1, pp. 109-112, figs, (with O. anduzei Matheson 1941 as synonym). Ornithodoros anduzei Matheson, 1941, Bol. Ent. Venez., 1: 3-5. According to Matheson (1935), larvae have been taken on the bats Carollia perspicillata azteca and Desmodus rotundus murinus, while nymphs and adults were found in cracks and crevices in a culvert at Summit (Canal Zone), in a cave on Taboga Island, and in the Chilibrillo Caves. Larvae from Noctilio labialis in an old building at Summit, 9 Jan. 1944, K. W. Cooper and W. Kirkland, were possibly this species. Recent collections, all of larvae, have been 6 from Lonchorhina aurita, railroad culvert east of Summit Golf Club, 26 Oct. 1959, V. J. Tipton; 20 from Desmodus rotun- dus, 1.5 miles W. of Santa Clara (Code) , 27 Oct. 1959, V. J. Tipton ; 2 from Peropteryx macrotis, Quebrada Bonita (Colon), 9 Mar. 1962, GML; 2 from bats, cave near Cement Plant (Colon), 15 Feb. 1962, C. M. Keenan. In addition to Panama, this species has been recorded from Cuba, Ja- maica, and Venezuela. The Rocky Mountain Laboratory has several lots from bats and from a cave in Trinidad collected by T. H. G. Aitken of the Trinidad Regional Virus Laboratory. Ornithodoros brodyi Matheson Ornithodoros brodyi Matheson, 1935, Jour. Parasit., 21: 351-352, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6) , p. 583. Cooley and Kohls, 1944, Amer. Midi. Nat. Monog., no. 1, pp. 80-81, figs. Larvae were taken from the short-tailed bat, Carollia perspicillata, and nymphs and adults from crevices in the walls and ceilings of the Chilibrillo Caves (Matheson, 1935). Fairchild (1943) records the species as common in the Chilibrillo Caves. Specimens, not now available, were taken 4 Jan., 18 Feb. 1940. Subsequent collections are as follows : Chilibrillo Caves, near Chilibre (Panama), 27 Aug. 1944, W. W. Middlekauf, 4 adults on walls of cave; same locality, 6 and 18 Jan. 1941, K. W. Cooper and W. Kirkland, FAIRCHILD, KOHLS AND TIPTON : TICKS 173 larvae from Carollia perspicillata; same locality and host, 28 Oct. 1959, V. J. Tipton and C. M. Keenan, larvae ; same locality, 4 Dec. 1943, GML, 2 fe- males on walls of cave; same locality, Dec. 1946, H. Trapido, 1 female; same locality, June 1957, GML, 1 male, 4 females, 1 nymph, on walls of cave; same locality, Nov. 1954, GML, 1 larva, from Desmodus rotundus; Cerro Hoya (Los Santos), 6 Feb. 1962, V. J. Tipton, 1 larva, from Trachops cir- rhosus; same locality and host, 24 Feb. 1962, 1 larva; same locality, 10 Feb. 1962, from Carollia perspicillata, 1 larva; bat cave, Quebrada Bonita (Colon), 25 May 1962, 1 male, 1 female, 5 nymphs; and same locality, 25 July 1962, from ? Rhynchonycteris sp., 1 larva. As far as we know, all collections of O. brodyi from the Chilibrillo Caves came from "Cave B," the middle cave of the complex of three which make up the Chilibrillo Caves. All three open from the sides of a sink hole in limestone about 100 meters west of the present Trans-Isthmian Highway. The fauna of these caves was extensively studied and collected by L. H. Dunn, but references to his findings are scattered in the literature. For- merly the caves could be reached only by hours of travel in dugout canoes up the Chagres and Chilibrillo Rivers, and at that time they contained a large and varied bat population. In recent years their accessibility has resulted in much disturbance, including treatment with smoke bombs in connection with an anti-rabies campaign against all bats. This has resulted in the virtual disappearance of bats from Cave B and no adult ticks have been re- covered on visits to the cave in recent years. The species also occurs in Guatemala as evidenced by a nymph in the Rocky Mountain Laboratory collection from cave wall, Cueva de Lanquin, Lanquin (Alta Vera Paz), 1000 feet elevation, 15 June 1948, R. D. Mitchell and Luis de la Torre, Chicago Natural History Museum Guatemala Zoo- logical Expedition. Ornithodoros hasei (Schulze) Argas hasei Schulze, 1935, Zeitschr. Morph. Okol. Tiere, 30: 34, fig. (May) ; 1941, op. cit., 37: 534, 547 (with O. dunni Matheson as synonym). Ornithodoros dunni Matheson, 1935, Jour. Parasit., 21:347-349, figs. (October). Fair- child, 1943, Amer. Jour. Trop. Med., 23, (6), p. 583. Cooley and Kohls, 1944, Amer. Midi. Nat. Monog. no. 1, pp. 103-105, figs. Schulze's material of hasei consisted of an unspecified number of larvae off the bat Myotis nigricans from "La Gueiira" (misprint for La Guaira?) , Venezuela. Kohls has compared reared larvae of O. dunni with the larva of "Argas" hasei figured by Schulze and concurs with Schulze (1941) that dunni is a synonym. Matheson's original specimens of dunni were reared from larvae taken from the bat Dirias albiv enter ( = Noctilio labialis) in Panama City and Sum- mit (Canal Zone) . Adults, nymphs and larvae have since been taken in some numbers from bat guano in the roof of an old church at Pacora (Panama) , June 21, 22, July 26, 1961. The bats inhabiting the church were Noctilio labialis, and one nymph and several larvae were taken from one of the bats. Several larvae, from which two nymphs later emerged, were secured from 174 ECTOPARASITES OF PANAMA the same species of bat taken at the Navy firing point, Galeta Point (Canal Zone) , 19 Nov. 1959. More recently, three lots of larvae were secured from Noctilio leporinus, five lots from Noctilio labialis, and one each from Uro- derma bilobatum, Vampyrops helleri and Tonatia silvicola, all from the vicinity of Las Palmitas (Los Santos), Jan.-Feb. 1962. Also, nine lots of larvae were collected from Noctilio sp. taken in nets, Gamboa (Canal Zone) , pipeline road, 14 May 1962, by C. Yunker. This species has also been recorded from Brazil (Mara jo Island near Belem), and the Rocky Mountain Laboratory collection contains several lots from Trinidad from bats, principally Noctilio I. leporinus, and their roosts, collected by T. H. G. Aitken. Ornithodoros puertoricensis Fox Ornithodoros puertoricensis Fox, 1947, Jour. Parasit., 33, (3), pp. 253-259; 1951, Jour. Parasit., 37, (1), pp. 85-95. Davis, 1955, Jour. Parasit., 41, (1), pp. 76-79. Fox and Garcia-Moll, 1961, Amer. Jour. Trop. Med. Hyg., 10, (4), pp. 566-573. This species was described by Fox (1947) from larvae and from nymphs and adults reared from larvae collected off domestic rats in Puerto Rico. He suggested the possibility that the species was also present in Panama but that it was being confused with O. talaje. The presence of this species in Panama is confirmed by the finding of a larva on Sylvilagus brasiliensis at Los Santos, 19 February 1962, and by re-examination of a lot labeled 0. talaje consisting of numerous larvae, 10 males, 1 female and 7 nymphs, collected by Dunn in Panama but without further data, and by another lot consisting of 7 larvae labeled O. talaje, from "rat", 27 March, 1931. All specimens in these two lots proved to be O. puertoricensis. The adults and nymphs of puertoricensis and talaje are so similar that we are as yet unable to distinguish the two species with certainty on the basis of these stages alone, although available specimens of puertoricensis are rather consistently smaller. The larvae, however, are quite distinct and are readily distinguished by characters of the hypostome. The hypostome of puertoricensis is much longer and narrower and ranges from approxi- mately 0.244 to 0.257 mm. long by 0.038 to 0.045 mm. wide as compared to 0.165 to 0.177 mm. long by 0.047 to 0.065 mm. wide in talaje larvae reared from adults collected at and near the type locality in Guatemala. The Rocky Mountain Laboratory has several collections of larvae from rodents (Proechimys and Nectomys) in Trinidad (T.H.G. Aitken), and larvae that were reared from adults collected at Ayacucho in Colombia. It is of interest to note that Davis (1955) interbred O. puertoricensis from Puerto Rico and "O. talaje" from Colombia and obtained fertile progeny. Ornithodoros rudis Karsch Ornithodoros rudis Karsch, 1880, Mitt. Munch. Ent. Ver., 4: 141-142. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6) , pp. 582-583. Cooley and Kohls, 1944, Amer. Midi. Nat. Monog. no. 1, pp. 101-103, figs. Ornithodoros talaje, Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), pp. 92-93 (in part). Ornithodoros venezuelensis, Dunn, 1933, Amer. Jour. Trop. Med., 13, (2), p. 203; 1933, Amer. Jour. Trop. Med., 13, (5), pp. 476, 482. FAIRCHILD, KOHLS AND TIPTON : TICKS 175 Dunn (1933) considers this species to be primarily a biter of man, at least in the post-larval stages. He records it as common in houses in the interior of Panama, hiding in cracks in furniture and walls and coming out to bite at night. Larvae were taken once on a chicken, and a few post- larval stages from crevices in chicken coops in the Panama City market. Fairchild collected adults and nymphs from crevices in furniture in a native house near Capira (Panama), many years ago, and there are specimens at the Rocky Mountain Laboratory from native houses at New San Juan, Chagres River, 7 July 1939, W. Trager ; from native house at Donoso (Colon) , March 1947, G. B. Fairchild; and from native houses 40 miles west of Panama City, April 1954, G. B. Fairchild. No recent material has been seen, doubtless because it has not been searched for. It is also likely that extensive spraying of houses with insecticides to control malaria, beginning about 1946, has greatly reduced this domestic species. Dunn believed O. rudis to be the main vector to man of relapsing fever in Panama. In addition to Panama, this species has been recorded from Colombia, Venezuela, Paraguay (Cooley and Kohls, 1944), and from Ecuador (Leon and de Leon, 1947), and the Rocky Mountain Laboratory has several col- lections from rodent nests from several localities in the Lancones District (Piura), Peru, October 1946 (Dr. A. Macchiavello). Ornithodoros talaje (Guerin-Meneville) Argas talaje Guerin-Meneville, 1849, Rev. Mag. Zool., 1 : 342-344, pi. 9. Ornithodoros talaje, Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 92 (in part) ; 1927, Jour. Parasit., 13: 177-182; 1931, Psyche, 38, (4), pp. 170-173; 1933, Amer. Jour. Trop. Med., 13, (5) , pp. 475-483. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6) , p. 582. Cooley and Kohls, 1944, Amer. Midi. Nat. Monog., no. 1, pp. 82-88, figs. 38-39, pi. 8. Fox, 1947, Jour. Parasit., 33: 253-259. ? Ornithodoros dugesi Mazzotti, 1943, Rev. Inst. Salub. Enferm. Trop., 4, (4), pp. 371-374; 1949, ibidem, 10, (3), pp. 277-281. With the recognition that O. puertoricensis occurs in Panama, all previ- ous records of O. talaje in Panama must be regarded as questionable. Future reports must also be so regarded unless verified on the basis of larvae in view of the close similarity of the post-larval stages of the two species. Unfortunately, none of the larvae reported by Dunn (1923, 1927, 1931, 1933) are available for study, and a positive identification as 0. talaje of the adults now remaining in the early and more recent collections cannot be made. Consequently, the following records may apply either to talaje, or to puertoricensis, or to both, although some of the specimens appear to be the former because of their size. Dunn (loc. cit.) recorded larvae from domestic rats( Mus rattus, M. alexandrinus and M. norvegicus) as well as dogs, cats, opossums, monkeys, chickens, and snake. Adults were occasionally taken in houses, once (Dunn 1931) in sufficient abundance to cause annoyance to the inhabitants by their bites. Adults were also taken in small numbers in crevices in chicken coops in the Panama City market. Dunn was of the opinion that the tick was widespread in Panama, the larvae attacking a variety of animals, the adults mainly attacking rats. He reported material from Parita (Herrera) , 176 ECTOPARASITES OF PANAMA Santa Rosa (Colon), Gatun (Canal Zone), Chorrera and San Juan (Pan- ama), and the cities of Panama and Colon. Fairchild has notes on speci- mens, not now available, from a native house at Villa Rosario (Panama) , 27 June 1941, sifted from dirt floor and in cracks in furniture, and a single engorged female from Juan Diaz (Panama), July 1941, taken in a native house. Further specimens collected in houses at Villa Rosario, April 1954, are in the Rocky Mountain Laboratory collection. An adult specimen was taken while biting a man in bed, Panama City, 14 August 1944. Cooley and Kohls (1944) stated that the range of O. talaje extends from Kansas and California to Argentina and noted that in the United States the species has been found only on wild rodent hosts and in associa- tion with them. Specimens taken in nests of "ratas silvestres 4" near Sabinas (Coahuila), northern Mexico, were described by Mazzotti (1943) as a new species, O. dugesi. However, Kohls has been unable to detect any significant differences between the larval and post-larval stages of dugesi and those of talaje from the United States, from native houses in southern Mexico, and from the type area in Guatemala, and believes dugesi may be a synonym. Ornithodoros viguerasi Cooley and Kohls Ornithodoros viguerasi Cooley and Kohls, 1941, Pub. Hlth. Kept., 56:396-399, figs; 1944, Amer. Midi. Nat., Monog. no. 1, pp. 106-109, figs. Our records for this species are based on larvae taken from bats, four from Pteronotus parnellii and one from Pteronotus sp. at Cueva de los Murcielagos, near Penonome (Cocle) , 4 Mar. 1955, A. Quinonez, and 15 Dec. 1961, V. J. Tipton. This species has been previously recorded from Cuba, and the Rocky Mountain Laboratory has several lots of larvae collected by T. H. G. Aitken from bats in Trinidad. Genus Antricola Cooley and Kohls, 1942 Antricola mexicanus Hoffmann Antricola mexicanus Hoffmann, 1959, An. Esc. Nat. Cienc. Biol., 9:97-102, figs. (1958) (Mexico). This species was first taken (27 males, 21 females, 4 nymphs) in De- cember, 1961, from Cueva de los Murcielagos (bat cave) about 1 km. NW. of Penonome (Cocle). Ticks were present in fair numbers and were ac- tively crawling about on the guano in the first chamber in the cave. Bats of several species, predominantly Pteronotus spp. with a few Carollia sp., were present in very large numbers, forming a nearly solid sheet on the ceiling and walls of the cave. A later visit, on 24 Jan. 1962, yielded sev- eral hundred adult ticks of both sexes and nymphs of several sizes as well as 20 larvae from Pteronotus psilotis. A fire had been built in the cave a short time previously and bats were much less numerous. Ticks were 4 Neotoma micropus canesccns according to Mazzotti, 1949. FAIRCHILD, KOHLS AND TIPTON : TICKS 177 more in evidence, crawling actively on walls and floors of small side tunnels and packed into available crevices. The cave was hot and damp, the atmos- phere almost unbreathable with ammonia fumes from the guano. Later, one female and one nymph were taken in a cave at Cerro Punta (Chiriqui) , at an elevation of 5800 feet, and larvae from Myotis nigricans at the same locality. This species was described from a female and a male found on bat guano in Gruta de Juxtlahuaca (Guerrero) , Mexico. The Rocky Mountain Labora- tory has a female and a nymph taken in a bat cave, elevation 6300 feet, at Chocoyos (Chimaltenango), Guatemala, 28 April 1948, by R. L. Wenzel, R. D. Mitchell and L. de la Torre, Chicago Natural History Museum Guate- mala Zoological Expedition. The species resembles A. coprophilus (Mcln- tosh) of the southwestern United States and Mexico, but it can be readily distinguished by the presence in females of tubercles, each bearing a tuft of long barbed hairs, on the posterior and posterolateral margins of the body. Family Ixodidae The Ixodidae or hard ticks are characterized by having a hard sclerotized scutum in all stages. This structure is small in the larval and nymphal stages and in the female, but covers the whole dorsum in the males. All stages attach to their hosts for relatively long periods of feeding. A few species stay on the same host in developing from larva to nymph to adult, but most species, in each stage, drop from the host on completion of feed- ing and after molting may parasitize the same or widely different host species. Seven genera, keyed below, have representatives in Panama. KEY TO PANAMANIAN GENERA MALES AND FEMALES 1. Anal groove distinct and contouring the anus anteriorly. Eyes, ornamentation, and festoons absent. Venter of male more or less completely covered by seven hardened non-salient plates Ixodes Anal groove distinct or indistinct but contouring the anus posteriorly. Eyes, orna- mentation and festoons present or absent. Venter of male with or without salient plates 2 2. Eyes absent, scutum inornate. Palps short and broad, base of second segment projecting laterally beyond the basis capituli. Festoons present. Venter of male without plates Haemaphysalis Eyes present 3 3. Palps usually long and slender, longer than the basis capituli; segment 2 at least one and one-half times longer than segment 3. Scutum usually ornate. Venter of male without extensive salient plates, rarely with small sclerotized non-salient plaques near the festoons. Coxa IV of males not greatly enlarged. . . .Amblyomma Palps short and broad, not longer than the basis capituli ; segment 2 about as long as segment 3 4 4. Basis capituli rectangular. Scutum ornate or inornate. Male without ventral plates. Coxa IV of male much larger than other coxae 5 Basis capituli hexagonal. Scutum inornate. Male ventrally with adanal and acces- sory plates. Coxa IV of male not greatly enlarged 6 5. Scutum ornate. Festoons 11 in number. Denticles of hypostome arranged in three longitudinal rows on each side of the median line. Spiracular plate not or only slightly raised above body surface; goblets numerous and small Dermacentor 178 ECTOPARASITES OF PANAMA Scutum inornate. Festoons seven in number. Denticles of hypostome arranged in four longitudinal rows on each side of the median line. Spiracular plate promi- nently elevated above surface of body Anocentor 6. Festoons absent. Palpi very short and ridged dorsally and laterally. Coxa I with two very short spurs. Anal groove obsolete in female, indistinct in male. Male very small Boophilus Festoons present. Palpi not unusually short, not ridged. Coxa I with two long spurs. Anal groove distinct. Male moderate in size Rhipicephalus Genus Ixodes Latreille, 1795 KEY TO PANAMANIAN SPECIES FEMALES 1. Hypostome with only files 1 and 2 extending the full length, files 3 never extending more than one-third the total length 2 Hypostome with files 3 extending for half the length or more; files 4 may be present 4 2. Medium-sized tick of usual appearance; basis capituli subrectangular, not ex- panded laterally; coxa I with internal spur slender and much longer than the external rubidus Large ticks of unusual appearance : basis capituli expanded laterally ; dorsally with a pair of anteriorly converging ridges ; coxa I with spurs flattened and robust . . 3 3. Coxa I with spurs subequal in length loricatus Coxa I with external spur much longer than the internal luciae 4. Coxa II without spurs 5 Coxa II with an external spur 7 5. Coxae III and IV without spurs. Auriculae mild ridge-like protuberances. Para- site of tapirs tapirus Coxae III and IV each with a short external spur. Auriculae long retrograde processes. Parasites of small mammals 6 6. Auriculae as sharply pointed curved horns. Hypostome situated on a conical base about as long as the hypostome itself. Punctations of scutum rather uniformly distributed venezuelensis Auriculae as long, straight, blunt, retrograde processes. Base of hypostome about half as long as the hypostome. Large punctations of scutum grouped near the posterior margin lasallei 1. Coxa I with a short internal spur, less than twice as long as the external spur. Trochanters I to III with small, but distinct, ventral spurs. Scutum, postscutal areas, and venter with numerous conspicuous long white hairs. Parasites of birds brunneus Coxa I with internal spur much longer than the external spur. Trochanters with- out spurs. Parasite of mammals 8 8. Cornua absent, posterior margin of basis capituli an even, concave, salient edge. Auriculae mild lateral saliences. Punctations near posterior margin of scutum conspicuous, deep circular affinis Cornua present 9 9. Palpal segment 1 with a long, sharp, ventral process. Auriculae long, thin curved horns. Parasite of Sylvilagus pomerantzi Palpal segment 1 with a suboval ventral plate or a short ventral spur similar to the external spurs of the coxae 10 10. Palpal segment 1 with a suboval ventral plate. Auriculae stout, curved, slightly longer than broad. Internal spur of coxa I long, slim, reaching nearly across coxa II in unfed specimens boliviensis Palpal segment 1 with a short ventral spur similar to the external spurs of the coxae. Auriculae mild, slightly pointed elevations. Internal spur of coxa I FAIRCHILD, KOHLS AND TIPTON I TICKS 179 moderate, slightly overlapping coxa II. Parasite of squirrels (Sciurus gran- atensis so far as known) tiptoni MALES (Males of rubidus, tapirus, and venezuelensis unknown) 1. Hypostome with large lateral denticles conspicuous and well differentiated from the small median denticles which are in diagonal or transverse crenulations. . .affinis Hypostome with smaller lateral denticles not well differentiated from the medians which may be in longitudinal files as in females or with small, mild teeth in in- definite files, or in diagonal transverse crenulations 2 2. Median denticles of hypostome in definite lineal files. Large ticks of unusual ap- pearance: basis capituli expanded laterally, dorsally with a pair of anteriorly converging lateral ridges; lateral body folds wide; coxa I with spurs flattened and robust 3 Median denticles in diagonal or transverse crenulations, or faint or indefinite lineal files. Smaller ticks of usual appearance 4 3. External spur of coxa I much longer than internal luciae External and internal spur of coxa I about equal loricatus 4. Hypostome distinctly notched apically 5 Hypostome not notched apically 7 5. Coxae I and II without external spur, posterior margins broadly rounded and salient. Internal spur of coxa I long. Scutum with large punctations limited mainly to lateral and posterior areas lasallei Coxae I and II with external spur 6 6. Trochanters each with a small but distinct ventral spur. Spurs of coxa I short, subequal. Small tick, up to about 2 mm. long exclusive of capitulum. Parasite of birds brunneus Trochanters without ventral spurs. Coxa I with internal spur medium and much longer than external spur. Larger tick, up to at least 2.4 mm. long exclusive of capitulum. Parasite of squirrels (Sciurus granatensis so far as known) . . . .tiptoni 7. Cornua present, short, pointed. Posterointernal margin of coxa IV salient and spur-like. Hypostome slender, longitudinally grooved ventrally, with about 12 diagonal rows of crenulations. Parasite of Sylvilagus pomerantzi Cornua absent. Posterointernal margin of coxa IV smoothly rounded. Hypostome broad, heavy, with about eight transverse rows of crenulations plus a basal row of large irregular teeth which are well elevated above the smooth basal portion of the hypostome boliviensis Ixodes affinis Neumann Ixodes affinis Neumann, 1899, Mem. Soc. Zool. Fr., 12: 120-121. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 583. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 29-34 (with I. ricinus aragaoi Fonseca, 1935 as synonym). Fonseca, 1960, Acarologia, 2, (1), pp. 9-10. Ixodes ricinus aragaoi Fonseca, 1935, Mem. Inst. Butantan, 9: 131-135. Ixodes aragaoi Fonseca. Aragao and Fonseca, 1952, Mem. Inst. Oswaldo Cruz, 50: 727-728. Fonseca, 1960, Acarologia, 2, (1), pp. 9-10. Dunn (1923, 1934) recorded specimens, probably of this species, as I. ricinus, from near Camp Pital and from the Boqueron River area. Fair- child (1943) recorded adults from Mazama and Odocoileus from El Real (Darien) and Alhajuela (Canal Zone), while Cooley and Kohls (1945) re- corded adults from Felis concolor (puma) from Almirante (Bocas del Toro) . We have secured considerable material listed below, all collected by per- sonnel of the Gorgas Memorial Laboratory unless otherwise stated, which indicates a wide distribution in Panama and considerable variety of hosts, 180 ECTOPARASITES OF PANAMA though deer and the larger carnivores seem preferred. Juan Mina (Canal Zone), 4 Sept. and 6 Jan. 1944, from Odocoileus, 4 females; Santa Rosa, Rio Chagres, 24 Feb. 1944, from ocelot, 1 female; Moja Polio, Rio Chagres, 19 Feb. 1941, from dog, 1 female; Gamboa pipeline road (Canal Zone), 16 Oct. 1960, V. J. Tipton, from Felis onca, 1 female; Pina (Colon), 22 Dec. 1957, R. M. Altman, from Didelphis marsupialis, 1 male, 1 female; Tacarcuna Yellow Fever Station (Darien), 3 Sept. 1958, from Mazama americana, 1 female; Rio Teribe (Bocas del Toro), 10 Aug. 1962, from Mazama americana, 3 males, 9 females; Cerro Pirre (Darien), 4 Feb. 1961, C. M. Keenan and C. E. Yunker, from ocelot, 1 female; Rio Chucunaque (Darien), 17 Feb. 1958, from Mazama, 1 male, 1 female; Rio Mandinga (San Bias), 30 May 1957, from Felis onca, 2 males, 1 female; Almirante (Bocas del Toro), 13 March 1961, from Felis pardalis, 2 males, 7 females. The range of /. affinis extends from Georgia (Lund, Marshall and Hayes, 1962) and Florida (Kohls and Rogers, 1953) to Brazil. Cotype females of aragaoi (Brazil) have the punctations near the posterior margin of the scutum somewhat smaller than in affinis but we do not regard this and other minor differences noted by Aragao and Fonseca (1952) and Fonseca (1960) as sufficient to justify recognition of aragaoi as a separate species. Ixodes boliviensis Neumann Ixodes boliviensis Neumann, 1904, Arch. Parasit., 8: 457-458. Kohls, 1956, Proc. Ent. Soc. Wash., 58: 232-233 (with I. bicornis as synonym). Ixodes bicornis Neumann, 1906, Arch. Parasit., 10: 196-197. Fairchild, 1943, Amer. Jour. Trop. Med., 23: 583. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 183-186. Kohls, 1956, Proc. Ent. Soc. Wash., 58: 232-233. Fairchild (1943) and Cooley and Kohls (1945) reported this species as /. bicornis, from man, domestic cat and dog from several localities in Chiriqui Province. Since then we have secured 44 lots from the following hosts : jaguar (3 lots, 3 males, 51 females) ; dog (15 lots, 21 males, 143 females, 1 nymph) ; tapir (3 lots, 12 females) ; Didelphis marsupialis (3 lots, 4 fe- males) ; Procyon lotor (1 lot, 3 males, 9 females, 8 nymphs, 15 larvae) ; cattle (4 lots, 1 male, 18 females) ; Odocoileus (1 lot, 4 females) ; mule (1 lot, 5 females) ; domestic cat (1 lot, 1 female) ; man (4 lots, 1 male, 4 females, only 1 female attached) ; Coendou mexicanus (4 lots, 10 females) ; Nasua nasua (3 lots, 15 females) ; in camp building (1 lot, 1 female). All were collected at elevations of over 2500 feet in Chiriqui Province or in neigh- boring Bocas del Toro Province. /. boliviensis appears to be the principal tick on dogs in this area, and on several occasions large numbers have been taken from a single dog. Its range extends from Mexico to Bolivia. Ixodes brunneus Koch Ixodes brunneus Koch, 1844, Arch. Naturg., 10: 232. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 205-211. Anastos and Smith, 1957, Jour. Parasit., 43: 535-541 (male, nymph, and larva described). Ixodes calif ornicus Banks, 1904, Proc. Cal. Acad. Sci., 3, (3), p. 369. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 215-216. New synonymy. FAIRCHILD, KOHLS AND TIPTON : TICKS 181 A single female from an undetermined species of tinamou, Bambito (Chiriqui), 6 Feb. 1960, collected by V. J. Tipton and C. M. Keenan, ap- pears to be the first record of this bird tick from Panama. The specimen is somewhat larger than those from the United States, the scutal hairs are shorter, fewer and finer, and the median field of the scutum is glossy and has fewer and smaller punctations. This species is known from several States in the United States (exclud- ing Alaska and Hawaii) and from Venezuela. Boero (1945) identified as 7. brunneus a female from the marsupial Dromiciops australis australis and a nymph from a supposed bat's roost, both from Victoria Island in Lake Nahuel (Neuquen), Argentina, but Ringuelet (1947) examined the speci- mens and described them as a new species, Ixodes neuquenensis . Boero's (1957) identification as brunneus of a female from a bird, Turdus nigriceps, Jujuy Province, Argentina, may be correct but his description and figures suggest that it may be another species. Kohls has recently compared the syntype nymphs of 7. calif ornicus with authentic nymphs of 7. brunneus and found them to be the same. Ixodes lasallei Mendez and Ortiz Ixodes lasallei Mendez and Ortiz, 1958, Mem. Soc. Cienc. Nat. LaSalle, 18: 198-202. This species was described from females from Venezuela, taken from Agouti paca. We have secured 10 lots, including the undescribed male, 8 from Dasyprocta punctata and 2 from Agouti paca: Almirante (Bocas del Toro) , 23 Jan. 1960, V. J. Tipton and C. M. Keenan, 1 female ; same locality, 24 Jan. 1962, 1 female; same locality, 18 July 1960, GML, 1 female; Rio Teribe (Bocas del Toro), 10 Aug. 1962, GML, 3 females; Cerro Hoya (Los Santos), 15 Feb. 1962, V. J. Tipton and C. M. Keenan, 7 nymphs, 2 larvae; same locality, 17 Feb. 1962, 1 female ; same locality, 22 Feb. 1962, 2 nymphs ; same locality, 23 Feb. 1962, 1 nymph; Timi de Boa, Rio Teribe (Bocas del Toro), from Agouti paca, GML, May 1962, 1 male, 8 females, and 4 nymphs, and 10 Aug. 1962, 3 nymphs. The salient morphological features of the male of this species are given in the key to Ixodes males. Ixodes loricatus Neumann Ixodes loricatus Neumann, 1899, Mem. Soc. Zool. Fr., 12: 139-142. Nuttall and War- burton, 1911, Ticks, pt. 2, pp. 266-269. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 187-193. Although this species was erroneously reported from Panama as 7. loricatus spinosus by Fairchild (1943), the following seems to be the first authentic record of the species from Panama : Tacarcuna Yellow Fever Sta- tion (Darien) , 1 Sept. 1958, GML, from Metachirus nudicaudatus, 1 female. The range of this species extends from Mexico to Argentina. Its princi- pal hosts appear to be opossums (Family Didelphidae) but it has been re- corded from a variety of other animals. Ixodes didelphidis Fonseca and Aragao, 1952, described from adults 182 ECTOPARASITES OF PANAMA taken on Metachirus sp. and Didelphis Paraguay 'ensis at Anapolis (Goiaz), Brazil, differs from loricatus in having somewhat larger spiracular plates with slightly smaller and more numerous goblets, and probably represents only a local population of that species. There are numerous lots from Anapo- lis in the Rocky Mountain Laboratory collection. Ixodes luciae Senevet Ixodes luciae Senevet, 1940, VI Cong. Intern. Ent., Madrid, (1935), pp. 896-898. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 175-180. Kohls, 1957, Proc. Ent. Soc. Wash., 59, (6), p. 259 (with 7. loricatus vogelsangi Dias, 1954 as synonym). Ixodes loricatus var. spinosus Nuttall, 1910, Parasitology, 3: 411-412, preoccupied by 7. spinosus Neumann, 1899, Mem. Soc. Zool. Fr., 12: 146-147, = 7. fuscipes Koch, 1844. Fairchild, 1943, Amer. Jour. Trop. Med., 23: 583. Ixodes scuticrenatus Vazquez, 1946, An. Inst. Biol., 17: 237-245. New synonymy. Aside from the specimens from Didelphis marsupialis from Alhajuela (Canal Zone), L. H. Dunn, reported by Fairchild (1943), and Cooley and Kohls (1945), we have secured the following additional material. From Didelphis marsupialis, 20 lots totaling 33 males, 27 females, from the following localities : Tacarcuna Station (Darien), Aug.-Sept. 1958; Rio Mandinga (San Bias) , 27 May 1957 ; Cerro Azul (Panama) , May, Aug. 1955, Jan.-Apr. 1956; Cerro Campana (Panama), March 1961; Camp Pifia (Ca- nal Zone) , May, Aug. 1955, Jan., March 1956. From Philander opossum, 6 lots totaling 7 females, from Cerro Azul (Panama), 21 Dec. 1955; Camp Pina (Canal Zone), 19 Jan., 9 March, 1956; Fort Gulick (Canal Zone), 21 March 1961. From Marmosa robinsoni, 1 nymph, Almirante (Bocas del Toro) , 22 Feb. 1960. From Zygodontomys microtinus, 1 nymph, Tacarcuna Station (Darien) , 3 Sept. 1958. From Oryzomys sp., 2 nymphs, Cerro Pirre. 1600 feet (Darien), 6 Feb. 1961. The above records indicate that opossums are the favored hosts for the adults, but suggest that rodents are preferred by the earlier stages. The lo- calities are all in areas of high rainfall, from sea level to about 2000 feet ele- vation. In addition to Panama, this species has been recorded from southern Mexico, British Honduras, Guatemala, Colombia, Venezuela, Peru, French Guiana, Brazil and Argentina. The Rocky Mountain Laboratory has numerous lots, including many larvae and nymphs from rodents, that were collected in Trinidad by Dr. T. H. G. Aitken of the Trinidad Regional Virus Laboratory. Ixodes pomerantzi Kohls Ixodes pomerantzi Kohls, 1957, Jour. Parasit., 42, (6), pp. 639-642 (Dec. 1956). Only three lots of this tick have been taken, all from Sylvilagus brasilien- sis, as follows: Casa Tilley, Cerro Punta (Chiriqui), 1 May 1960, V. J. Tip- ton and C. M. Keenan, 2 males, 1 female, 13 nymphs; Bambito (Chiriqui), 5100 feet, 7 March 1962, V. J. Tipton, 1 male, 2 females, 17 nymphs; same locality, 13 March 1962, V. J. Tipton, 1 male, 1 female, 9 nymphs. FAIRCHILD, KOHLS AND TIPTON : TICKS 183 The species was described from adults from Sylvilagus brasiliensis in Peru and S. floridanus chiapensis in Guatemala. Ixodes rubidus Neumann Ixodes rubidus Neumann, 1901, Mem. Soc. Zool. Fr., 14: 282-283. Nuttall and War- burton, 1911, Ticks, pt. 2, pp. 175-176. Cooley and Kohls, 1945, Nat. Inst. Hlth. Bull., no. 184, pp. 153-156. In addition to the 13 females from Eira barbara reported by Cooley and Kohls (1945) from Boquete (Chiriqui) , we have secured five additional lots as follows: Nueva Colonia (Chiriqui), 30 Jan. 1960, V. J. Tipton and C. M. Keenan, from Procyon lotor, 7 females, 7 nymphs, 15 larvae ; Bambito (Chir- iqui), 15 Feb. 1960, V. J. Tipton and C. M. Keenan, from Mustela frenata, 1 female; Casa Tilley, Cerro Punta (Chiriqui), 5300 feet, 12 March 1962, V. J. Tipton, from Conepatus semistriatus, 9 females, 17 nymphs; Bambito (Chiriqui) , 5100 feet, 12 March 1962, V. J. Tipton, from Bassaricyon gabbii, 3 females, 4 nymphs ; and 13 March 1962, from Nasua nasua, 1 female. All of our material has come from small, wild carnivores of the families Pro- cyonidae and Mustelidae, and from localities over 5000 feet in Chiriqui Province. The species was described from a female from Bassariscus as- tutus, Guanajuato, Mexico, and the Rocky Mountain Laboratory has 2 fe- males from Didelphis sp., Yepocapa (Chimaltenango), Guatemala, 19 Jan. 1949, H. T. Dalmat. The male remains unknown. Ixodes tapirus Kohls Ixodes tapirus Kohls, 1957, Jour. Parasit., 42, (6), pp. 642-643, (Dec. 1956). Two lots of this recently described species, both from tapir, Tapirus bairdii, have been secured. Upper Rio Changuinola (Bocas del Toro), 10 May 1959, R. Hartmann, 1 female, and Rio Candela (Chiriqui), 6600 feet, Oct. 1953, R. Hartmann, 5 females. This species was previously known from a female from the woolly, or mountain tapir, Tapirus pinchaque, Rio Majuas, about 8600 feet, San Agustin (Huila), Colombia. Ixodes tiptoni Kohls and Clifford Ixodes tiptoni Kohls and Clifford, 1962, Jour. Parasit., 48, (2), pp. 182-184. Records of this recently described species will be found listed in detail in the original description. Subsequent collections include larvae, nymphs, females and a single male, from Sciurus granatensis and from rodent and bird nests, the latter probably in use by mammals, possibly squirrels. One lot is from Rio Changena (Bocas del Toro), 6000 feet, the others from the vicinity of Cerro Punta (Chiriqui). The important morphological features of the male are given in the key to the males of this genus. Ixodes venezuelensis Kohls Ixodes venezuelensis Kohls, 1953, Jour. Parasit., 39, (3), pp. 300-303. We have taken only females of this species and all but one lot came from a 184 ECTOPARASITES OF PANAMA camp on the slopes of Mount Tacarcuna (Darien) . Tacarcuna Yellow Fever Station, 31 Aug. 1958, GML, from Zygodontomys microtinus, 3 females; 1 Sept. 1958, same host, GML, 2 females ; 2 Sept. 1958, same host, GML, 1 female ; 7 Sept. 1958, same host, GML, 3 females ; 20 March 1959, GML, from Monodelphis adusta, 1 female; Cerro Azul (Panama), 14 March 1961, C. E. Yunker, from Oryzomys sp., 1 female. Kohls (1953) recorded females and nymphs of this species from Hete- romys anomalus anomalus and Monodelphis brevicaudata palliolata, State of Aragua, Venezuela, and females from Oryzomys caliginosus and Nec- tomys alfari, Antioquia, Colombia. Vogelsang and Bias (1953) recorded a female from Mus musculus, State of Aragua, Venezuela. The male is un- known. Genus Anocentor Schulze, 1937 The one known species in this genus is widely distributed in the New World from southern Texas, Florida, and the West Indies southward to Brazil. It is commonly referred to as the "tropical horse tick." Anocentor nitens (Neumann) Dermacentor nitens Neumann, 1897, Mem. Soc. Zool. Fr., 10: 376-378. Dunn, 1915, Ent. News, 26: 214-219; 1923, Amer. Jour. Trop. Med., 3, (2), pp. 93-94. Fair- child, 1943, Amer. Jour. Trop. Med., 23, (6), p. 583. Otocentor nitens (Neumann), Cooley, 1938, Nat. Inst. Hlth. Bull., no. 171, pp. 65-68, figs. Anocentor columbianus Schulze, 1937, Zool. Anz., 120: 24-27, figs. Dunn (1923,1934) reports this tick as primarily a parasite of horses and mules in Panama ; often all stages are found on the host simultaneously. He also indicates that the ticks seem to prefer the ears. Fairchild (1943) noted that it is often taken on cattle and that is was taken twice from deer. Sub- sequent records include thousands of specimens collected by Field in 1954- 1955 from horses in horse-baited mosquito traps at Fort Davis, and lesser numbers from horses at Fort Clayton. It is a curious fact that this New World tick is now very largely confined to domestic animals of Old World origin. Observations on the biology of this tick in Panama are given by Dunn (1915). Genus Dermacentor Koch, 1844 KEY TO PANAMANIAN SPECIES MALES AND FEMALES 1. Spurs of coxa I with proximal edges parallel or only slightly divergent latus Spurs of coxa I widely divergent 2 2. Males ventrally with a short retrograde tubercle on some of the festoons. Female scutum with whitish ornamentation indistinct; tarsi II to IV each with strong, sharp, subterminal and apical ventral spurs imitans Males ventrally without tubercles on the festoons. Female scutum with distinct whitish ornamentation; tarsi II to IV each with the subterminal ventral spur blunt and much shorter than the apical ventral spur halli FAIRCHILD, KOHLS AND TIPTON : TICKS 185 Only one species of this genus has been hitherto recorded from Panama, D. latus Cooley (Fairchild, 1943) . Extensive collecting in the highlands of Chiriqui and Darien Provinces has added two species and has indicated that the species in the genus appear to prefer a cool environment. Dermacentor latus Cooley Dermacentor latus Cooley, 1937, Jour. Parasit., 23: 262-264, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 583. Arthur, 1960, Ticks, pt. 5, pp. 102-103, figs. (fig. 185 is of scutum of D. dispar, not D. latus as stated) . Clifford and Kohls, 1962, Jour. Parasit., 48: 486-488, figs. This species was described from a single male from tapir in Costa Rica. Fairchild (1943) reported a male from dog, Boquete (Chiriqui). We have since secured additional material as follows : Rio Candela (Chiriqui) , >5000 feet, 20 Aug. 1954, R. Hartmann, from man, 1 male; Rio Candela, >6000 feet, Dec. 1953, R. Hartmann, from tapir, 12 males, 12 females ; Upper Rio Changuinola (Bocas del Toro) , >6000 feet, 10 May 1959, R. Hartmann, from tapir, 36 males, 12 females; Rio Changena (Bocas del Toro), 2400 feet, 16 Sept. 1960, C. M. Keenan and C. E. Yunker, from tapir, 16 males, 12 females ; Rio Changena, >6000 feet, 8 Sept. 1961, V. J. Tipton, from man, 1 female. We feel that the tapir is probably the true host of this tick in the adult stage. The female of the species was described by Clifford and Kohls from speci- mens from Rio Candela and Rio Changuinola. Dermacentor halli Mclntosh Dermacentor halli Mclntosh, 1931, Jour. Parasit., 18: 124 (brief description) ; 1932, Proc. U. S. Nat. Mus., 82, art. 4, pp. 1-6, figs. Cooley, 1938, Nat. Inst. Hlth. Bull., no. 171, pp. 55-58, figs. Arthur, 1960, Ticks, pt. 5, pp. 69-74, figs. The previous range of this species was from Yucatan, Mexico, north to extreme southern Texas, and the hosts were peccary and skunk. Our rec- ords are all from the vicinity of Cerro Punta (Chiriqui) , at about 5000 feet, collected by V. J. Tipton and C. M. Keenan (unless otherwise indicated), as follows : 3 May 1960, from rodent nest, 4 males, 1 female, 1 nymph ; 4 May 1960, from rodent nest, 2 males; 10 Jan. 1961, C. E. Yunker, from mouse, 1 male reared from nymph ; 6 March 1962, from Coendou mexicanus, 2 males, 1 female; 11 March 1962, same host, 1 male; 8 March 1962, same host, 7 males, 2 females ; 6 March 1962 from Myotis nigricans, 1 nymph ; Bambito (Chiriqui), 12 March 1962, from Coendou mexicanus, 1 male, 2 females; same data, 5 males, 1 nymph. The specimen from Myotis is possibly a stray. The Rocky Mountain Laboratory has records of this species as follows : from peccary, Los Pozos (Sinaloa) , Mexico, 26 Dec. 1937, 1 male ; from man or vegetation, Taninul (San Luis Potosi) , 24 Aug. 1961, 1 male, R. B. Eads ; from Coendou mexicanus, San Pedro Yepocapa (Chimaltenango), Guate- mala, April 1949, 5 males, 3 females, H. T. Dalmat ; same host, locality and collector, 9 Feb. 1950, 1 female; and 15 April 1951, 8 males, 3 females; same host and collector, Acatenango (Chimaltenango), Guatemala, 8 Feb. 1951, 1 male, 2 females ; from sloth, near San Jose, Costa Rica, March 1962, J. J. Shaw, 1 female. 186 ECTOPARASITES OF PANAMA The specimens from Panama, Costa Rica, and Guatemala differ some- what from those from Texas and Mexico. They are more slender and nar- rowed anteriorly, the palpi are thinner, the porose areas tend to be larger, and the females from Panama have the cornua much reduced or absent. If these ticks are actually D. halli as we assume them to be, it is strange that there are no records of this species from peccaries in Panama or elsewhere in Central America. Dermacentor imitans Warburton Dermacentor imitans Warburton, 1933, Parasitology, 24: 559-560, figs. Cooley, 1937, Jour. Parasit., 23: 261, figs. Arthur, 1962, Ticks, pt. 5, pp. 97-98, figs. This species, previously known only from the type lot consisting of 12 males and 1 female from peccary, Tayassu tajacu, (=Pecari angulatus), at Turrialba, Costa Rica, is now known from Panama, Guatemala and Mexico as detailed below. Our Panama material consists of 6 lots as follows: Rio Candela, (Chiri- qui) , 20 Aug. 1954, R. Hartmann, from man, 1 male ; near Almirante (Bocas del Toro) , 9 Feb. 1960, C. M. Keenan and V. J. Tipton, from collared peccary, Tayassu tajacu, 2 males ; Timishik, Rio Teribe (Bocas del Toro) , May 1962, from Tayassu tajacu, 10 males, 4 females ; Cerro Pirre (Darien) , 2 Feb. 1961, C. M. Keenan and C. E. Yunker, from Tayassu tajacu, 1 female; same lo- cality, host and collectors, 5 Feb. 1961, 3 males; same locality, 6 Feb. 1961, C. E. Yunker and V. J. Tipton, from work table with animal skins, host un- known, 1 male. All the localities are in the highlands, over 3000 feet, ex- cept the collection from Bocas del Toro Province. The collections of the Rocky Mountain Laboratory contain 1 male of this species from Tayassu sp. taken at Santa Clara near Cabanas (Zacapa), Guatemala, 1 Aug. 1948, at 5500 feet elevation by R. D. Mitchell and Luis de la Torre, Chicago Natural History Museum Guatemala Zoological Expedi- tion. There are also 3 lots from El Ocote, Ocozocoautla (Chiapas), Mexico, 2000 feet elevation, collected by Miguel Alvarez del Toro, with data as fol- lows : 1 male, 2 females from Tayassu sp., 4 May 1946 ; 1 male, same host and date ; and 2 males from Mazama americana, 25 May 1950. The latter speci- mens are probably strays. Genus Rhipicephalus Koch, 1844 Rhipicephalus sanguineus (Latreille) Ixodes sanguineus Latreille, 1806, Gen. Crust. Ins., 1: 157. Rhipicephalus sanguineus (Latreille), Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 94. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6) , p. 583. Cooley, 1946, Nat. Inst. Hlth. Bull., no. 187, pp. 24-29, figs. The common brown dog tick, is found on nearly all dogs throughout Panama. Fairchild removed nearly a pint of ticks from a stray mongrel several years ago. Infestation of houses by this tick is not uncommon, and often leads to excited inquiries on the part of the householders, though we have not encountered evidence of the ticks actually biting man. We have FAIRCHILD, KOHLS AND TIPTON : TICKS 187 occasional records from animals in captivity, such as marmosets, capybara, and domestic rabbit, but these are no doubt accidental. Genus Boophilus Curtice, 1891 Boophilus microplus (Canestrini) Haemaphysalis micropla Canestrini, 1887, Atti Soc. Veneto Trent. Sci. Nat., 11 : 104. Margaropus annulatus australis (Fuller) , Dunn, 1923, Amer. Jour. Trop. Med., 3, (2) , p. 95. Boophilus microplus (Canestrini), Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 586. Cooley, 1946, Nat. Inst. Hlth. Bull., no. 187, pp. 17-22, figs, (with Uro- boophilus cyclops Minning as synonym). Uroboophilus cyclops Minning, 1934, Zeitschr. Parasitenk., 7: 33, figs. This is the most abundant tick on cattle throughout Panama. We have records from as high as Cerro Punta, (5800 feet), in Chiriqui Province. Aside from twenty-three records from cattle, we have three records from horse, three from pig, and one each from goat and deer. Dunn (1923) re- corded it, as Margaropus annulatus australis Fuller, from dogs as well. He also recorded the North American cattle tick, B. annulatus, from a recently imported bull, but it does not seem to have become established here. Genus Haemaphysalis Koch, 1844 KEY TO PANAMANIAN SPECIES MALES AND FEMALES Palpal segment 3 with a very short retrograde ventral spur; ventral cornua present; hypostome with dentition 3/3 leporispalustris Palpal segment 3 with a long retrograde ventral spur; ventral cornua absent; hypostome with dentition 4/4 or 5/5 juxtakochi NYMPHS Coxa I with a small external spur ; palpal segment 2 ventrally with 4 or more stout hairs on the internal margin leporispalustris Coxa I with external spur absent; palpal segment 2 ventrally with 2 fine hairs on the internal margin juxtakochi LARVAE Cornua present on basis capituli dorsally; scutum relatively narrower (approxi- mately 0.25 mm. long, 0.30 mm. wide) leporispalustris Cornua absent; scutum relatively broader (approximately 0.24 mm. long, 0.37 mm. wide) juxtakochi Haemaphysalis juxtakochi Cooley Haemaphysalis kochi Aragao, 1908, Trab. Inst. Manguinhos, pp. 3-6 (reprint). Cooley, 1946, Nat. Inst. Hlth. Bull., no. 187, pp. 44-47, figs. Haemaphysalis juxtakochi Cooley, 1946, loc. cit., no. 187, pp. 48-51, figs. Kohls, 1960, Jour. Parasit. 46, 3, pp. 356-358, figs. (H. kochi Aragao and H. kohlsi Aragao and Fonseca, 1951 reduced to synonyms of H. juxtakochi.) Haemaphysalis kohlsi Aragao and Fonseca, 1951, Mem. Inst. Osw. Cruz, 49: 574 (new name for H. kochi Aragao preoccupied by H. concinna var. kochi Neumann, 1905) . Dunn (1923) and Fairchild (1943) recorded this species (as H. kochi) from deer (Odocoileus) and tapir, and on vegetation, while Kohls (1960) re- 188 ECTOPARASITES OF PANAMA corded the species from Odocoileus and Mazama. Subsequent collections are all from the Canal Zone area, as follows : Gamboa road, 14 March 1961, N. B. Gale, from Nasua nasua dead on road, 3 nymphs ; Juan Mina, 10 March 1944, GML, from peccary, 1 nymph ; Rio Casaya, 27 Feb. 1944, GML, from collared peccary, 1 nymph ; Summit Gardens, 29 Aug. 1959, V. J. Tipton and C. M. Keenan, from O. virginianus, 24 males, 15 females ; vicinity of Gamboa, 24 Oct. 1959, same collectors and host, 7 males, 6 females ; Cerro Tigre, 4 July 1961, same collectors and host, 7 males, 3 females; Curundu, 10 March 1961, same collectors, from Coendou rothschildi, 1 nymph ; Barro Colorado Island, 15 July 1960, same collectors, from Tapirus bairdii, 1 female. Deer, especially Odocoileus, seem to be the preferred hosts of the adults, while nymphs have been secured from a variety of animals. Haemaphysalis leporispalustris (Packard) Ixodes leporis-palustris Packard, 1869, Ann. Rept. Peabody Acad. Sci., p. 67. Haemaphysalis leporis-palustris (Packard), Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 96. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 586. Cooley, 1946, Nat. Inst. Hlth. Bull., no. 187, pp. 31-36, figs. Kohls, 1960, Jour. Parasit., 46, (3), pp. 355-361. Dunn (1923) recorded specimens from domestic rabbit and Dasyprocta sp., while Kohls (1960) gave a number of records from Sylvilagus and birds. Nearly all of our specimens were taken from Sylvilagus brasiliensis, as fol- lows: Cerro Punta (Chiriqui), 1 May 1960, V. J. Tipton and C. M. Keenan, 2 females; Rio Mandinga (San Bias), 29 May 1957, P. Galindo, 6 males, 3 nymphs, 1 larva ; 2 May 1957, same locality and collector, 2 females ; Cerro Azul (Panama) , 15 March 1961, C. E. Yunker, 1 female held for oviposition, eggs and larvae secured ; 22 March 1961, same locality and collector, 8 males, 2 females, 2 nymphs; Bambito (Chiriqui), 5100 feet elevation, 7 and 13 March 1962, V. J. Tipton, 1 male, 4 females, 4 nymphs, 1 larva ; Cerro Punta (Chiriqui), 5300 feet elevation, 11 March 1962, V. J. Tipton, 2 males, 2 fe- males, 2 nymphs; Las Palmitas (Los Santos), 19 and 28 Feb., and 1 March 1962, V. J. Tipton (4 lots), 47 males, 13 females, 40 nymphs, 2 larvae; El Hato (Chiriqui), 7 Jan. 1961, L. C. Wislocki and C. E. Yunker, from Pero- myscus sp., 1 larva. Genus Amblyomma Koch, 1844 KEY TO PANAMANIAN SPECIES MALES 5 1. Marginal groove complete, limiting all festoons 2 Marginal groove incomplete or absent 10 2. Coxa I with two long equal or subequal spurs, both spurs at least twice as long as broad 3 Coxa I with one or both spurs short or medium 6 5 Excluding crassum Robinson, 1926. We have not seen males but we believe the description by Mendez and Ortiz (1957) may be that of another species, perhaps sab- anerae. FAIRCHILD, KOHLS AND TIPTON : TICKS 189 3. Spurs on coxa I very long and acute, much longer than the spur on coxa IV, the external spur slightly curved outward near the tip. Body elongate oval ovale Spurs on coxa I as long as spur on coxa IV or shorter 4 4. Coxa I with broad stout spurs. Scutum with punctations numerous, large and deep coelebs Coxa I with slender acute spurs. Scutum with punctations few and fine 5 5. Body broad oval. Elements of scutal pattern all of about equal intensity. Medium sized species tapirellum Body elongate oval, lateral margins subrectilinear. Longitudinal elements of scutal pattern accentuated, giving a more striped appearance. Very small species oblong oguttatum 6. Scutum with elongate, keel-like ridge in posteromedian area pecarium Scutum without keel-like ridge 7 7. Palpal segment 1 with a stout ventral retrograde spur. Trochanters I-IV each with a small ventral spur. Scutum inornate or very slightly ornate. Small species 8 Palpal segment 1 without ventral spur. Trochanters without spurs. Scutum distinctly ornate 9 8. Coxa I with internal spur much shorter than the external. Scutum inornate parvum Coxa I with short sub-equal spurs. Scutal ornamentation indistinct or absent auricularium 9. Capitulum sub-rectangular. Coxae II and III each with spurs in the form of a broad salient ridge ; a long stout spur on coxa IV cajennense Capitulum trapezoidal. Coxae II-IV each with a short triangular spur. Ventral plaques large geayi 10. Marginal groove incomplete, terminating posteriorly at the third festoon on each side. Body large, elongate oval. Ventral plaques large. Coxa I with two small unequal spurs; coxae II-IV each with a single short, pointed spur longirostre Marginal groove absent 11 11. Coxae II-IV each with two spurs (internal spur of IV sometimes absent). Para- sites of reptiles and amphibians 12 Coxae II-IV each with a single spur. Parasites of mammals 13 12. Dentition of hypostome 3/3. Coxal spurs pointed and not notably flattened; in- ternal spurs much smaller than the externals. Scutum smooth; ornamentation not limited primarily to the anterolateral fields; punctations scattered, unequal in size, larger in the peripheral areas dissimile Dentition of hypostome 4/4. Coxal spurs broadly rounded, blunt, subequal. Scutum roughened by small areas of punctation-free elevations; ornamentation limited primarily to the anterolateral fields; punctations dense, coarse and deep sabanerae 13. Coxa I with two long contiguous equal or sub-equal spurs 14 Coxa I with one or both spurs short or medium 15 14. Palpal segment 1 ventrally with a broad flattened expansion; segment 2 with a pronounced salient ridge surrounding the posterior border and forming dors'ally a strong retrograde spine nodosum Palpal segment 1 ventrally without broad expansion ; segment 2 without posterior salient ridge but posterodorsal margin pointed calcaratum 15. Coxa I with external spur long and pointed, the internal spur short and blunt. Festoons, except the first and median, each bearing a small posteriorly-directed tubercle. Scutum flat; punctations numerous, shallow. Small species, .naponense Coxa I with subequal spurs 16 16. Small species; scutum less than 4.5 mm. long, indistinctly ornate; punctations numerous, small. Parasite of pacas primarily pacae Large species, scutum more than 4.5 mm. long 17 190 ECTOPARASITES OF PANAMA 17. Spurs of coxa I short, approximately as long as wide, well separated. Integument yellowish. Scutum with indistinct ornamentation and with numerous small deep punctations. Cornua very short and broad. Parasite of anteaters pri- marily pictum Spurs of coxa I medium, approximately twice as long as broad, contiguous. Scutum with ornamentation much reduced but distinct. Punctation numerous, coarse, absent from three or four small isolated elevated areas on each side and a similar median area. Cornua long. Parasite of sloths primarily varium FEMALES 1. Coxa I with equal or subequal spurs 2 Coxa I with the external spur much longer than the internal 14 2. Coxa I with long spurs, more than twice as long as broad 3 Coxa I with medium or short spurs 6 3. Coxa I with slender spurs 4 Coxa I with stout spurs 11 4. Coxa I with very long acute spurs, the external slightly curved outward near its tip ovale Coxa I with moderately long spurs, apical portion of external spur not curved .... 5 5. Genital apron overlaid on each side posterolaterally by a conspicuous, blunt, flattened projection darker in color than the apron and adjacent integument. Punctations not limited to anterior half of scutum tapirellum Genital apron overlaid on each side with an inconspicuous, long, slender projection. Punctations very scant on posterior half scutum oblong 'oguttatum 6. Coxa II-IV with two spurs (internal spur of IV sometimes absent). Parasites of reptiles and amphibians 7 Coxa II-IV with one spur. Parasites of warm-blooded hosts 9 7. Hypostome dentition 3/3. Internal spur of coxa IV very small, sometimes absent dissimile Hypostome dentition 4/4 8 8. Very large species; scutum over 4 mm. wide. Coxae II-IV each with a pair of short rounded plate-like spurs connected by a salient sharp-edged ridge . . . crassum Smaller species; scutum less than 3.5 mm. wide. Spurs of coxae II-IV distinctly separated sabanerae 9. Large species; scutum approximately 3 mm. wide; punctations numerous, coarse or medium. Basis capituli with cornua. Hypostome dentition 4/4 10 Small species; scutum approximately 2 mm. wide; punctations numerous, fine. Cornua absent. Hypostome dentition 3/3 auricularium 10. Spurs of coxa I short, about as long as wide. Scutum with punctations medium in size, uniformly distributed. Parasite of anteaters primarily pictum Spurs of coxa I about twice as long as wide. Scutum with large deep punctations, these usually few or absent from median field near posterior margin. Para- site of sloths primarily varium 11. Scutum with extensive pale ornamentation. Palpal segment 1 ventrally with a large elongate flattened plate coelebs Scutum extensively dark-colored 12 12. Scutum inornate or indistinctly ornate. Coxa I with external spur the longer, .pacae Scutum usually distinctly ornate. Coxa I with spurs equal 13 13. Scutum with ornamentation consisting of an irregular pale spot in the posterior angle and sometimes one also in each posterolateral field. Palpi slender and smooth calcaratum Scutum with ornamentation in the form of a pale spot in the posterior angle and a Y-shaped figure in each lateral field. Palpi heavier and more rugose, segment 2 with an oblique ridge dorsally nodosum 14. Scutum ornate 15 Scutum inornate; small, about 1.5 mm. wide parvum FAIRCHILD, KOHLS AND TIPTON : TICKS 191 15. Coxae II and III with broad flat ridge-like spurs much broader than long 16 Coxae II and III with spurs scarcely broader than long 17 16. Palpal segment 2 about two and one-half times as long as segment 3. Festoons ventrally somewhat rugose and poorly denned, first four on each side of the median each with a well-developed tubercle at the posterointernal angle. Internal spur of coxa I broad, blunt pecarium Palpal segment 2 about twice as long as segment 3. Festoons ventrally smooth and clearly defined, each except the median, with a much smaller tubercle at the posterointernal angle. Internal spur of coxa I narrower and more sharply pointed cajennense 17. A small to medium-sized species. Coxa I with external spur long, slender, pointed; internal spur, short, blunt, stout. Scutum with extensive pale ornamentation naponense Large species. Coxa I with both spurs short, flat; internal spur very small. Scutum mostly dark colored 18 18. Scutum cordiform, about as long as wide, with irregular pale areas in the postero- median field geayi Scutum elongate oval, indistinctly ornate with an irregular longitudinal pale area in the median field. Hypostome very long and sharply pointed. Legs, especially IV, notably long longirostre Amblyomma auricularium (Conil) Ixodes auricularius Conil, 1878, Act. Acad. Nac. Cienc. Exact., 3: 99-110, figs. Amblyomma concolor Neumann, 1899, Mem. Soc. Zool. Fr., 12: 222. Robinson, 1926, Ticks, pt. 4, pp. 66-69, figs. Amblyomma auricularium (Conil), Lahille, 1905, An. Minist. Agric., Secc. Zootec. Bact., Vet., Zool., 2, (2), pp. 34, 145-148, figs. Aragao, 1935, Mem. Inst. Osw. Cruz, 30, (3), p. 528 (with A. concolor as synonym). Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), pp. 583-584. Amblyomma curruca Schulze, 1936, Zeitschr. Parasitenk., 8, (6), pp. 621-622. New synonymy. This is an abundant species whose chief hosts appear to be armadillos, although it has been taken in smaller numbers on a variety of other mam- mals, especially marsupials and edentates. It appears to range throughout the Republic, though most of our records are from the drier Pacific side and all are from elevations below 500 feet. We have specimens from the fol- lowing hosts : Dasypus novemcinctus (22 lots, 164 males, 122 females, 28 nymphs, 7 larvae) ; Cabassous centralis (1 lot, 1 male, 2 nymphs) ; Taman- dua tetradactyla (12 lots, 14 males, 13 females) ; Didelphis marsupialis (2 lots, 3 males, 1 female) ; Philander opossum (3 lots, 4 males, 4 females, 13 nymphs, 5 larvae) ; Sigmodon hispidus (2 lots, 2 females reared from 2 nymphs) ; Hydrochaeris hydrochaeris (1 lot, 1 male) ; Nasua nasua (1 lot, 2 males) ; domestic dog (1 lot, 1 male) ; animal burrows, probably dug by armadillos (2 lots, 1 male, 1 female, many nymphs). The three lots from Philander consisted, in part, of engorged nymphs from which adults were reared. The range of this species extends from Mexico to Argentina. Examination of the types of A. curruca by Kohls revealed that this spe- cies is a synonym of A. auricularium, not of A. parvum as stated by Aragao and Fonseca (1953). 192 ECTOPARASITES OF PANAMA Amblyomma cajennense (Fabricius) Acarus cajennensis Fabricius, 1787, Mant. Insect., p. 372. Amblyomma cajennense (Fabricius), Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 96; 1934, Psyche, 41, (3), p. 182; 1934, Jour. Parasit., 20, (5), p. 312. Robin- son, 1926, Ticks, pt. 4, pp. 48-54, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. Cooley and Kohls, 1944, Jour. Parasit, 30, (2), pp. 83-87, figs. Aragao and Fonseca, 1953, Mem. Inst. Osw. Cruz, 51: 485-488. Kohls, 1958, Jour. Parasit., 44, (4), pp. 430-433, figs. Although this is an abundant species, it is interesting that adults are not often taken from wild hosts. Our records, therefore, do not adequately indi- cate its abundance, since collections from domestic animals have been few and confined to small samples of the population of ticks on any one animal. For this reason no numbers of ticks collected are given below in the host list. The species seems to be present in most parts of the country up to elevations of over 5000 feet, but is most abundant in the drier areas and where cultiva- tion has reduced the original forest cover. Our records are, as with most species of Amblyomma, confined almost wholly to adults, as nymphs and larvae are not yet identifiable with certainty: Horse (24 lots), cattle (20 lots), man (13 lots), dog (5 lots), horse-baited mosquito traps (3 lots), Tamandua (3 lots), Didelphis (2 lots), deer (2 lots), domestic hog (2 lots), chicken (2 lots) , on vegetation and ground (2 lots) , tapir, peccary, Dasypus, goat and a hawk, Buteo magnirostris (1 lot each). As can be seen by the above, horses and cattle seem to be the preferred hosts of the adults. Most of the collections from man consist of one or a few specimens taken crawling on the clothing, though the species will attack if given the opportunity. It is probably the larvae of this species which form the bulk of the seed ticks which attack man in enormous numbers during the dry season. The range of A. cajennense extends from southern Texas and islands of the Caribbean to Argentina. In Panama, natural infection of this tick with the etiologic agent of Rocky Mountain spotted fever, Rickettsia rickettsi, has been demonstrated by Rodaniche (1953) and the species is believed to be a vector here as in Mexico, Colombia and Brazil. Amblyomma calcaratum Neumann Amblyomma calcaratum Neumann, 1899, Mem. Soc. Zool. Fr., 12: 226. Robinson, 1926, Ticks, pt. 4, pp. 191-194, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. This large tick is almost restricted to anteaters ; of 24 lots, 20 are from Tamandua tetradactyla and one from Myrmecophaga tridactyla, the latter a rare animal in Panama. Two of the remaining lots, collected by L. H. Dunn from Choloepus hoffmanni, include 5 adults, Panama, and 1 female from Rio Abajo (Panama), 28 Sept. 1931. One lot is from Mazama americana, 1 male, Rio Teribe (Bocas del Toro) , 10 Aug. 1962. We have studied 62 males, 24 females, from localities on both coasts and from Darien to Bocas del Toro Provinces, but not from any elevations over 2500 feet. This species is often found together with A. nodosum on the same animal, and has a similar geo- graphic range in Panama. FAIRCHILD, KOHLS AND TIPTON : TICKS 193 In addition to Panama, this species has been recorded from Venezuela, French Guiana, Ecuador, Brazil, and Paraguay. The Rocky Mountain Lab- oratory collection contains 1 male, San Rafael, Trinidad, 27 Feb. 1947, from Tamandua tetradactyla longicauda, Frank Wonder, Chicago Natural His- tory Museum; 5 males, Rio Guapaya, La Macarena (Meta), Colombia, 21 March 1957, from Tamandua sp., K. von Sneidern, Chicago Natural History Museum; 1 male, 1 female, San Jose, Costa Rica, 5 Feb. 1934, from "small anteater", F. Nevermann ; 7 males, 2 females, Kate's Lagoon, British Hon- duras, 26 Feb. 1940, from "lesser anteater", I. T. Sanderson, Chicago Natural History Museum. Amblyomma coelebs Neumann Amblyomma coelebs Neumann, 1899, Mem. Soc. Zool. Fr., 12: 223. Robinson, 1926, Ticks, pt. 4, pp. 30-33. Dunn, 1934, Jour. Parasit., 20, (5), p. 312; Psyche, 41, (3), p. 182. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. Dias, 1958, An. Inst. Med. Trop., 15, (2), pp. 507-508 (with A. bispinosum Neumann as synonym). Amblyomma bispinosum Neumann, 1906, Arch. Parasit., 10, (2), p. 204. Dunn (1934, 1934a) recorded this species from tapir and horse, Progreso (Chiriqui), and from three tapirs from Summit (Canal Zone), and Aguas Buenas (Panama). Fairchild (1943) added records of single specimens from tapir and horse. Subsequent records, mostly from tapir, are as fol- lows : Cerro Azul (Panama) , 12 May 1957, GML, from tapir, 1 female and 4 nymphs presumably this species; Rio Chico Hydrographic Station, Upper Rio Chagres (Panama), 20 March 1948, GML, from tapir, 8 males, 2 fe- males ; Rio Tuira at mouth of Rio Paya (Darien) , 25 Feb. 1958, P. Galindo, from Agouti paca, 1 female ; same locality, 3 July 1958, P. Galindo, no host, free; Rio Mandinga (San Bias), 18 May 1957, GML, no host, free; Rio Changena (Bocas del Toro), 2400 feet, 2 Aug. 1961, R. Hartmann, from tapir, 5 males ; same locality 16 Sept. 1961, C. M. Keenan and C. E. Yunker, from tapir, 1 male; Rio Teribe (Bocas del Toro), 10 Aug. 1962, GML, 6 males, 6 females, from Tapirus bairdii; Porto Bello Trail, Continental Di- vide (Panama), 30 May 1914, Hallinan, no host, 1 female; Barro Colorado Island (Canal Zone) , 30 March 1924, in woods, no host, 3 females. The range of this species extends from Mexico to Brazil and northern Argentina. In addition to hosts mentioned above, we have seen specimens from Mazama americana in Mexico and from cattle in Nicaragua. Amblyomma crassum Robinson Amblyomma crassum Robinson, 1926, Ticks, pt. 4, pp. 177-179, figs. Osorno-Mesa, 1941, An. Acad. Nac. Med., 1938-40, pp. 413, 422, 425-426. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. Mendez and Ortiz, 1957, Mem. Soc. Cienc. Nat. LaSalle, 17: 190-199, figs. Dias-Ungria, 1957, Rev. Sanid. Asist. Social, 22, (5-6), p. 459. Dias, 1958, An. Inst. Trop. Med., 15, (3), pp. 496-497 (as synonym of A. humerale Koch). Although we have not taken this species in Panama, we include it because it was described from "Darien Country, Colombia," which may have been within the present boundaries of Panama. The description was based on a female found on a "land-tortoise". Available specimens recorded by Fair- 194 ECTOPARASITES OF PANAMA child (1943) as this species or sabanerae Stoll or humerale Koch prove to be sabanerae, and there are as yet no authentic records of the occurrence of either crassum or humerale in Panama. Mendez and Ortiz (1957) described the male and redescribed the female which they believed to be crassum found on "land-tortoise (Testudo sp.)" from the territory of the Delta Amacuro in Venezuela but their descriptions and figures appear to be those of another species, perhaps sabanerae. Os- orno-Mesa (1941) included both sexes in his key and recorded males from land turtles taken in two localities in Intendencia del Meta, Colombia. Dias- Ungria (1957) recorded specimens from Testudo sp. in Venezuela. The Rocky Mountain Laboratory collection contains a female taken from "Tes- tudo tabulata" (=Geochelone sp.), Unguia, Golfo de Uraba, Choco, Colom- bia, March, 1950, P. Hershkovitz, Chicago Natural History Museum ; and a female found on a log, Sheshea River Basin at the headwaters of the Peru- vian Amazon, Peru, 1960 or 1961, by G. E. Dickinson and received from Dr. R. E. Ryckman of Loma Linda University, Loma Linda, California. Amblyomma dissimile Koch Amblyomma dissimile Koch, 1844, Arch. Naturg., 10: 225. Dunn, 1918, Jour. Parasit., 5, (1), pp. 1-10; 1923, Amer. Jour. Trop. Med., 3, (2), p. 97. Robinson, 1926, Ticks, pt. 4, pp. 163-167. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. Cooley and Kohls, 1944, Jour. Parasit., 30, (2), pp. 98-102, figs. Aragao and Fonseca, 1953, Mem. Inst. Osw. Cruz, 51: 489 (with A. deminutivum Neumann, 1899 as synonym). Dias, 1958, An. Inst. Med. Trop., 15, (2), pp. 494-496 [as synonym of A. bibroni (Gervais), 1842]. This is the common reptile tick in Panama. Dunn (1923) found that over 60 percent of the snakes, 72 percent of the toads and 84 percent of the iguana lizards he examined were infested. Dunn (1918) also made a study of the life history of this tick. Fairchild (1943) noted that laboratory infes- tations on captive snakes were severe enough to kill the snakes. In one case, 190 ticks were removed from a single small fer-de-lance (Bothrops atrox) . Tipton more recently secured 1800 ticks of all stages from a single snake. All stages are frequently found on a single host animal. The bulk of our material has come from the Canal Zone and vicinity, with no specimens from elevations over 800 feet. We have about 95 lots containing adults, mostly from snakes and iguanas, and an additional 22 lots of nymphs and/or larvae, mostly from lizards, which are probably this species. Two unusual records are 1 male, 1 nymph, 2 larvae from a hatchling aquatic turtle, Pseudemys scripta, Barro Colorado Island, Canal Zone, 20 Aug. 1961, J. M. Legler, and 1 male from boat-billed heron, Cochlearius cochlearius, Juan Mina, Chagres River, 5 April 1962, C. L. Hayward. The range of this species extends from Florida, Mexico, and the West Indies, to Argentina. Dias (1958) maintains that Neumann's (1899) summary of the Gervais (1842) and Packard (1869) descriptions of Ixodes bibroni applies to A. dis- simile but we prefer to retain dissimile as the valid name of this species. FAIRCHILD, KOHLS AND TIPTON : TICKS 195 Amblyomma geayi Neumann Amblyomma geayi Neumann, 1899, Mem. Soc. Zool. Fr., 12: 223-224; 1901, Mem. Soc. Zool. Fr., 14: 299. Robinson, 1926, Ticks, pt. 4, pp. 59-61, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. Bias, 1961, An. Serv. Veter. Mozambique, 7: 240-242 [as synonym of A. perpunctatum (Packard), 1869]. We have secured some 60 lots of this species, practically all from our two species of sloth, Bradypus infuscatus, the three-toed sloth, and Choloepus hoffmanni, the two-toed sloth. The former is much more often seen, as it feeds on the foliage of the Guarumo (Cecropia pentandra) a common sec- ond-growth tree in the Canal Zone and elsewhere in the lowlands of Panama. Sloths are frequently encountered crossing highways and are then easily captured, or their corpses may be searched for ticks. However, they are seldom seen and difficult to secure in less inhabited areas, since they are hard to collect from the tree tops. This we believe accounts for the fact that with the exception of three lots from Bocas del Toro Province, all our material is from the Canal Zone or immediate vicinity. We have 35 lots from Bradypus, 227 males, 36 females, 8 nymphs ; 12 lots from Choloepus, 36 males, 12 fe- males ; and 8 lots from unidentified sloths, 47 males, 8 females, 2 nymphs, 1 larva. In addition we have a single nymph from the woolly opossum, Calu- romys derbianus, and 2 males, 1 female, said to be from "Eaton's opossum", which may refer to Didelphis marsupialis etensis. The remaining lots are without host, but are probably from sloths. Nymphs, possibly of this spe- cies, are occasionally taken with adults on sloths. This species, originally described from specimens from Brazil and "Darien (Colombie)" which may have been within the present boundaries of Panama, is also recorded from British Guiana and French Guiana. In addition, the Rocky Mountain Laboratory collection contains specimens from Choloepus, Iquitos (Loreto) , Peru, Sept. 3, 1956, C. Kalinowski, Chicago Nat- ural History Museum. After reviewing the literature, Bias (1961) concluded that A. geayi is the species that was very inadequately described by Packard (1869) as Ixodes perpunctatus. We cannot accept his conclusion and we regard A. geayi as the valid name of this species. Amblyomma longirostre (Koch) Haemalastor longirostris Koch, 1844, Arch. Naturg., 10, (1), p. 223. Amblyomma longirostre (Koch), Robinson, 1926, Ticks, pt. 4, pp. 137-140, figs., syn- onymy. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 585. Amblyomma avecolens Cooley and Kohls, 1944, Jour. Parasit., 30, (2), pp. 107-109, figs. New synonymy. This species appears to be a specific parasite of porcupines, all our rec- ords of adults and those of Dunn (1923) being from Coendou rothschildi. The males are sometimes found attached to the spines of the host as noted by some earlier workers. Aragao (1936) says that the larvae and generally the nymphs are parasites of birds, and we have nymphs from seven species of birds in Panama. We have records only from the Canal Zone and Darien Province, listed below, but porcupines are uncommon or at least seldom 196 ECTOPARASITES OF PANAMA taken, so that we know little of the distribution of either host or tick in Pan- ama. It is noteworthy that a series of the highland porcupine, Coendou mexicanus, was not infested with this tick, but with Dermacentor halli. From the Canal Zone : Gatun, 2 Jan. 1932, L. H. Dunn, 1 male, 1 nymph ; Juan Mina, 24 April 1940, GML, 2 females ; Fort Sherman, 30 July 1959, V. J. Tipton and C. M. Keenan, 1 male, 1 female ; same locality and collectors, 29 July 1960, 1 female; same locality, F. S. Blanton, 29 June 1951, 1 male; Curundu, 10 March 1961, V. J. Tipton and C. M. Keenan, 5 males, 1 female; same locality, 19 April 1962, C. E. Yunker, 1 male; Pedro Miguel River, 28 Feb. 1962, V. J. Tipton, 9 males ; Barro Colorado Island, 4 March 1955, C. Rettenmeyer, without host data, 1 female, Kansas University 955. Of nymphs, we have records from the following: Saltator albicollis, Saltator maximus, Icterus chrysater, Xiphorhynchus sp., Malacoptila panamensis, Cymbilaimus lineatus, Cacicus microrhynchus and Querula purpurata, all but one belonging to the Passerif ormes. Five of the 12 lots of nymphs were from the vicinity of Cerro Pirre (Darien) , at about 1600 feet, the rest from Canal Zone localities. Some of the undetermined nymphs and larvae from birds discussed elsewhere in this paper may also be longirostre. The known breeding range of this species, as evidenced by collections of adults, extends from Panama to Brazil but nymphs are found occasionally on birds as far north as the United States. The Rocky Mountain Laboratory collection contains nymphs reported by Cooley and Kohls (1944) as ave- colens from British Honduras and southern Texas, as well as a nymph taken off the base of the bill of a cardinal at Nashville, Tennessee, 17 July 1947, and several nymphs from birds in Chiapas, Mexico. Kohls has seen a nymph that was found on the throat of a white-eyed vireo in Leon County, Florida, 2 April 1957, L. J. Stannard, Illinois Natural History Survey. Amblyomma naponense (Packard) Ixodes naponensis Packard, 1869, Ann. Rept. Peabody Acad. Sci., p. 65. Amblyomma naponense (Packard), Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 99. Osorno-Mesa, 1941, An. Acad. Nac. Med., 1938-1940: 426 (with A. mantiquirense Aragao, 1908 as synonym). Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 585. Amblyomma mantiquirense Aragao, 1908, Brazil. Med., 22: 251-252. Robinson, 1926, Ticks, pt. 4, pp. 212-215, figs. Dunn, 1934, Psyche, 41, (3) , p. 182. Dunn (1923, 1934) records the species from Tamandua and collared pec- cary. Our collections, with few exceptions, are from collared peccary and are summarized as follows: Tayassu tajacu (13 lots, 11 males, 27 females) ; no host (3 lots, 6 males, 2 females) ; man (1 lot, 1 male, 1 female) ; Tamandua tetradactyla (1 lot, 1 female) ; Nasua nasua (1 lot, 2 females) ; raccoon (1 lot, 1 female) . This species appears to prefer the wetter areas of Panama, as we have material only from the Provinces of Darien, San Bias, Colon, Bocas del Toro and the Canal Zone. Only two collections, from Balboa and Fort Clayton in the Canal Zone, are from relatively dry areas. Only one collec- tion was made above 2000 feet. This species has also been recorded (as mantiquirense) from British FAIRCHILD, KOHLS AND TIPTON : TICKS 197 Guiana, French Guiana, and Brazil. The Rocky Mountain Laboratory col- lection contains specimens from Tayassu tajacu, Socorre, Upper Rio Sinu (Bolivar), Colombia, 12 March 1949, P. Hershkovitz, Chicago Natural His- tory Museum ; and from "wild pig", Sheshea River basin at the head of the Peruvian Amazon, Peru, 1960 or 1961, G. E. Dickinson, received from Dr. R. E. Ryckman, Loma Linda University, Loma Linda, Calif. Amblyomma nodosum Neumann Amblyomma nodosum Neumann, 1899, Mem. Soc. Zool. Fr., 12: 224-225. Dunn, 1923, Amer. Jour. Trop. Med., 3, (2), p. 98. Robinson, 1926, Ticks, pt. 4, pp. 196-199, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 585. This species is rather easily confused with A. calcaratum, as the two are quite similar in gross appearance and both occur on the same host, often to- gether. We have 19 lots from Tamandua tetradactyla, one lot from Myrme- cophaga tridactyla, and one lot from unknown host, totaling 207 males, 48 fe- males, of which 95 males, 13 females came from one animal. It occurs throughout the lowlands, including at least San Jose Island in the Pearl Islands, but we have no records from elevations over 1000 feet. This species, the adults of which are parasitic only on anteaters so far as known, was described from specimens from Costa Rica and besides Panama, it has also been recorded from Guatemala, Colombia, Venezuela, and Brazil. The Rocky Mountain Laboratory collection contains 1 male, 2 females from "lesser anteater", El Recreo, Nicaragua, 9 Feb. 1951, W. H. Dickinson. Amblyomma oblongoguttalum Koch Amblyomma oblong oguttatum Koch, 1844, Arch. Naturg., 10: 228. Robinson, 1926, Ticks, pt. 4, pp. 33-36, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 585. Kohls, 1958, Jour. Parasit., 44, (4), pp. 430-433. This is probably the commonest and most ubiquitous tick in Panama, at least at lower elevations. Our records indicate that it prefers forested coun- try, or at least areas of moderately high rainfall, since we have relatively few records of its occurrence in the drier areas of the Pacific coast. Dunn ( 1923 ) records it, as darlingi, from wild turkey, Crax panamensis (Crax rubra) from the Boqueron River region and from a black vulture, Catharista urubu (=Coragyps atratus), while Robinson (1926) records specimens from a turkey buzzard, Catharista atratus (=Coragyps atratus) from Empire (Canal Zone) . We have examined 131 lots of adults from 21 different mam- mal hosts listed below. Deer (Odocoileus) (18 lots) ; horse (17 lots) ; cattle (15 lots) ; dog (19 lots) ; peccary (13 lots) ; man (9 lots) ; Nasua nasua (10 lots) ; Tamandua tetradactyla (7 lots) ; tapir (7 lots) ; no host, free on vegetation, etc. (12 lots) ; domestic hog (3 lots) ; Mazama (2 lots) ; Dasyprocta (3 lots) ; sloth (Choloepus), Myrmecophaga tridactyla, goat, armadillo, Procyon cancri- vorus, Chironectes minimus, Cebus capuchinus, Eira barbara, and domestic cat, (1 lot each). According to Robinson (1926) and data of the Rocky Mountain Labora- tory, A. oblong oguttatum occurs from Sinaloa, Mexico to Brazil. The species 198 ECTOPARASITES OF PANAMA strongly resembles A. cajennense (Fabricius) and A. tapirellum Dunn but is separated by the characters given in the keys. Amblyomma ovale Koch Amblyomma ovale Koch, 1844, Arch. Naturg., 10: 227. Robinson, 1926, Ticks, pt. 4, pp. 25-29, figs, (with synonyms including A. fossum Neumann, 1899, and A. striatum Koch, 1844). Vogelsang and Dias, 1953, Revista Med. Vet. Parasit., Caracas, 12, (1-4), pp. 70-74, figs, (with synonyms including A. fossum). Dias, 1958, An. Inst. Med. Trop., 15, (2), p. 507 (with A. fossum as synonym). Aragao and Fonseca, 1961, Mem. Inst. Osw. Cruz, 59, (2) , pp. 131-148, figs, (detailed review of synonomy; redescribed). This tick ranges throughout the wetter forested regions of Panama at lower elevations, and we have one record from San Jose Island in Panama Bay. Dunn (1923, 1934) records specimens from Tamandua and tapir, while Fairchild (1943) believed dogs to be the most common host. We have examined 39 lots of adults from 16 different hosts, listed below in order of their abundance. One of the largest single lots, consisting of 17 males and 9 females, was from Nasua nasua, which also seems to be a favorite wild host. Over 60 percent of the recorded lots and over 80 percent of the total ticks were from carnivores. Nasua nasua (12 lots, 44 males, 15 females) ; dog (9 lots, 14 males, 10 females) ; man (7 lots, 5 males, 4 females) ; tapir (6 lots, 13 males, 11 fe- males) ; Felis pardalis (3 lots, 1 male, 2 females) ; Eira barbara (3 lots, 11 males, 4 females) ; horse (2 lots, 1 male, 1 female) ; Oryzomys sp. (2 lots, 2 males) ; Felis concolor (1 lot, 25 males, 1 female) ; Felis onca (1 lot, 2 males, 1 female) ; sloth (1 lot, 2 males) ; Felis yagouaroundi (1 lot, 1 male, 1 female) ; Procyon lotor (1 lot, 1 male, 1 female) ; and one specimen each from Procyon cancrivorus, armadillo, Galictis allamandi, and a bird, Arremonops conirostris. Engorged nymphs which subsequently molted to adults have been taken four times from Proechimys semispinosus, and once from Zygodontomys microtinus, so that rodents are perhaps important hosts for the earlier stages. Nymphs and larvae associated with adults on the same host, taken on several occasions, are not certainly ovale. Several authors refer to this species as A. fossum Neumann, 1899, de- spite the fact that Robinson (1926) compared numerous specimens of fossum from various places in South America with the types of ovale and found them to be the same. Dias (1958) examined the types of fossum and, in accordance with Robinson and Vogelsang and Dias (1953), found that this species is the same as ovale. Although Robinson also reduced striatum Koch, 1844, to a synonym of ovale, he noted that striatum differed somewhat but attributed this to variation. Most subsequent authors have regarded striatum as distinct, the most recent of these being Aragao and Fonseca (1961) who include fossum among the synonyms of ovale and recognize aureolatum Pallas, 1772, as the valid name for striatum. We have not seen specimens referable to aureolatum in our Panama material but a male from an unspecified locality in Darien, possibly within the present boundaries of Panama, was doubtfully determined by Neumann (1899) as striatum. FAIRCHILD, KOHLS AND TIPTON : TICKS 199 The range of ovale extends from Mexico to Argentina. A single male was removed from a dog on the Tama Indian Reservation in Iowa (Eddy and Joyce, 1942) but there is no evidence to suggest that the species is estab- lished there or elsewhere in the United States. Amblyomma pacae Aragao Amblyomma pacae Aragao, 1911, Mem. Inst. Osw. Cruz, 3, (2), pp. 170-172, figs. Robinson, 1926, Ticks, pt. 4, pp. 209-211, figs. Aragao and Fonseca, 1953, Mem. Inst. Osw. Cruz, 51 : 490 (with A. nigrum Tonelli Rondelli, 1939 as synonym) . Fairchild (1943) tentatively recorded this species from Agouti paca, a determination subsequently confirmed by Kohls. Since then, in spite of the ubiquity and abundance of its hosts, only a few additional collections of this species have been secured, as follows: Almirante (Bocas del Toro), 26 Jan. 1950, V. J. Tipton and C. M. Keenan, from Agouti paca, 1 female; Volcan (Chiriqui), 15 June 1959, V. J. Tipton and C. M. Keenan, from Dasyprocta punctata, 3 males, 3 females; vicinity of Juan Mina (Canal Zone), Jan. 1962, GML, from Agouti paca, 1 female; Fort Kobbe (Canal Zone), 18 Aug. 1955, G. Field, from paca, 1 female; Panama, L. H. Dunn, T-12, no other data, 1 female; Timishik, Rio Teribe (Bocas del Toro), May 1962, GML, from Agouti paca, 1 female. In addition to Panama this species is recorded from Brazil, Paraguay and Colombia, and the Rocky Mountain Laboratory collection contains a male from Agouti paca, Humming Bird Highway, British Honduras, 25 Dec. 1953, Dr. F. Manolson. Amblyomma parvum Aragao Amblyomma parvum Aragao, 1908, Trab. Inst. Manguinhos, pp. 18-19 (reprint). Robinson, 1926, Ticks, pt. 4, pp. 37-38. Aragao and Fonseca, 1953, Mem. Inst. Osw. Cruz, 51: 491 [with A. curruca Schulze, 1936, erroneously as synonym. Actually, A. curruca — A, auricularium (Conil), 1878]. This appears to be a rare species in Panama. Dunn (1923) recorded it from deer (Odocoileus) and cotton rats (Sigmodon) and Fairchild (1943) added a single record from cattle. Review of some old material of Dunn's and a few additional lots taken more recently suggests that the species occurs mainly on the drier Pacific coast and that we do not know its favored host. Our scanty records follow : Parita (Herrera), 13 June 1931, L. H. Dunn, from cats, 3 females; Tumba Muerta (? Panama), 13 Sept. 1932, L. H. Dunn, from Tamandua, 1 female; Cocle Province, 3 Dec. 1942, H. S. Eakins, from cattle, 1 male, 1 female; Balboa (Canal Zone), 8 Dec. 1914, no host, American Museum of Natural History, 1 female; Paraiso (Canal Zone), May 1955, G. Field, from sloth, 1 female ; Fort Clayton (Canal Zone) , 7 Nov. 1960, C. E. Yunker, from man, attached, feeding, 1 female. Besides Panama, this species has been recorded from a wide variety of mammalian hosts from Venezuela, French Guiana, Brazil, and Argentina. In addition, the Rocky Mountain Laboratory collection contains a male and a female taken on man, Finca Santa Cristina (Escuintla), Guatemala, 10 Sept. 1948, H. T. Dalmat, and 3 males from Urocyon cineroargenteus 200 ECTOPARASITES OF PANAMA guatemalae, Hacienda Miramar (Nenton) , Guatemala, 18 Nov. 1948, L. de la Torre, Chicago Natural History Museum. Excellent illustrations of adults of this tick are given by Boero (1957) . Amblyomma pecarium Dunn Amblyomma pecarium Dunn, 1933, Parasitology, 25, (3) , pp. 356-358, figs. Three male paratypes in bad condition remain at Gorgas Memorial Laboratory labeled Miraflores (Canal Zone), 11 Apr. 1932, R. Isaacs, Dunn no. 859, from Pecari angulatus bangsi. We have seen the following material, all from collared peccary, or wild pig, very probably collared peccary : Almi- rante (Bocas del Toro) , 9 Feb. 1960, V. J. Tipton and C. M. Keenan, 1 male ; Rio Tuira, mouth of Rio Paya (Darien), 28 Feb. 1958, GML, 1 male; Juan Mina (Canal Zone) 29 March 1946, many males, females, and nymphs; Almirante (Bocas del Toro), 1 March 1937, United Fruit Company Medical Department, 2 males; Fort Clayton (Canal Zone), 13 Sept. 1954, G. Field, 18 males, 1 female; Barro Colorado Island (Canal Zone), 2 Feb. 1961, C. E. Yunker, 7 males, 4 females. The Rocky Mountain Laboratory collection contains a female from "peccary", Humming Bird Highway, British Honduras, 25 Dec. 1953, F. Manolson, and three lots from jungles of El Ocote, Ocozocautla (Chiapas), Mexico, Miguel Alvarez del Toro, with data as follows: 7 males, 1 female from Tayassu tajacu, 4 May 1946; about 40 adults from Tayassu pecari, 4 May 1946; and 19 males, 12 females from Mazama americana, 25 May 1950. Whether this last lot is actually from a deer seems questionable since all other collections of pecarium are from peccaries. Amblyomma pictum Neumann Amblyomma pictum Neumann, 1906, Arch. Parasit., 10, (2), pp. 204-206. Robinson, 1926, Ticks, pt. 4, pp. 238-240 (with A. conspicuum Aragao, 1913, as synonym). A singe female which appears to be this little-known species was found on an anteater, Myrmecophaga tridactyla, Rio Mandinga (San Bias), 25 May 1957, P. Galindo. This species has been previously reported from Brazil off Myrmecophaga tridactyla ( = Myrmecophaga jubata) and dog; from Argentina off dog though not included in Boero's (1957) list; and from French Guiana off Tamandua sp. The Rocky Mountain Laboratory collec- tion contains one of two males from giant anteater, upper New River (trib- utary of Courentyne), southern British Guiana, Sept. 1938, E. R. Blake, Chicago Natural History Museum. The dentition of the hypostome of these males is %, not % as stated by Neumann for this species. Aragao (1913) gave it as % in his description of A. conspicuum; Floch and Fauran (1958) gave it as % for their single male from French Guiana. Amblyomma sabanerae Stoll Amblyomma sabanerae Stoll, 1890, Biol. Centr.-Amer., Arach., p. 23, figs, (female). Robinson, 1926, Ticks, pt. 4, pp. 182-183, figs. Schulze, 1937, Zeitsch. Parasitenk., FAIRCHILD, KOHLS AND TIPTON : TICKS 201 9, (6), pp. 692-694, fig. (male). Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 584. Dias, 1958, An. Inst. Trop. Med., 15, (2), pp. 496-497 (as synonym of A, humerale Koch). Dunn (1923) recorded as A. humerale Koch, 2 males, 2 females taken from a tortoise (which he believed to be Testudo tabulata) on the Boqueron River (Panama), H. C. Clark. Later (1934) he recorded as humerale a single adult, sex not stated, from a tapir. We have not seen Dunn's speci- mens from Testudo but we have a sabanerae male, without data, determined by Bishopp as humerale, which may well be the specimen Dunn reported from tapir. Recent examination of specimens of Geochelone (= Testudo) from the San Bias coast did not yield any ticks, though several sabanerae were taken from specimens of Geoemyda from the same area. Whether ticks from Geochelone in Panama will be humerale, crassum (q.v.) , or sabanerae, must await re-examination of Dunn's specimens or the collection of new material. We have taken A. sabanerae fairly frequently on turtles of the genus Geoemyda, less frequently on other turtles and on iguanas and once on an opossum. The ticks are often attached on the carapace around the margin where they leave characteristic pits in the shell, recognizable for a consider- able period after the ticks have detached. Males outnumber females about three to one in our collections, perhaps due to their remaining attached longer. Adults may attach on the shell or on the skin of head and appendages. Dr. John M. Legler informs us that Geoemyda annulata is largely terrestrial, G. funeria, Kinosternon spp. and Pseudemys scripta are aquatic, at least as adults, although they come on land to lay their eggs or when moving from one body of water to another. Most of our material has come from the vi- cinity of the Canal Zone, though the species appears to range throughout the country at low elevations, as we have material from Darien and Bocas del Toro Provinces. In the subjoined list, the lots under "turtle" and Geoemyda sp. probably are largely from G. annulata, the commonest terrestrial turtle in this area. Geoemyda annulata (6 lots, 23 males, 4 females, 1 larva) ; Geoemyda funeria (2 lots, 6 males, 3 females, 3 nymphs, 11 larvae) ; Geoemyda sp. (6 lots, 21 males, 3 females, 3 nymphs) ; turtle (12 lots, 40 males, 15 females, 4 nymphs, 76 larvae) ; Pseudemys scripta (juveniles, 2 lots, 2 males) ; Kinosteron sp. (4 nymphs) ; Iguana iguana (2 lots, 7 males, 1 female) ; Marmosa robinsoni (1 male) ; crawling on man (1 female). We have also seen 1 female and 2 nymphs from Los Diamantes, vicinity of Guapiles (Limon) , Costa Rica, 29 July 1961, J. M. Legler, from Geoemyda funeria (juvenile) ; 1 nymph, Rio Sucio, Quezaltepeque, El Salvador, 4 July 1961, J. M. Legler, from Geoemyda pulcherina; and the Rocky Mountain Laboratory collection has 3 records from Mexico as follows : 1 male, 1 female from Island of Cozumal, Quintana Roo, 16 July 1951, L. J. Stannard, from Geoemyda areolata; 2 males from Chichen Itza (Yucatan), 24 July 1937, B. W. Andrews, from Terrapene yucatana, and 1 male, 1 female from Naya- rit, Dec. 1955, J. Cook, from Geoemyda sp. The types, 2 females from Guatemala, were from "a small terrapin known to the natives by the name of 202 ECTOPARASITES OF PANAMA la Sabanera." Schulze described the male from specimens from Colombia without further data. Dias (1958) reduced A. sabanerae and A. crassum to synonyms of A. humerale but we regard them as valid species separable by characters given in the keys. Amblyomma tapirellum Dunn Amblyomma tapirellum Dunn, 1933, Parasitology, 25, (3), pp. 353-355, figs.; 1934, Jour. Parasit., 20, (5), p. 312. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), p. 586. Kohls, 1958, Jour. Parasit., 44, (4), pp. 431-432. This species was described from tapir, and we have taken it from col- lared peccary, giant anteater and horse as well. The bulk of our material, however, consists of specimens found crawling on man or on the ground. It appears to be quite strictly limited to forest areas below about 2500 feet, mostly from the Canal Zone eastward into Darien Province ; we have only one record from western Panama (La Vaca, Chiriqui Province, 18 Feb. 1930, Dunn, from peccary, 3 males, 1 female). We did not take it on several tapirs examined in the highlands of Chiriqui and Bocas del Toro. To a certain extent it seems to replace A. cajennense as a human parasite in for- ested areas such as Barro Colorado Island, from which we have 27 lots of tapirellum totaling 101 specimens, but only 8 of cajennense, totaling 18 specimens. We have taken it only once from horse, in that case a pack animal used on a jungle field trip, and have secured no material from cattle or deer, in marked contrast to cajennense. We list our material below, by numbers of lots and specimens. Free on ground or vegetation (11 lots, 11 males, 21 females) ; man, on clothing (14 lots, 22 males, 32 females) ; collared peccary (5 lots, 9 males, 5 females) ; tapir (5 lots, 180 males, 36 females, 1 nymph) ; man, attached (1 lot, 1 male) ; Myrmecophaga tridactyla (1 lot, 3 males, 2 females) ; horse (1 lot, 5 males, 6 females) ; bat (Carollia perspicillata) (1 lot, 1 male) ; no recorded host (16 lots, 20 males, 31 females plus two large lots not counted). The specimen from bat is probably a stray from the collector. One of the large lots with no host data is labeled simply "T-l", and is from Dunn's collecting. It may well be part of the paratype series. Dunn (1933) described the species from an adult tapir collected at Summit (Canal Zone) and mentions additional specimens from two young tapirs taken subse- quently, but the dates of neither collection are given. Later (1934), he discusses the ticks taken from three tapirs, an adult killed at Summit (Canal Zone) , and two young specimens from Aguas Buenas (Panama) , again with no dates given. In the later paper, A. tapirellum is listed, with a reference to the earlier paper. We suspect that both references allude to the same lots of ticks, and that T-l refers to "Tapir No. 1" of the 1934 paper, the adult tapir from Summit. Data of the Rocky Mountain Laboratory extend the range of A. tapirel- lum to Nicaragua, Colombia and Venezuela and add cattle to the list of hosts parasitized by this tick. FAIRCHILD, KOHLS AND TIPTON I TICKS 203 Amblyomma varium Koch Amblyomma varium Koch, 1844, Arch. Naturg., 10: 224 (male). Neumann, 1899, Mem. Soc. Zool. Fr., 12: 246-247 (redescription of male and description of var. albida, Chile) ; 1901, Mem. Soc. Zool. Fr., 14: 304-305 (female). Robinson, 1926, Ticks, pt. 4, pp. 205-209, figs. Amblyomma gertschi Cooley and Kohls, 1942, Pub. Hlth. Kept., 57, (46), pp. 1733- 1735, figs. Fairchild, 1943, Amer. Jour. Trop. Med., 23, (6), pp. 584-585. New synonymy. Robinson (1926) records specimens from Panama off Choloepus, as does Dunn (1923) . In the adult stage, this species, like geayi, appears to be restricted mainly to sloths but is less abundant. Both species appear to be slightly more abundant on Bradypus than on Choloepus. All material we have seen, except one female from Isla Bastimentos (Bocas del Toro) , has come from the Canal Zone or nearby Panama, as fol- lows: from Choloepus hoffmanni, two-toed sloth (16 lots, 30 males, 9 fe- males) ; Bradypus infuscatus, three-toed sloth (12 lots, 30 males, 9 females) ; from sloth (4 lots, 10 males, 2 females) ; no host (1 lot, 5 males) ; from "Eaton's opossum," probably Didelphis m. etensis (1 female). This species, originally described from specimens from Brazil, has been recorded from Nicaragua, Panama, Colombia, Venezuela, the Guianas, Argentina, and Chile (var. albida}. Vargas (1955) included it in his list of species occurring in Mexico, but its principal hosts, Bradypus and Choloe- pus, do not range that far northward and we doubt its presence there. The Rocky Mountain Laboratory collection contains a male and female (and reared larvae) from a sloth from near San Jose, Costa Rica, March 1962, J. J. Shaw; and a male said to be from "wild pig", Sheshea River basin at the headwaters of the Peruvian Amazon, Peru, 1960 or 1961, G. E. Dickin- son, received from Dr. R. E. Ryckman, Loma Linda University, Loma Linda, California. Dunn (loc. cit.) noted an engorged female of varium that weighed over 5 grams, the largest tick he had seen from Panama. We have two engorged females which weigh 6.4 grams each. Floch and Fauran (1958) had a speci- men from French Guiana measuring 32 mm. long, 30 mm. wide and weighing 7.5 grams. All males of A. varium that we have seen differ from the type of var. albida (Robinson, 1926, fig. 100) in being more broadly oval and in having long stout cornua as shown in fig. 2 of Cooley and Kohls (1942). Some males also have a long rather than a short spur on coxa IV. The dentition of the female hypostome is usually arranged % rather than % as stated by Robinson. In all males that we have seen it is % as given by all previous authors, except Floch and Fauran (1958) who stated that it is %• Host-Parasite Relationships We have secured ticks from most of the larger mammals that are known to occur in Panama, as well as from a number of birds and reptiles. Specific identifications of the larger host mammals are reasonably accurate ; we have not attempted to identify subspecies. Many of the smaller rodents have been identified to species, but in some cases only a guess as to their generic 204 ECTOPARASITES OF PANAMA identity has been possible. Many of the specimens of ticks have been ac- companied by only the common name of the host. Names such as "tapir," "ocelot," "jaguar," and "armadillo" permit identification to species with fair certainty. Others such as "sloth," "wild boar," "wild turkey" pose dif- ficulties, as two or more species are referable to these names. Names such as "bird," "lizard," "turtle," "wild rat" hardly permit conjecture. A host-parasite list is given below. The scientific names of the mammals follow Charles O. Handley's checklist, which appears elsewhere in this volume. Most of the birds have been determined by Dr. Alexander Wetmore. The turtles were identified by Dr. John M. Legler, the other reptiles by var- ious individuals. In the list, following the scientific name of each host we have given : (1) the number of specimens of the host from which ticks have been taken ; (2) the species of ticks in order of their frequency on the host, followed by a number in parentheses indicating the number of individual host specimens that were found infested with that species. Many lots of ticks either were not accompanied by host determinations or the host was merely listed as "mouse," "bird," "lizard," etc. These have not been listed below, but they are included under the discussion of each species of tick. Table 5 summarizes the information in the host lists. The ticks are listed by genera, with the number of species in each genus. The mammalian hosts are listed by order. The number of genera from which ticks have been secured is indicated for each order. For birds and reptiles we have shown the number of orders and genera from which ticks have been taken. Data for ticks from Amphibia (Bufo marinus) are not shown. The numbers in table 5 indicate the number of determined species of tick in each genus that have been taken from hosts of each order or class of animal. "A" indicates adults, "NL" nymphs and/or larvae. A query ( ?) indicates that nymphs or larvae probably belonging to the indicated genus, but not determined spe- cifically, were secured from the indicated order. In the case of nymphs and larvae, it is probable that many more species were taken than indicated, but in many cases larvae and nymphs are not determinable to species, and were not always determined to genus. Certain associations are apparent from this table. The Marsupialia are hosts to but two genera of ticks, the Edentata to but one, yet both orders are well represented in species and individuals in Panama. In Panama, the Chiroptera exceed all other orders of mammals in number of genera and species and yet they are true hosts to ticks of only two genera. The scattered records of other genera from bats probably represent strays or errors of association. The Primates, with the exception of man, are rarely attacked by ticks. We have never found ticks on wild primates in Panama ; all of our records are from animals in captivity. The Lagomorpha are represented in Panama only by one species of Syl- vilagus. These rabbits are infested by two species of ticks, Ixodes pomerantzi and Haemaphysalis leporispalustris, both apparently limited to Sylvilagus in Panama. Larvae of several other genera have been taken occasionally on FAIRCHILD, KOHLS AND TIPTON : TICKS 205 TABLE 5. HOST PREFERENCES OF PANAMANIAN TICKS (by genera) (See text for detailed explanation.) Genera of ticks and number of species in each Host groups and numbers of gen- era from which ticks were col- lected : Marsupialia: 6 Insectivora: 1 Chiroptera : 14 Primates (man) Edentata: 7 Lagomorpha: 2 Rodentia: 17 Carnivora: 11 Perissodactyla : 2 Artiodactyla : 5 Reptilia: 15 (from 3 orders) Aves: 26 (from 8 orders) •e a. o o 05 5/1 1/1 /I I/ A/NL A/NL A/NL A/NL A/NL A/NL A/NL A/NL A/NL A/NL 6/2 1/1 3/4 /? .. /I 2/ .. 2/ I/ .. 2/ 6/ 1 9 / /I 1/1 .. /? I/ .. 1/1 l^/ / /I 5/2 .. 1/1 I/ .. /2 6/ 3/2 .. I/ 1/1 .. /I 6/ 2/ 1/1 I/ .. 1/1 I/ 5/2 2/ 1/1 I/ 1/1 1/1 7/ 3/: 3/1 A = adults; NL = nymphs and/or larvae; ?=Probable members of indicated genus, but of undetermined species these rabbits. The record for Rhipicephalus on Lagomorpha is based on the domestic rabbit and is doubtless due to a laboratory infestation. The Insectivora are poorly represented in Panama. Only undetermined tick larvae were secured from the few specimens of shrews examined. The Rodentia are hosts to thirteen species of ticks belonging to six genera. However, probably the only species whose adults regularly parasi- tize rodents are those of Ixodes and Amblyomma, while only the earlier instars of species of the other genera listed parasitize these hosts. Again, the record of Rhipicephalus refers to a rodent (capybara) kept in heavily in- fested quarters in the laboratory. The Carnivora are hosts to adult ticks of eleven species belonging to four genera; an additional genus is recorded only from an early instar. Rhipicephalus has been taken only on domestic dogs and cats in Panama and does not belong to the native fauna. 206 ECTOPARASITES OF PANAMA The Perissodactyla in Panama include only the horse and tapir, but together these are hosts to eleven species of six genera of ticks. The horse is host to seven species representing three genera, the tapir to nine species belonging to four genera. Of these, Dermacentor, Ixodes, and Haema- physalis have been taken only on the tapir, Anocentor and Boophilus only on the horse. The Artiodactyla, with five genera positive for ticks, have yielded thirteen species belonging to six genera. We have not had the opportunity to examine the white-lipped peccary (Tayassu), the only large terrestrial mammal from which we have not secured ticks. The Reptilia, in Panama, are hosts to but two species of Amblyomma (three, if A. crassum actually occurs here), one on turtles and the other on a wide variety of lizards and snakes, as well as on the common toad, Bufo marinus. We have not examined any of the local Crocodilia. These are almost exclusively aquatic and not likely to be infested by ticks. The Aves are rarely hosts to adult ticks. Probably only Argas persicus and Ixodes brunneus regularly infest birds in this stage. Birds are, however, frequently attacked by larval ticks of at least one and probably several species of Amblyomma. Search of a few nests of birds — such as woodpeckers and toucans, which utilize treeholes — has so far not yielded any ticks. Host Specificity It is exceptional to find adults of any Panamanian species of ticks (ex- clusive of Argasidae) , parasitizing hosts of more than one class of verte- brates. Amblyomma dissimile apparently is the only tick to do so regularly, since it attacks species of both Amphibia and Reptilia. Adults of A. dis- simile and A. cajennense have also been taken occasionally on birds. It is very doubtful that these hosts are satisfactory or that they are regularly parasitized by these ticks. Most Panamanian Ixodidae do not regularly parasitize hosts of more than one order of vertebrates, though there are exceptions. Among those species of which we have adequate material, two species of Ixodes regularly parasitize mammals of more than one order. Another species has been taken once on a small marsupial, but otherwise only on rodents, while the remaining species appear to be confined to hosts of a single order. Anocentor nitens and Boophilus microplus parasitize both Artiodactyla and Perissodactyla, while Rhipicephalus sanguineus is almost restricted to dogs. It is probable that all three species of Dermacentor regularly parasitize species of but a single order of mammals. However, all have been taken as strays on members of other orders. Each of the two species of Haemaphysalis also appear to be confined to hosts of a single order, though H. leporispalustris , which is normally restricted to Lago- morpha, has been secured once from a rodent. In the dominant genus Amblyomma, with nineteen species in Panama, host selectivity varies greatly. Species such as cajennense and oblongo- guttatum occur on hosts of seven or eight different orders, while longirostre and pecarium are known from only a single species of host. The six species parasitizing Edentata are of some interest. Two, geayi and varium, are practically confined to the almost entirely arboreal sloths. Three others — FAIRCHILD, KOHLS AND TIPTON : TICKS 207 calcaratum, pictum, and nodosum — parasitize the less arboreal anteaters. There are two records of calcaratum from sloths. The last species, auricular- ium, is primarily a parasite of the burrowing armadillos, though it is often taken on anteaters and occasionally on hosts of other orders. Our information on the hosts of the early instars of Panamanian ticks is meager. Most species of Ornithodoros can now be determined in the larval stage, but our collections of larvae, other than of species that occur on bats, are limited. In the case of the latter, there is some indication of host preference, but whether this is due to the environmental necessities of the essentially free-living nymphs and adults, or to true host selection by the larvae is not clear from our scanty data. In the case of the Ixodidae, it has been possible to determine the larvae of Haemaphysalis and of a few species of Amblyomma. It is usually possible to place larvae of the other genera in the correct genus, though we have not critically studied all the numerous lots consisting only of nymphs and larvae. We have secured adults from engorged nymphs of numerous species. In most cases these nymphs have been taken from hosts different from those generally favored by the adults. Thus, of five reared nymphs of A. auricu- larium, three were from Philander (a marsupial) , and two from Sigmodon (a rodent). The adults are usually found on edentates (armadillos and anteaters) . All but one of the adults of H. juxtakochi obtained were from deer (Artiodactyla) but no early stages were taken from this host. On the other hand, nymphs of this tick were taken from three other species of mam- mals, two of which belong to other orders (Rodentia, Carnivora). The case of A. longirostre, a specific parasite of the lowland porcupine, Coendou rothschildi, has been noted previously by others (Aragao, 1936). We also have secured nymphs of this species, but only from birds, mostly passerines. It is almost certain that the sloth ticks, and undoubtedly other species whose early instars are seldom or never taken on the host with adults, have larval and nymphal hosts far different from those of the adult. Work on the taxonomy of the early instars is an absolute prerequisite to the solving of these problems. Abstract Forty-seven species of ticks belonging to 10 genera are recorded from Panama. Keys to genera and species are given, and host and locality records for each species are de- tailed. Extensive host lists and comments on the apparent host preferences of the genera and species are given. The altitudinal and climatic preferences of the various species of Ixodoidea are also discussed and tabulated. No new species are reported, but the follow- ing 12 species are new for the fauna of Panama : Antricola mexicanus Hoffmann, Ornitho- doros puertoricensis Fox, O. viguerasi Cooley and Kohls, Ixodes brunneus Koch, /. lasallei Mendez and Ortiz, I. loricatus Neumann, /. pomerantzi Kohls, /. tapirus Kohls, /. venezuelensis Kohls, Dermacentor halli Mclntosh, D. imitans Warburton, and Am- blyomma pictum Neumann. Ixodes calif ornicus Banks is synonymized under I. brunneus Koch, and /. scuticrenatus Vazquez under /. luciae Senevet. Amblyomma avecolens Cooley and Kohls and A. curruca Schulze are reduced to synonyms of A. longirostre Koch and A. auricularium (Conil), respectively. A. gertschi Cooley and Kohls is synonymized under A. varium Koch. 208 ECTOPARASITES OF PANAMA HOST-PARASITE LIST (N m nymph; L = larva) Class AMPHIBIA Order SALIENTIA Bufo marimis: 3 Amblyomma dissimile Koch (3) Class REPTILIA Order TESTUDINATA Geoemyda annulata: 6 Amblyomma sabanerae Stoll (6) Geoemyda funeria: 3 Amblyomma sabanerae Stoll (2) sp., NN and LL (2) Pseudemys scripta: 3 Amblyomma sabanerae Stoll (2) dissimile Koch (1) Kinosternon sp. : 1 Amblyomma sabanerae Stoll Order SQUAMATA Iguana iguana: 18 Amblyomma dissimile Koch (16) sabanerae Stoll (2) Basiliscus basiliscus: 1 Amblyomma sp., LL Ameiva ameiva: 7 Amblyomma dissimile Koch (1) " sp., NN and/or LL (6) lizards, probably Ameiva spp. : 17 Amblyomma sp., NN and/or LL (17) Order SERPENTES Constrictor constrictor: 3 Amblyomma dissimile Koch (3) Epicrates sp. : 1 Amblyomma sp., NN and LL Spilotes pullatus: 1 Amblyomma dissimile Koch Pseustes poecilonotus : 1 Amblyomma dissimile Koch Chironius carinatus: 1 Amblyomma dissimile Koch Oxybelis sp. : 1 Amblyomma sp., N Thalerophis richardi: 1 Amblyomma sp., LL Bothrops atrox: 4 Amblyomma dissimile Koch (4) Lachesis muta: 3 Amblyomma dissimile Koch (3) Class AVES In addition to the birds listed by name below, we have a considerable number of additional lots, all Amblyomma nymphs or larvae, from unidentified birds. Thus, birds seem to be important hosts for the early stages of Amblyomma. The few records of adult Amblyomma from birds appear to be strays. Order TINAMIFORMES Crypturellus soui: 1 Amblyomma sp., L "Tinamou": 1 Ixodes brunneus Koch Order CICONIIFORMES Cochlearius cochlearius: 1 Amblyomma dissimile Koch Order FALCONIFORMES Buteo magnirostris: 1 Amblyomma cajennense (Fabricius) Spizaetus tyrannus: 1 Amblyomma sp., N Order GALLIFORMES Crax rubra: 2 Amblyomma oblongoguttatum Koch sp., LL Penelope purpurascens: 1 Amblyomma sp., LL domestic fowl: 2 Amblyomma sp., NN and LL chicken coops: 1 Argas persicus (Oken) FAIRCHILD, KOHLS AND TIPTON : TICKS 209 Order CUCULIFORMES Neomorphus geoffroyi salvini: 1 Argas persicus (Oken), N Order CORACIIFORMES Chloroceryle americana: 1 Amblyomma sp., LL Order PICIFORMES Malacoptila panamensis: 2 Amblyomma longirostre (Koch),NN Capito maculicoronatus : 1 Amblyomma sp., N Ramphastos sp. : 1 Amblyomma sp., N Order PASSERIFORMES Xiphorhynchus sp. : 1 Amblyomma longirostre (Koch), N Cymbilaimus lineatus: 1 Amblyomma longirostre (Koch),N Thamnophilus nigriceps: 1 Amblyomma sp., N Rhytipterna holerythra: 1 Amblyomma sp., N Querula purpurata: 1 Amblyomma longirostre (Koch), N Cacicus uropygialis microrhynchus : 2 Amblyomma longirostre (Koch), N Icterus chrysater: 1 Amblyomma longirostre (Koch), N Ramphocelus passerinii: 8 Amblyomma sp., NN Tachyphonus rufus: 1 Amblyomma sp., N Eucometis penicillata: 1 Amblyomma sp., L Saltator maximus: 1 Amblyomma longirostre (Koch), N Saltator albicollis: 1 Amblyomma longirostre (Koch), N Sporophila aurita corvina: 9 Amblyomma sp., NN Arremonops conirostris: 1 Amblyomma ovale Koch, 2 , engorged Class MAMMALIA Order MARSUPIALIA Family Didelphidae Caluromys derbianus: 4 Amblyomma geayi Neumann (1) sp., NN and/or LL (3) Monodelphis adusta: 1 Ixodes venezuelensis Kohls Marmosa robin son i: 2 Ixodes luciae Senevet, N (1) Amblyomma sabanerae Stoll (1) Philander opossum: 24 (Early records from "Philander" generally refer to Caluromys derbianus, q.v.) Ixodes luciae Senevet (6) Amblyomma auricularium (Conil) (3) geayi Neumann (1) sp., NN and/or LL (22) Metachirus nudicaudatus: 2 Ixodes loricatus Neumann (1) " sp., NN (1) Didelphis marsupial is: 43 Ixodes luciae Senevet (20) " boliviensis Neumann (3) " affinis Neumann (2) " sp., NN and/or LL (3) Amblyomma auricularium (Conil) (2) Amblyomma cajennense (Fabricius) (2) geayi Neumann (1) varium Koch (1) sp., NN and/or LL (20) Chironectes minimus: 1 Amblyomma oblong oguttatum Koch Order INSECTIVORA Family Soricidae Cryptotis nigrescens: 2 Dermacentor sp., L ? Ixodes sp., L Order CHIROPTERA Infestations of bats with ticks other than Ornithodoros were probably accidental. Many of the bats were caught in mist nets and rested on or near the ground for some time. Here they could easily be infested with larval ticks of other genera. In addition to the following, we have a number of records of ticks from unidentified bats. Family Emballonuridae Peropteryx macrotis: 2 Ornithodoros azteci Matheson, LL 210 ECTOPARASITES OF PANAMA Family Noctilionidae Noctilio labialis: 7 Ornithodoros hasei (Schulze), NN and LL (7) Noctilio leporinus: 3 Ornithodoros hasei ( Schulze ),LL (3) Family Phyllostomidae Pteronotus parnellii: 1 Ornithodoros viguerasi Cooley and Kohls, LL Pteronotus psilotis: 1 Antricola mexicanus Hoffmann, LL Amblyomma sp., L Pteronotus sp. : 1 Ornithodoros viguerasi Cooley and Kohls, L Lonchorhina aurita: 2 Ornithodoros azteci Matheson, LL Tonatia silvicola: 1 Ornithodoros hasei (Schulze), L Trachops cirrhosus: 3 Ornithodoros brodyi Matheson, LL (3) hasei (Schulze), L (1) Carollia perspicillata: 4 Ornithodoros brodyi Matheson, LL (3) Amblyomma tapirellum Dunn (1) Uroderma bilobatum: 1 Ornithodoros hasei (Schulze), LL Vampyrops helleri: 2 Amblyomma cajennense ( Fabricius ),$ sp., N Ornithodoros hasei (Schulze), LL Chiroderma salvini: 1 Amblyomma sp., N Artibeus sp. : 1 Ixodes sp., L Family Desmodidae Desmodus rotundus: 2 Ornithodoros azteci Matheson, LL (1) " brodyi Matheson, L (1) Family Vespertilionidae Myotis nigricans: 3 Dermacentor halli Mclntosh, N Antricola mexicanus Hoffmann, LL (3) Family Molossidae Molossus sp. : 1 Ornithodoros hasei (Schulze), L Order PRIMATES With the exception of man, ticks are very seldom found on Primates in Panama. We have records of Amblyomma nymphs and/or larvae from one individual each of Alouatta villosa, Cebus capuchinus, and Aotus trivirgatus. Two Saguinus geoffroyi have yielded ticks, Rhipicephalus sanguineus (Latreille) in one case, and Amblyomma larva in the other. Family Hominidae Homo sapiens: 54 Our records do not generally indicate whether the ticks were attached or merely crawling on the skin or clothing. Larvae of Amblyomma species are a great pest in many areas, especially during the dry season. Amblyomma tapirellum Dunn (15) cajennense (Fabricius) (13) naponense (Packard) (1) oblong oguttatum Koch. (9) ovale Koch (7) parvum Aragao (1) sabanerae Stoll (1) Sp., NN (11) Ixodes boliviensis Neumann (4) Rhipicephalus sanguineus (Latreille) (2) Dermacentor latus Cooley (2) imitans Warburton (1) Order EDENTATA Family Myrmecophagidae Myrmecophaga tridactyla: 1 Amblyomma calcaratum Neumann nodosum Neumann oblong oguttatum Koch tapirellum Dunn pictum Neumann sp., NN Tamandua tetradactyla: 30 Amblyomma calcaratum Neumann (20) nodosum Neumann (19) auricularium (Conil) (12) oblong oguttatum Koch (7) cajennense (Fabricius) (3) naponense (Packard) (1) parvum Aragao (1) sp., NN and/or LL (1) Cyclopes didactylus: 2 Amblyomma sp., NN and/or LL (2) FAIRCHILD, KOHLS AND TIPTON : TICKS 211 Family Bradypodidae Bradypus infuscatus: 35 Amblyomma geayi Neumann (35) varium Koch (12) sp., NN and/or LL (7) Choloepus hoffmanni: 25 Amblyomma varium Koch (16) geayi Neumann (12) calcaratum Neumann (2) oblong oguttatum Koch (1) Family Dasypodidae Cabassous centralis: 1 Amblyomma auricularium (Conil) Dasypus novemcinctus : 25 Amblyomma auricularium (Conil) (22) cajennense ( Fabricius ) (1) oblongoguttatumKoch (1) ovale Koch (1) sp., NN and/or LL (2) Ixodes sp., L (1) Order LAGOMORPHA Sylvilagus brasiliensis : 14 Haemaphysalis leporispalustris (Packard) (14) Ixodes pomerantzi Kohls (3) Dermacentor sp., N (1) Amblyomma sp., N (1) Ornithodoros puertoricensis Fox, L (1) Oryctolagus cuniculus: 3 Rhipicephalus sanguineus (Latreille) (3) Order RODENTIA Family Sciuridae Sciurus granatensis: 23 Ixodes tiptoni Kohls and Clifford (14) sp., probably tiptoni, LL (3) Amblyomma sp., NN and/or LL (6) Family Heteromyidae Liomys adspersus: 20 Amblyomma sp., LL (20) Heteromys sp. : 5 Amblyomma sp., NN and/or LL (3) Dermacentor sp., NN (2) Family Cricetidae Oryzomys spp. : 21 Amblyomma ovale Koch (2) sp., NN and/or LL (13) Ixodes luciae Senevet (1) Ixodes venezuelensis Kohls (1) " sp., NN and/or LL (4) Dermacentor sp., LL (1) Reithrodontomys creper and spp. : 12 Dermacentor sp., NN and/or LL (8) Amblyomma sp., LL (3) Ixodes sp., N (1) Peromyscus spp. : 23 Dermacentor sp., NN and/or LL (11) Amblyomma sp., NN and/or LL (7) Ixodes sp., NN and/or LL (5) Haemaphysalis leporispalustris (Packard), L (1) Zygodontomys microtinus: 16 Ixodes venezuelensis Kohls (4) luciae Senevet, N Amblyomma ovale Koch, N (1) " sp., NN and/or LL (11) Scotinomys xerampelinus: 4 Ixodes sp., LL (2) Dermacentor sp., L (1) ? Amblyomma sp., L (1) Sigmodon hispidus: 40 Amblyomma auricularium (Conil), N (2) sp., N and/or LL (38) ? Ixodes sp., L (1) Family Muridae Rattus sp. : 1 Ornithodoros puertoricensis Fox, LL Family Erethizontidae Coendou mexicanus: 5 Dermacentor halli Mclntosh (5) Ixodes boliviensis Neumann (4) " sp., N and LL (2) Coendou rothschildi: 10 Amblyomma longirostre (Koch) (8) sp., NN and/or LL (5) Haemaphysalis juxtakochi Cooley, N (1) Family Hydrochaeridae Hydrochaeris hydrochaeris : 2 Amblyomma auricularium (Conil) (1) Rhipicephalus sanguineus (Latreille) (1) Family Dasyproctidae Agouti paca: 10 Amblyomma pacae Aragao (6) coelebs Neumann (1) sp., NN (3) Ixodes lasallei Mendez and Ortiz (2) " sp., NN and/or LL (3) 212 ECTOPARASITES OF PANAMA Dasyprocta punctata: 14 Ixodes lasallei Mendez and Ortiz (8) " sp., NN (1) Amblyomma oblongoguttatum Koch (3) pacae Aragao (1) sp., NN and/or LL (5) Family Echimyidae Proechimys semispinosus: 52 Amblyomma ovale Koch, NN (4) sp., NN and/or LL (48) Ixodes sp., N (1) Haemaphysalis juxtakochi Cooley, N (1) Hoplomys gymnurus: 1 Amblyomma sp., L Order CARNIVORA Family Canidae Canis familiaris: 38 Rhipicephalus sanguineus (Latreille) (20) Amblyomma oblongoguttatum Koch (19) ovale Koch (9) cajennense (Fabricius) (5) auricularium (Conil) (1) " sp., NN and/or LL (5) Ixodes affinis Neumann (1) " boliviensis Neumann (15) Family Procyonidae Procyon lotor: 1 Amblyomma ovale Koch Ixodes boliviensis Neumann rubidus Neumann Procyon cancrivorus: 3 Amblyomma ovale Koch (1) oblongoguttatum Koch (1) naponense (Packard) (1) sp., NN and/or LL (2) Nasua nasua: 30 Amblyomma ovale Koch (12) oblongoguttatum Koch (10) auricularium (Conil) (1) cajennense (Fabricius) (1) naponense (Packard) (1) sp., NN and/or LL (8) Ixodes rubidus Neumann ( 1 ) boliviensis Neumann (3) " sp., NN and/or LL (2) Haemaphysalis juxtakochi Cooley (2) Potos flavus: 1 Amblyomma sp., N Bassaricyon gabbii: 1 Ixodes rubidus Neumann Family Mustelidae Mustela frenata: 1 Ixodes rubidus Neumann Eira barbara: 4 Amblyomma ovale Koch (3) oblongoguttatum Koch (1) Ixodes rubidus Neumann (1) Galictis allamandi: 1 Amblyomma ovale Koch sp., N Conepatus semistriatus : 1 Ixodes rubidus Neumann Lutra annectens: 1 Ixodes sp., L Family Felidae Felis concolor: 1 Amblyomma ovale Koch Felis onca: 5 Ixodes boliviensis Neumann (3) affinis Neumann (2) Amblyomma ovale Koch (1) Sp., NN Felis pardalis: 4 Ixodes affinis Neumann (3) Amblyomma ovale Koch (3) sp., NN andLL (1) Felis yagouaroundi: 1 Amblyomma ovale Koch Felis cattus: 4 Amblyomma parvum Aragao (2) oblongoguttatum Koch (1) sp., NN and/or LL (2) Rhipicephalus sanguineus (Latreille) (1) Ixodes boliviensis Neumann (1) Order PERISSODACTYLA Family Tapiridae Tapirus bairdii: 11 Amblyomma oblongoguttatum Koch (7) ovale Koch (6) coelebs Neumann (5) tapirellum Dunn (5) cajennense (Fabricius) (1) Sp., NN (2) Dermacentor latus Cooley (3) Ixodes boliviensis Neumann (3) tapirus Kohls (2) Haemaphysalis juxtakochi Cooley (1) FAIRCHILD, KOHLS AND TIPTON : TICKS 213 Family Equidae Equus caballus: 43 Amblyomma cajennense (Fabricius) (24) oblong oguttatum Koch (17) ovale Koch (2) tapirellum Dunn (1) coelebs Neumann (1) sp., NN (1) Anocentor nitens (Neumann) (7) Boophilus microplus (Canestrini) (3) (Numerous specimens of A. nitens collected by Field not included) Order ARTIODACTYLA Family Tayassuidae Tayassu tajacu: 19 Amblyomma naponense (Packard) (13) oblong oguttatum Koch (13) pecarium Dunn (7) tapirellum Dunn (5) cajennense (Fabricius) (1) sp., NN and/or LL (7) Dermacentor imitans Warburton (4) Haemaphysalis juxtakochi Cooley (3) Family Suidae Sus scrofa: 7 Anocentor nitens (Neumann) (7) Boophilus microplus (Canestrini) (3) Amblyomma oblong oguttatum Koch (3) cajennense (Fabricius) (2) " ovale Koch (2) Amblyomma tapirellum Dunn (2) coelebs Neumann (1) sp., NN (1) Family Cervidae Odocoileus virginianus: 21 Amblyomma oblong oguttatum Koch (18) cajennense (Fabricius) (1) Haemaphysalis juxtakochi Cooley (6) Ixodes affinis Neumann (4) boliviensis Neumann (1) Anocentor nitens (Neumann) (2) Mazama americana: 5 Ixodes affinis Neumann (3) Amblyomma oblongoguttatum Koch (2) calcaratum Neumann (1) sp., NN (1) Haemaphysalis juxtakochi Cooley (1) Family Bovidae Bos taurus: 33 Boophilus microplus (Canestrini) (23) Amblyomma cajennense (Fabricius) (20) oblongoguttatum Koch (15) parvum Aragao (1) sp., NN and/or LL (4) Anocentor nitens (Neumann) (5) Ixodes boliviensis Neumann (4) Capra hircus: 3 Amblyomma cajennense (Fabricius) (1) oblongoguttatum Koch (1) Boophilus microplus (Canestrini) (1) References ANASTOS, G., AND SMITH, C. 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Parasitology, 3: 408-416. NUTTALL, G. H. F., AND WARBURTON, C. 1911. Ixodidae, Section 2, The Genus Ixodes. In Ticks, a monograph of the Ixodoidea, by Nuttall, Warburton, Cooper and Robinson. Cambridge University Press. Pt. 2. pp. 133-348, 4 pis. OKEN, L. 1818. Sogenannte giftige Wanze in Persien. Isis, pp. 1567-1570, pi. 19, figs. 1-4. OSORNO-MESA, E. 1941. Las garrapatas de la Republica de Colombia. Anuaro Acad. Nac. Med., 1938- 1940: 398-434. Bogota. PACKARD, A. S. 1869. List of hymenopterous and lepidopterous insects collected by the Smithsonian Expedition to South America, under Prof. James Orten; appendix to report on Articulates. Ann Rept. Peabody Acad. Sci., pp. 56-69. RlNGUELET, R. 1947. La supuesta presencia de Ixodes brunneus Koch en la Argentina y descripcion de una nueva garrapata Ixodes neuquenensis nov. sp. Notas Mus. La Plata, 12: 207-216. ROBINSON, L. E. 1926. The genus Amblyomma. In Ticks, a monograph of the Ixodoidea, by Nuttall, Warburton, Cooper and Robinson. Cambridge University Press. Pt. 4. pp. xii + 302, 7 pis. DE RODANICHE, E. 1953. Natural infection of the tick Amblyomma cajennense with Rickettsia rickettsii in Panama. Amer. Jour. Trop. Med. Hyg., 2: 696-699. SCHULZE, P. 1935. Zur vergleichenden Anatomic de Zecken. (Das Sternale, die Mundwerkzeuge, Analfurchen und Analbeschilderung und ihre Bedeutung, Ursprunglichkeit und Luxurieren. Zeitschr. f. Morph. u. Okol. der Tiere, 30: 1-40. 1936. Neue und wenig bekannte Amblyommen und Aponommen aus Afrika, Siida- merika, Indien, Borneo und Australien. (Ixodidae.) Zeitschr. f. Parasitenk., 8, (6), pp. 619-637. FAIRCHILD, KOHLS AND TIPTON : TICKS 219 1937. Beitrage zur Kenntnis der Zeckengattung Amblyomma. Ibidem, 9, (6), pp. 690-694. . Anocentor columbianus n. g. n. sp. (Ixod.). Zool. Anz., 120, (1-2), pp. 24-27. 1941. Das Geruchsorgan der Zecken. Untersuchungen iiber die Abwandlungen eines Sinnesorgans und seine stammesgeschichtliche Bedeutung. Zeitschr. f. Morph. u. Okol. der Tiere, 37: 491-564. SENEVET, G. 1940. Quelques Ixodides de la Guyane franchise. Especies nouvelles d'lxodes et d'Amblyomma. VI Congr. Intern. Ent. pp. 891-898, Madrid (1935). STOLL, 0. 1886-1893. Arachnida Acaridea. Biologia Centrali-Americana, Zoologia. London, pp. v-xxi, 1-55, 21 plates. VARGAS, L. 1955. Relacion del papel patogeno de las garrapatas y lista de las especies Mexicanos. Gaceta Med. de Mexico, 85, (4-5), pp. 489-502. VAZQUEZ, L. 1946. Ixodes scuticrenatus, una especie nueva de garrapata de Mexico. An. Inst. Biol., 17, (1-2), pp. 237-245. VOGELSANG, E. G., AND BIAS, J. A. T. SANTOS. 1953. Nueva contribucion al estudio de la fauna ixodologica en Venezuela. Rev. Med. Vet. Parasit., 12, (1-4), pp. 63-89. WARBURTON, C. 1933. On five new species of ticks (Arachnida, Ixodoidea). Parasitology, 24, (4), pp. 558-568. The Chiggers of Panama (Acarina : Trombiculidae) JAMES M. BRENNAN l AND CONRAD E. YuNKER2 Ewing (1925) described the first chigger from Panama. Fairchild (1943) prepared the first list in which he recorded eight species and was followed by Wharton and Fuller (1952) who recorded 12 species. Both in- cluded Trombicula cavernarum Ewing, 1933 and T. trifurca Ewing, 1933, known only as adults and therefore not further considered here. Brennan and Jones (1961a) added 17 new species and three new genera. The present report concerns large numbers of chiggers collected in Pan- ama (most after 1954 and the great majority from 1960 to 1962), from nearly 5000 vertebrate hosts of about 70 mammalian, 50 avian and a few reptilian species. Seventy-six species of chiggers, distributed among 29 genera, are recorded. Five of these genera and 16 species are described as new. Several other undescribed species and new genera have been rec- ognized, but are not included because of inadequate material. The active support and cooperation of the following individuals and their organizations are acknowledged : Major Vernon J. Tipton and Mr. Charles M. Keenan, Preventive Medicine Division, United States Army Caribbean ; Dr. Graham B. Fairchild and Mr. Pedro Galindo, Gorgas Memorial Labora- tory ; Dr. Nathan Gale, Veterinary Division, Canal Zone Government. They assisted the authors, either directly or indirectly, in obtaining permission to collect in the Canal Zone and the Republic of Panama, by arranging field trips, by providing transportation via land, air, and water, by provid- ing various items of essential supplies and equipment, by generously lend- ing laboratory and field personnel, and by collecting many vertebrate hosts and their parasites. For specimens collected after May 1960, identifications of Panamanian *• 2 U. S. Department of Health, Education, and Welfare, Public Health Service, Na- tional Institutes of Health, National Institute of Allergy and Infectious Diseases, Rocky Mountain Laboratory, Hamilton, Montana, and Middle America Research Unit, Ancon, Canal Zone. 221 222 ECTOPARASITES OF PANAMA reptiles were by Mr. Hymen Marx, Chicago Natural History Museum; of birds by Dr. Alexander Wetmore, Smithsonian Institution; of mammals by Dr. Charles 0. Handley, United States National Museum. We are un- able to credit accurately host determinations prior to 1960, although Dr. Handley identified many of the mammals collected since 1957. Holotypes of all new species are deposited in the Rocky Mountain Lab- oratory (RML). Paratypes are deposited there and in the United States National Museum, the British Museum (Natural History), and the Chicago Natural History Museum, as indicated. If not otherwise indicated, collecting of new forms is credited to organ- izations: Environmental Health Branch, Preventive Medicine Division, United States Army Caribbean (EHB) ; Middle America Research Unit (MARU). KEY TO PANAMANIAN GENERA 1. Leg I with six segments; coxa I with two setae; spiracles and tracheae pres- ent; scutum with six setae (Leeuwenhoekiinae) 2 Leg I with seven segments; coxa I with one seta; spiracles and tracheae ab- sent ; scutum with three, four, five, or seven setae 3 2. Scutum with anteromedian projection; cheliceral blade with series of teeth Odontacarus Ewing Scutum without anteromedian projection ; cheliceral blade with tricuspid cap only Sasacarus Brennan and Jones 3(1). Scutum with a pair of anterosubmedian setae (Apoloniinae) . .Vargatula n. gen. Scutum with a single anteromedian seta (Trombiculinae) 4 4. Sensillae expanded 5 Sensillae flagelliform 18 5. With dorsal platelets in addition to scutum. . Polylopadium Brennan and Jones Without additional dorsal platelets 6 6. PL'S and most dorsal setae broad-foliate Cordiseta Hoffmann PL'S and dorsal setae not foliate 7 7. Sensillae with elongate flagelliform setules, parasitic on bats Perissopalla Brennan and White Sensillae without flagelliform setules 8 8. Cheliceral blades with tricuspid cap only or a single minute subapical dorsal tooth or hook 9 Cheliceral blades with one or a series of dorsal teeth plus tricuspid cap 16 9. Posterior margin of scutum obsolescent or absent; coxa II with two setae; intranasal habitat Kymocta Yunker and Brennan Posterior margin of scutum present; coxa II with one seta; habitat not intra- nasal 10 10. Tarsi with nude subapical setae; sensillae slightly expanded, but with large swollen setules, parasitic on bats Speleocola Lipovsky Tarsi without nude subapical setae; sensillae greatly expanded; setules not swollen 11 11. Integumental striae encroaching on posterior half of scutum Neoschoengastia Ewing Scutum without striae 12 12. Leg segmentation 7-7-7 ; ventral humeral setae absent 13 Leg segmentation usually 7-6-6, rarely 7-7-6 or 7-7-7 in which cases ventral humeral setae are always present, but may be present or absent if leg segmentation is 7-6-6 14 13. PL'S extrascutal Ascoschoengastia Ewing PL'S on scutum Euschoengastia Ewing BRENNAN AND YUNKER : CHIGGERS 223 14 (12) . Ventral humeral setae and parasubterminala present Pseudoschoengastia Lipovsky Ventral humeral setae and parasubterminala absent 15 15. PL'S on scutum; genualae II and III absent; intradermal habitat Intercutestrix n. gen. PL'S off scutum; genualae II and III present; habitat not intradermal Vanidicus Brennan and Jones 16(8). Cheliceral blade with one dorsal tooth in addition to tricuspid cap; genualae II and III present; habitat intranasal Blix n. gen. Cheliceral blade with a series of dorsal teeth; genualae II and III absent. ... 17 17. Ventral humeral setae present; cheliceral blade sharply curved; palpal tibial claw bifurcate; scutum subquadrate; intranasal habitat. . .Myxacarus n. gen. Ventral humeral setae absent; cheliceral blade nearly straight; palpal tibial claw trif urcate ; scutum much wider than deep ; habitat not intranasal .... Aniatrus Brennan and Jones 18(4). With seven scutal setae Hoffmannina Brennan and Jones With five scutal setae 19 19. PL'S off scutum Tecomatlana Hoffmann PL'S on scutum 20 20. Palpal tibial claw with a single prong 21 Palpal tibial claw with two or three prongs 23 21. Eyes large, 2/2, in a plate ; anterior scutal setae branched ; sensillae branched ; a mastitarsala III; habitat not intranasal Crotiscus Ewing Eyes absent or only a single reduced pair; anterior scutal setae nude; sensillae nude or with few vestigial barbs; no mastitarsala III; intranasal habitat, on bats 22 22. Scutum cuneiform; eyes absent; all leg segments with one or more elongate nude setae; palpal femur greatly enlarged Alexfainia Yunker and Jones Scutum subquadrate; eyes 1/1 ; leg segments without nude setae; palpal femur not enlarged Vergrandia Yunker and Jones 23(20). With a mastifemorala I; basifemora and telofemora semi-fused to fused; in- tranasal, on bats Perates Brennan and Dalmat Without a mastifemorala I; basifemora and telofemora not fused, but dis- tinctly articulated 24 24. Cheliceral blade with hood-like serrate distal expansion ; posterior margin of scutum with a sharp median tip; on bats Beamerella Brennan Cheliceral blade otherwise, usually with a subapical dorsal tooth or a tricuspid cap ; posterior margin of scutum without a tip, although it may be angulate . 25 25. Scutum cuneiform with a medially cleft posterior margin; all scutal setae nude; coxae II and III multisetose; intranasal habitat; on rodents Crotonasis n. gen. Scutum not cuneiform and without a cleft posterior margin; all scutal setae branched ; coxae II and III unisetose ; habitat not intranasal 26 26. Palpal claw bifurcate, accessory prong inner and ventral. . Eutrombicula Ewing Palpal claw normally trifurcate, but if bifurcate, accessory prong outer and dorsal 27 27. Mastitarsala III present; three genualae I; scutum deep, frequently pentag- onal, the AL'S never in anterolateral angles and usually set considerably behind margin; sensillae branched; palpal femoral seta branched; on birds Blankaartia Oudemans Without the above combination of characters 28 28. Mastitarsala III absent; two genualae I; scutum roughly rectangular, the AL'S always in anterolateral angles; sensillae branched; palpal femoral, genual, and tibial setae nude ; galeal seta branched Leptotrombidium Nagayo et al. Without the above combination of characters Trombicula Berlese 224 ECTOPARASITES OF PANAMA Genus Odontacarus Ewing Odontacarus Ewing, 1929, Man. Ext. Parasites, p. 188. Type-species: Trombicula dentata Ewing, 1925. KEY TO PANAMANIAN SPECIES Tarsus III with a tarsala chiapanensis (Hoffmann) Tarsus III without a tarsala fieldi Brennan and Jones Odontacarus chiapanensis (Hoffmann) Acomatacarus chiapanensis Hoffmann, 1948, Rev. Inst. Salub. Enferm. Trop., 9, (3), pp. 179-182, figs. 6-11. Twenty-nine specimens off (5) Proechimys semispinosus, Almirante (Bocas del Toro), 22 to 27 January and 16 July 1960. First Panamanian records. Odontacarus fieldi Brennan and Jones Odontacarus fieldi Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 105-106, fig. 1. Seventy specimens identified from 28 lots. Hosts: BIRDS, Neomorphus geoffroyi salvini, Odontophorus erythrops; MAMMALS, Didelphis marsu- pialis, Proechimys semispinosus, Liomys adspersus, Sciurus granatensis, Sigmodon hispidus, Zygodontomys microtinus, Felis pardalis. Localities : Cacao Plantation, Summit, and Miraflores (Canal Zone) ; Cerro Pirre (Dar- ien) ; Cerro Campana (Panama) ; Isla Bastimentos (Bocas del Toro) ; Cerro Hoya (Los Santos). Active throughout the year. See Brennan and Jones (1961a) for other Panamanian records. Genus Sasacarus Brennan and Jones Sasacarus Brennan and Jones, 1959, Ann. Ent. Soc. Amer., 52, (1), p. 8. Type-species : Chatia furmani Hoffmann, 1954. Sasacarus furmani (Hoffmann) Chatia furmani Hoffmann, 1954, Ann. Esc. Nac. Cienc. Biol. Mex., 8, (1-2), pp. 17-20, figs. 1-4. Sixty-eight specimens identified from 30 lots. Hosts: Didelphis mar- supialis, Proechimys semispinosus, Heteromys desmarestianus. Local- ities: Pina, Gamboa Road, Fort Gulick, and France Field, (Canal Zone) ; Cerro Azul (Panama) ; Bocas del Toro Province. First records for Panama. Sporadic collections, 1954 to 1962, November to April, suggest peak ac- tivity during the dry season. The Panamanian form differs from the typical form in that tarsala II is noticeably thicker and not longer than tarsala I. Vargatula, new genus Type-species : Vargatula hispida, new species. DIAGNOSIS : Apoloniine larvae with anterolateral, paired submedian, and displaced posterolateral scutal setae ; sensillae flagellif orm, branched. Che- liceral blades with minute dorsal tooth. Palpal tarsus with five branched BRENNAN AND YUNKER : CHIGGERS 225 setae, a subterminala, and a tarsala; tibial claw trifurcate. Eyes absent. No genuala II and no specialized setae on leg III. Differs from the other three genera of the subfamily Apoloniinae ( Whar- ton and Fuller, 1952) in lacking an anteromedian scutal projection. Vargatula hispida. new species. Figure 12. DIAGNOSIS : Scutum cuneiform, apex posterior, sensillae branched. One genuala I, no genualae II and III, no tibiala III, no parasubterminala. Coxal setae 1-2-1. Palpal setae branched. Dorsal, sternal, and ventral setae numerous. DESCRIPTION : Idiosoma. — Ellipsoidal. Length and width of holotype, nearly engorged, 515 by 290 /j.. Eyes absent. Anus at about the eighth row of ventral setae. Scutum. — As figured, cuneiform, moderately punctate, two pairs of pores at lateral margins in posterior half of scutum. Setae with long coarse branches. Sensillae branched to base. Measure- FIG. 12. Vargatula hispida, new species. Scutum. Palpal tarsus and tibia. Cheliceral blade. Specialized setae of legs I and II, with measurements in microns. ments of holotype : AW 41, SB 13, ASB 29, PSB 15, AM 12, AL 17, PL 32, s 24 /*. Gnathosoma. — Punctate. Capitular sternum transversely rugose. Cheliceral blade nearly straight, with minute dorsal tooth. Galeal seta nude. Palpal setae B/B/BBB; claw trifurcate; tarsus with five branched setae, a subterminala, and a tarsala. Legs. — Punctate. Specialized setae as figured. No pretarsala II. Non-specialized setae coarsely branched. Coxa I with one, II with two, and III with one branched setae. Empodia elongate, nearly filiform. Body setae. — Dorsal setae thickened and becoming thicker and longer posteriorly, heavily branched, 17 to 40 /*, four or five humerals on each side plus about 110 dorsals. Ventral setae, 12 to 16 sternals arranged 2-2 plus a fairly uniform group at the level of coxae III, and about 150 ventrals. Postanals similar to dorsals. TYPE MATERIAL : Holotype and 10 paratypes, RML no. 44425, off Dasypus novemcinctus, Paraiso (Canal Zone), 12 February 1962; 4 paratypes, same host and locality, 27 February and 26 March 1962, MARU. Holotype in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Mountain Laboratory, United States National Museum, Chicago Natural History Museum, British Museum (Natural History). Genus Alexf ainia Yunker and Jones Alexfainia Yunker and Jones, 1961, Jour. Parasit., 47, (6), p. 995. Type-species: Alexfainia chilonycteris Yunker and Jones, 1961. 226 ECTOPARASITES OF PANAMA KEY TO PANAMANIAN SPECIES Palpal femur with hamuli chilonycteris Yunker and Jones Palpal femur without hamuli munozi n. sp. Alexfainia chilonycteris Yunker and Jones Alexfainia chilonycteris Yunker and Jones, loc. cit., pp. 995-996, pi. I. More than 100 specimens (22 lots) from intranasal passages of (21) Pteronotus parnellii, Paraiso (Canal Zone), 18 August 1960, 4 January and FIG. 13. A lex fainia munozi, new species. Scutum. Cheliceral blade and palp. Specialized setae of legs. 2 March 1962, Madden Field (Canal Zone) , 7 July and 3 October 1961 ; 1 off Carollia perspicillata, Pacora (Panama) , 31 October 1960. See Yunker and Jones (1961) for other Panamanian records. Alexfainia munozi, new species. Figure 13. DIAGNOSIS : A large intranasal chigger, separated from A. chilonycteris Yunker and Jones by palpal femur lacking hamuli and scutum with more emarginate and narrow anterior margin. DESCRIPTION: Idiosoma. — Ellipsoidal, mildly constricted when engorged. Length and width of holotype, unengorged, 310 by 165 /u, of an engorged paratype, 1463 by 793 p.. Eyes absent. Anus at third row of ventral setae. Scutum. — As figured, densely punc- tate, cuneiform with a pronounced emarginate and narrow anterior margin. Anterior BRENNAN AND YUNKER : CHIGGERS 227 setae short, nude, the holotype anomalous in that the AM is absent; posterior setae much longer, with tiny barbs. Sensillae slender, elongate, with few minute barbs. Measure- ments of holotype: AW 44, pw 86, SB 54, ASB 39, PSB 38, AP 42, AM-, AL 8, PL 53, s 96 p. Gnathosoma. — Densely punctate. As in A. chilonycteris, a marked disparity in size be- tween the greatly reduced chelicerae and the exceedingly enlarged palpi, as figured. Palpal setae N/N/NNN; claw large, simple, recurved; tarsus with three heavily barbed thick setae, four small nude setae, and a tarsala. Galeal seta nude. Legs. — Punctate. Special- ized setae as figured. Non-specialized setae sparsely branched, some with few barbs, others apparently nude, a thick seta ventrally on telofemur and genu of leg III, much longer than homologous setae in chilonycteris. Coxal setae II much shorter than I and III. Body setae. — Dorsal setae with tightly appressed barbs, some appearing nude, 47 to 54 /u, arranged 2-6-6-8-6-6-4-2. Ventral setae, 2-2 sternals plus 34; sternals apparently nude, others minutely barbed. TYPE MATERIAL : Holotype, RML no. 44413, off Pteronotus psilotis, Pen- onome (Code), 8 February 1962; 8 paratypes, same host and locality, 30 January and 8 February 1962, V. J. Tipton, collector. Holotype in the col- lection of the Rocky Mountain Laboratory. Paratypes in the Rocky Moun- tain Laboratory, United States National Museum, Chicago Natural History Museum, and the British Museum (Natural History). Named for Mr. Angel Mufioz, Acarology Group, Middle America Re- search Unit. Genus Aniatrus Brennan and Jones Aniatrus Brennan and Jones, 1961, Jour. Parasit., 47, (1), p. 105. Type-species: Aniatrus bifax Brennan and Jones, 1961. Aniatrus bifax Brennan and Jones Aniatrus bifax Brennan and Jones, loc. cit., pp. 106-107, fig. 2. Five specimens off (2) Dasypus novemcinctus near Pedro Miguel River and Paraiso, El Rallo (Canal Zone) , 15 and 27 February 1962. A. bifax was described from a single specimen off Dasypus novemcinctus, Albrook Air Force Base (Canal Zone), 17 March 1959. Genus Ascoschoengastia Ewing Ascoschoengastia Ewing, 1946, Proc. Biol. Soc. Wash., 59: 71. Type-species: Neoschoengastia malayensis Gater, 1932. Ascoschoengastia dyscrita Brennan and Jones Ascoschoengastia dyscrita Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 107- 108, fig. 3. Twenty-eight specimens identified from 12 lots. Hosts : Didelphis mar- supialis, Heteromys desmarestianus , Hoplomys gymnurus, Liomys adsper- sus, Oryzomys capita, Tylomys watsoni. Localities : Pina, Cacao Plantation, Miraflores, Summit, and Corte Cule- bra Road (Canal Zone) ; Cerro Azul (Panama) . From 1959 to 1962, collec- tions were made from September to March with none from April to August. Brennan and Jones (1961a) record the following additional hosts : Oryzomys caliginosus, Sigmodon hispidus, and Proechimys semispinosus, the last from Bocas del Toro Province. 228 ECTOPARASITES OF PANAMA Genus Beamerella Brennan Beamerella Brennan, 1958, Jour. Kans. Ent. Soc., 31, (2), p. 71. Type-species : Beamerella acutascuta Brennan, 1958. Beamerella acutascuta Brennan Beamerella acutascuta Brennan, loc. cit., pp. 72-73, fig. 1. Six specimens off Micronycteris megalotis, Pacora (Panama), 31 Octo- ber 1960 ; 3 off Saccopteryx bilineata, Pacora, 31 October 1960 ; 7 off Carollia perspicillata, Los Santos Province, 27 January 1962. First Panamanian records. FIG. 14. Blix cabassoi, new species. Scutum and eyes. Cheliceral blade. Specialized setae of legs. Blix, new genus Type-species : Blix cabassoi, new species. DIAGNOSIS : Intranasal trombiculine larvae with leg segmentation 7-7-7. Legs without mastisetae, but with genualae II and III and a tibiala III. Scutum subquadrate, with five setae and expanded sensillae. Eyes present. Cheliceral blades with tricuspid cap and a prominent dorsal tooth. Palpal tarsus with five branched setae, a subterminala, and a tarsala; tibial claw trifurcate. This genus has affinities with Schoutedenichia from which it differs in the palpal tarsal setation, a tibiala III, unique form of scutum, etc. BRENNAN AND YUNKER : CHIGGERS 229 Blix cabassoi, new species. Figure 14. DIAGNOSIS : Fairly large intranasal species. Rectangular scutum deeper than wide, sensillae arising just within lateral margins. Cheliceral blades with large dorsal tooth in addition to tricuspid cap. Two genualae I and a tibiala III. Gnathosoma, scutum, and legs densely and conspicuously punctate. DESCRIPTION : Idiosoma. — Ellipsoidal, slightly constricted. Length and width of holo- type, slightly engorged, 885 by 390 /u. Eyes 2/2, no plate. Anus at sixth row of ventral setae. Scutum. — Approximately rectangular, with mildly convex posterior margin and lateral margins bulging opposite sensillary bases which they nearly touch. Puncta conspicuous and compact. AL'S reduced, with barbs suggested; AM and PL'S with ap- pressed barbs. Sensillae oblanceolate, apparently lacking setules. Measurements of holotype: AW 88, PW 76, SB 70, ASB 51, PSB 47, AP 60, AM 74, AL 18, PL 75, s 57 /*. Gnatho- soma.— Densely punctate. Chelicerae elongate, blades with tricuspid cap and prominent dorsal tooth. Palpi densely punctate, all setae barbed; tarsus with five branched setae, a subterminala, and short tarsala; claw trifurcate, one accessory prong smaller than the other. Galeal seta nude. Legs. — Densely punctate. Specialized setae as figured. Non-specialized setae long, with few appressed barbs. Body setae. — Dorsal setae barbed, 48 to 62 n, arranged 2-6-8-8-4-2-2. Ventral setae, 2-2 sternals plus about 70. TYPE MATERIAL : Holotype and 16 paratypes, RML no. 40076, from nasal mucosa of Cabassous centralis, Gamboa (Canal Zone), 29 November 1960, Dr. Nathan Gale, collector. Holotype in the collection of the Rocky Moun- tain Laboratory. Paratypes in the Rocky Mountain Laboratory, United States National Museum, Chicago Natural History Museum, and the British Museum (Natural History). Genus Cordiseta Hoffmann Cordiseta Hoffmann, 1954, An. Esc. Nac. Cienc. Biol. Mex., 8: 26. Type-species: Walchiella (Cordiseta) mexicana Hoffmann, 1954. Cordiseta mexicana (Hoffmann) Walchiella (Cordiseta) mexicana Hoffmann, loc. cit., pp. 27-30, figs. 15-20. One specimen off Scotinomys teguina, Cerro Punta (Chiriqui), 4 May 1960; one off Peromyscus nudipes, same locality, 18 January 1961. First records for Panama. Genus Crotiscus Ewing Crotiscus Ewing, 1944, Proc. Biol. Soc. Wash., 57: 102. Type-species: Trombicula desdentata Boshell and Kerr, 1942. Crotiscus desdentatus (Boshell and Kerr) Trombicula desdentata Boshell and Kerr, 1942, Rev. Acad. Colomb. Cienc. Exacta, Fisico-Quim. y Nat., 5, (17), pp. 11-12 (in reprint), figs. 15-17. About 230 specimens identified from 35 lots. Hosts : Didelphis marsu- pialis, Philander opossum, Proechimys semispinosus, Hoplomys gymnurus, Heteromys desmarestianus, Nectomys alfari, Sigmodon hispidus. Local- ities: Pina, Road K-9, and Fort Gulick (Canal Zone); Almirante, Cayo Agua, and Isla Bastimentos (Bocas del Toro) ; Cerro Campana (Panama). Taken throughout the year, from 1954 to 1962. First Panamanian records. 230 ECTOPARASITES OF PANAMA This polymorphic species is widely distributed in the American tropics. A form common to Panama is C. desdentatus tissoti Fauran (1960), de- scribed as a variety. Another similar form known from Cerro Campana has a much longer and strongly curved tarsala I. Crotonasis, new genus Type-species : Crotonasis fissa, new species. DIAGNOSIS : Intranasal trombiculine larvae with leg segmentation 7-7-7 ; coxae II and III multisetose. Scutum cuneiform, with five nude setae and short, nude, flagelliform sensillae. Eyes absent. Cheliceral blades short, recurved, with a single dorsal hook. Palpal tarsus with four branched setae and a tarsala ; tibial claw trif urcate. FIG. 15. Crotonasis fissa, new species, setae of legs. Scutum. Palpal tarsus and tibia. Specialized Distinguished from other American genera of intranasal chiggers with flagelliform sensillae, by the multisetose coxae II and III, trifurcate palpal tibial claw, and all scutal setae nude. Crotonasis fissa, new species. Figure 15. DIAGNOSIS: Two genualae I, genualae II and III, tibiala III. Coxae II and III multisetose. Three pairs of sternal setae. Palpal, genual, and tibial setae nude. Posterior margin of scutum with a median cleft. DESCRIPTION : Idiosoma. — Broad-ovate. Length and width of hoiotype, engorged, 405 by 278 /JL. Eyes absent. Anus at third row of ventral setae. Scutum. — As figured, cuneiform, with numerous fine puncta. All setae nude. Sensillary bases nearly touching lateral margins. Sensillae short, flagelliform, nude. Measurements of holotype : AW 14, PW 39, SB 20, ASB 21, PSB 17, AP 25, AM 15, AL 15, PL 27, S 24 n. Gnathosoma. — Finely punctate. Blades small, recurved, with a subapical dorsal hook. Galeal seta nude. Palpal setae B/N/NNN; tarsus with four branched setae and a tarsala; claw trifurcate. Legs. — Finely punctate. Specialized setae as figured, tarsala II thicker than tarsala I. Non- BRENNAN AND YUNKER: CHIGGERS 231 specialized setae sparsely branched. Coxa I with one, II with two, and III with seven to nine branched setae. Body setae. — Dorsal setae bare, or with barbs merely suggested, 22 to 30 /i, arranged 2-6-6-4-2. Ventral setae, 2-2-2 sternals plus 32. Sternals and preanals barbed, postanals similar to dorsals. TYPE MATERIAL: Holotype and 7 paratypes, RML no. 43471, from nasal mucosa of Liomys adspersus, Summit (Canal Zone), 28 August 1961; 3 paratypes, same host and locality, 27 and 28 December 1961, MARU. Holo- type in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Mountain Laboratory, United States National Museum, Chicago Natural History Museum, and the British Museum (Natural History) . Genus Doloisia Oudemans Doloisia Oudemans, 1910, Ent. Ber., 3: 87. Type-species: Doloisia synoti Oudemans, 1910. Subgenus Kymocta Yunker and Brennan Kymocta Yunker and Brennan, 1962, Acarologia, 4, (4), p. 572. Type-species: Kymocta teratarsalis Yunker and Brennan, 1962. KEY TO PANAMANIAN SPECIES Genuala III present teratarsalis Yunker and Brennan Genuala III absent chironectes Yunker and Brennan Doloisia (Kymocta) chironectes Yunker and Brennan Doloisia (Kymocta) chironectes Yunker and Brennan, 1962, Acarologia, 4, (4), pp. 574-576, fig. 3. No records other than of the original description : 2 specimens off Chiro- nectes minimus, Pedro Miguel (Canal Zone), 19 February 1962. Doloisia (Kymocta) teratarsalis Yunker and Brennan Doloisia (Kymocta) teratarsalis Yunker and Brennan, loc. cit., p. 574, fig. 2. Described from 6 specimens off Heteromys desmarestianus, Pina (Canal Zone), 7 to 15 December 1960; 1 off Hoplomys gymnurus, Pina, 6 December 1960 ; 7 off Tylomys watsoni, Pina, 8 August 1961. No further records. Genus Euschoengastia Ewing Euschoengastia Ewing, 1938, Jour. Wash. Acad. Sci., 28:293. Type-species: Euschoengastia americana Ewing, 1938 (=Schoengastia sciuricola Ewing, 1925). KEY TO PANAMANIAN SPECIES 1. With a mastitarsala III 2 Without a mastitarsala III 3 2. Palpal genual and ventrotibial setae branched; scutum deep, with narrow lance- olate sensillae tragulata Brennan and Jones Palpal genual and ventrotibial setae nude; scutum shallow, with broad obovate sensillae nunezi (Hoffmann) 3(1). Palpal tibial claw with five prongs; tibiala III absent; coxa III with three setae . . libertatis Brennan and Dalmat 232 ECTOPARASITES OF PANAMA Palpal tibial claw with three prongs; tibiala III present; coxa III usually uni- setose, rarely multisetose 4 4. With two genualae 1 5 With three genualae 1 7 5. Coxa III with two to four setae ; ventral setae extend into area of sternals cunctata Brennan and Jones Coxa III with one seta; ventral setae not extending to sternals 6 6. Galeal seta branched; palpal dorso- and laterotibial setae branched; dorsal for- mula begins 2-10 ; on rodents enhebra n. sp. Galeal seta nude; palpal dorso- and laterotibial setae nude; dorsal formula begins 2-6 ; on bats desmodus Brennan and Dalmat FIG. 16. Euschoengastia belgicae, new species. Scutum and eyes. Specialized setae of legs. 7(4). A single pair of eyes; palpal tarsala unusually long (25 fj.) ; on bats megastyrax Brennan and Jones Two pairs of eyes ; palpal tarsala normal 8 8. Scutum subquadrate; sensillae broad, oblanceolate ; palpal genual seta branched; on bats colombiae (Boshell and Kerr) Scutum much wider than deep; sensillae fusiform; palpal genual seta nude; on rodents 9 9. Eyes large; dorsal setae 32, more than 55 /* long; setules of sensillae long and densely distributed spissa Brennan and Jones Eyes reduced; dorsal setae 24, less than 50 /j. long; setules of sensillae long, but moderately distributed belgicae n. sp. Euschoengastia belgicae, new species. Figure 16. DIAGNOSIS : Distinguished from E. lipoglena Brennan and Jones (off bat, Trinidad) by presence of eyes; palpal genual and tibial setae nude; sensil- lary bases at level of PL's; and about half as many dorsal setae; from E. spissa Brennan and Jones by greatly reduced eyes, dorsal formula, and the much smaller size of all setae. BRENNAN AND YUNKER : CHIGGERS 233 DESCRIPTION: Idiosoma. — Broad-ovate. Dimensions of fully engorged specimen can- not be accurately determined because of crushed mount; approximate length and width, 475 by 390 p. Eyes 2/2, small, in obscure plate. Anus at fourth row of ventral setae. Scutum. — As figured, densely punctate, with sinuous margins. Sensillae fusiform, with elongate setules. Measurements : AW 66, PW 78, SB 33, ASB 24, PSB 17, AP 18, AM 34, AL 28, PL 48, S 41 /j.. Gnathosoma. — Moderately punctate. Blades with tricuspid cap. Galeal seta nude. Palpal setae B/N/NNN ; tarsus with at least five branched setae and a tarsala ; tibial claw trifurcate. Legs. — Punctate. Specialized setae as figured. Nonspecialized setae moderately branched. Idiosomal setae. — Dorsal setae with semi-appressed branches, 38 to 42 M> arranged 2-6-6-4-4-2. Ventral setae, 2-2 sternals plus 18. TYPE MATERIAL: Holotype, RML no. 40556, off Heteromys desmares- tianus, Cerro Azul (Panama), 17 March 1961, MARU. In the collection of the Rocky Mountain Laboratory. Named for Miss Belgica Rodriguez, Acarology Group, Middle America Research Unit. Euschoengastia colombiae (Boshell and Kerr) Neoschoengastia colombiae Boshell and Kerr, 1942, Rev. Acad. Colomb. Cienc. Exacta, Fisico-Quim. y Nat., 5, (17), pp. 16-18 (in reprint), figs. 6-8. One specimen off Carollia subrufa, Cerro Pirre (Darien), 31 January 1961 ; 1 off same host, Rio Changena (Bocas del Toro), 24 September 1961. First records for Panama. Euschoengastia cunctata Brennan and Jones Euschoengastia cunctata Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 108- 109, fig. 4. Known only from the type series, 8 specimens off Oryzomys capita (= talamancae) , Cerro Azul (Panama) , 8 to 14 February 1956. Euschoengastia desmodus Brennan and Dalmat Euschoengastia desmodus Brennan and Dalmat, 1960, Ann. Ent. Soc. Amer., 53, (2), pp. 188-189, fig. 7. One specimen off Saccopteryx bilineata, Chepo (Panama), 8 October 1959; 4 off bat, El Valle (Code), 30 May 1961; 4 off Glossophaga soricina, Rio Changena (Bocas del Toro) , 19 September 1961 ; 3 off Carollia subrufa, Rio Changena, 26 September 1961 ; 2 off Carollia castanea, Cerro Hoya (Los Santos) , 9 February 1962 ; 2 off Micronycteris megalotis, Fort Kobbe (Canal Zone), 2 February 1962. First Panamanian records. Euschoengastia enhebra, new species. Figure 17. DIAGNOSIS : Two genualae I, no genualae II and III, tibiala III. Distin- guished from E. utahensis Brennan and Beck by the widely separated sensil- lae, longer scutal setae, more body setae, branched galeal seta, and longer tarsala I than tarsala II. DESCRIPTION : Idiosoma. — Broad-ovate. Length and width engorged, 463 by 288 /JL. Eyes large, 2/2, in a plate. Anus at about fourth row of ventral setae. Scutum. — As figured, trapezoidal with mildly sinuous margins, punctate. Sensillae obovate, anterior surface densely setulose, posterior surface with a denuded median area, stems barbed nearly to base. Setae with semi-appressed branches. Measurements: AW 64, PW 95, SB 51, ASB 31, PSB 24, AP 42, AM 38, AL 34, PL 45, s 37 M- Gnathosoma. — Densely punctate. 234 ECTOPARASITES OF PANAMA Cheliceral bases broad, blades with tricuspid cap. Galeal setae branched. Palpal setae B/B/BBB; tarsus with five branched setae, a subterminala and a tarsala; tibial claw trifurcate. Legs. — Punctate. Specialized setae as figured. Non-specialized setae sparsely to moderately branched. Body setae. — Dorsal setae similar to scutals, 32 to 46 /j., arranged 2-10-2-6-6-6-6-2-2. Ventral setae, 2-2 sternals plus about 50, postanals similar to dorsals. TYPE MATERIAL: Holotype, RML no. 44401, off Tylomys watsoni, Pina (Canal Zone) , 7 February 1962, EHB. In the collection of the Rocky Moun- tain Laboratory. FIG. 17. Euschoengastia enhebra, new species. Scutum and eyes. Specialized setae of legs. Euschoengastia libertatis Brennan and Dalmat Euschoengastia libertatis Brennan and Dalmat, 1960, Ann. Ent. Soc. Amer., 53, (2), pp. 189-190, fig. 8. Four specimens off Peromyscus nudipes, Highlands of Chiriqui, 12 Feb- ruary 1960. First Panamanian record. Euschoengastia megastyrax Brennan and Jones Euschoengastia megastyrax Brennan and Jones, 1960, Acarologia, 2, (4), pp. 506- 507, fig. 7. Seven specimens off unidentified bat, Los Santos, 26 January 1962 ; 1 off Carollia perspicillata, Cacao Plantation (Canal Zone), 12 December 1961; 1 off Didelphis marsupialis , Pina (Canal Zone), 18 December 1959. First records for Panama. Euschoengastia nunezi (Hoffmann) Neoschoengastia nunezi Hoffmann, 1944, Rev. Inst. Salub. Enferm. Trop., 9, (3), pp. 221-225, figs. 1-4. One specimen off Didelphis marsupialis, Madden Forest (Canal Zone), 24 February 1955; 15 off Philander opossum, Canal Zone, 16 and 17 March 1955. First records for Panama. BRENNAN AND YUNKER : CHIGGERS 235 Euschoengastia spissa Brennan and Jones Euschoengastia spissa Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 109-110, fig. 5. Known only from the holotype, off Peromyscus nudipes, Highlands of Chiriqui, 4 May 1960. Euschoengastia tragulata Brennan and Jones Euschoengastia tragulata Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 110- 111, fig. 6. One specimen off Didelphis marsupialis, Pina (Canal Zone) , 16 February 1962 ; 33 off (5) Coendou rothschildi, vicinity of Pedro Miguel River (Canal Zone) , 21 February to March 1962. Described from 5 specimens off Nasua nasua, Barro Colorado Island (Canal Zone), November 1956. Genus Eutrombicula Ewing Eutrombicula Ewing, 1938, Jour. Wash. Acad. Sci., 28: 293. Type-species: Microthrombidium alfreddugesi Oudemans, 1910. KEY TO PANAMANIAN SPECIES 1. With three mastitarsalae III and two mastitibialae III; dorsal formula begins 2-8 batatas (Linnaeus) With one mastitarsala III and no mastitibialae III; dorsal formula begins 2-6 2 2. Accessory prong of palpal claw arises at distal three-fourths of axial prong alfreddugesi (Oudemans) Accessory prong of palpal claw arises at distal one-half of axial prong goeldii (Oudemans) Eutrombicula alfreddugesi (Oudemans) Microthrombidium alfreddugesi Oudemans, 1910, Ent. Ber., 3: 84. Some 1000 specimens identified from 245 lots. Hosts : REPTILES, Oxybelis sp., Pseustes poecilonotus, Spilotes pullatus, Ameiva bifrontata, A. undulata, Anolis sp., Sceloporus sp., unidentified lizards; BIRDS, Odontophorus ery- throps, Tar aba, major; MAMMALS, Didelphis marsupialis, Marmosa robin- soni, Philander opossum, Aotus trivirgatus, Saguinus geoffroyi, Proechimys semispinosus, Heteromys desmarestianus, Coendou rothschildi, Agouti paca, Hydrochaeris hydrochaeris, Sciurus granatensis, Sigmodon hispidus, Zygo- dontomys microtinus, Reithrodontomys mexicanus, Oryzomys alfaroi, O. capita, Oryzomys sp., Peromyscus flavidus, P. nudipes, Scotinomys teguina, S. xerampelinus, mouse, Sylvilagus brasiliensis, Nasua nasua, Tayassu tajacu. Localities : Galeta Point, Fort Gulick, France Field, Pina, Cacao Planta- tion, Summit, Madden Forest, Juan Mina, Pedro Miguel, Fort Clayton, Miraflores, Curundu, Roads K-9 and K-10, Corte Culebra Road, and Nuevo Emperador (Canal Zone) ; Cerro Azul, Cerro Campana, and Pacora (Pan- ama) ; Parita (Herrera) ; Cerro Pirre (Darien) ; Achiote (Colon) ; Almi- rante and Rio Changena (Bocas del Toro) ; Cerro Hoya (Los Santos) ; Boquete Trail, El Hato, Bambito, and Cerro Punta (Chiriqui). This com- mon pest chigger is more or less active throughout the year, with peak 236 ECTOPARASITES OF PANAMA activity during the wet season, especially in the lowlands, according to col- lections from 1954 to 1962. Both the "lipovskyana" and "tropica" forms are included in the above records. The former, from the Highlands of Chiriqui, is distinguished by a longer than usual tarsala I, and the latter, principally from lowlands, is distinguished by nude genual and ventrotibial palpal setae. Eutrombicula batatas (Linnaeus) Acarus batatas Linnaeus, 1758, Syst. Nat. (10th ed.), p. 617. About 225 specimens identified from 95 lots. Hosts : BIRDS, Gallus gallus, Hypomorphnus urubitinga, Caracara plancus, Catharus mexicanus, Croto- phaga ani, Nyctidromus albicollis, Myiozetetes sp. ; MAMMALS, Homo sapiens, Hydrochaeris hydrochaeris, Sigmodon hispidus, Zygodontomys microtinus, Reithrodontomys sp., Mazama americana. Localities: Fort Gulick, France Field, Fort Davis, Gatun, Nuevo Em- perador, Fort Clayton, Fort Kobbe, and Curundu (Canal Zone) ; Tocumen and Pacora (Panama) ; Achiote (Colon) ; Almirante and Rio Changena (Bocas del Toro) ; El Hato (Chiriqui). Active during the wet season; no records available for January, February, or March. Eutrombicula goeldii (Oudemans) Microthrombidium goeldii Oudemans, 1910, Ent. Ber., 3: 84. Some 1000 specimens identified from 280 lots. Hosts : REPTILES, Pseustes poecilonotus, Ameiva bifrontata, A. festiva, A. undulata; BIRDS, Momotus momota, Neomorphus geoffroyi salvini, Malacoptila panamensis, Cyphor- hinus aradus phaeocephalus ; MAMMALS, Didelphis marsupialis, Marmosa mexicana, M. robinsoni, Metachirus nudicaudatus , Philander opossum, un- identified bats, Aotus trivirgatus, Saguinus geoffroyi, Dasypus novemcinc- tus, Proechimys semispinosus, Hoplomys gymnurus, Coendou rothschildi, Liomys adspersus, Heteromys australis, H. desmarestianus, Sciurus granat- ensis, Microsciurus alfari, Dasyprocta punctata, Rattus rattus, Sigmodon hispidus, Oryzomys caliginosus, O. capito, Nectomys alfari, Zygodontomys microtinus, Sylvilagus brasiliensis, Nasua nasua, Felis pardalis. Localities: Barro Colorado Island, Pina, Galeta Point, Fort Gulick, France Field, Fort Sherman, Fort Davis, Cacao Plantation, Summit, Mad- den Forest, Juan Mina, Miraflores, Pedro Miguel, Fort Clayton, Nuevo Em- perador, Road K-9, Corte Culebra Road, Fort Kobbe, and Ancon Hill (Canal Zone) ; Coiba Island; Cerro Azul, Cerro Campana, Capira, Guayabito, and Madden Airstrip (Panama) ; Cerro Pirre (Darien) ; Divisa and Parita (Herrera) ; Achiote (Colon) ; Cerro Hoya (Los Santos) ; Almirante, Rio Changena, Cayo Agua, Escudo de Veraguas, and Isla Bastimentos (Bocas del Toro) ; Chiriqui Province. Common throughout the year, as suggested by miscellaneous collections made from 1954 to 1962. BRENNAN AND YUNKER: CHIGGERS 237 Genus Hoffmannina Brennan and Jones Hoffmannina Brennan and Jones, 1959, Ann. Ent. Soc. Amer., 52: 8. Type-species: Novotrombicula suriana Hoffmann, 1954. KEY TO PANAMANIAN SPECIES With two genualae I handleyi Brennan and Jones With three genualae I suriana (Hoffmann) Hoffmannina handleyi Brennan and Jones Hoffmannina handleyi Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 111-112, fig. 7. Thirty-two specimens in 16 lots off Didelphis marsupialis, Peromyscus nudipes, Reithrodontomys sp., El Hato, Bambito, and Cerro Punta (Chi- riqui) , 7 to 31 January, 26 April to May 1961. Brennan and Jones (1961a) report this species also from Scotinomys teguina, Reithrodontomys mexi- canus, and Heteromys desmarestianus, all from the same general area in the Chiriqui Highlands. Hoffmannina suriana (Hoffmann) Novotrombicula suriana Hoffmann, 1954, Ann. Esc. Nac. Cienc. Biol. Mex., 8, (1-2), pp. 23-26, figs. 9-12. Nine specimens identified off Reithrodontomys sumichrasti, Cerro Baru, 10500 feet (Chiriqui), 2 May 1960; 1 off Scotinomys xerampelinus, 7 off Peromyscus nudipes, 9 off Reithrodontomys creper, Boquete Trail (Chi- riqui) , 3 to 5 May 1961. First Panamanian records. Intercutestrix, new genus Type-species: Euschoengastia tryssa Brennan and Jones, 1961. DIAGNOSIS: Intradermal trombiculine larvae with leg segmentation 7-6-6 ; legs without mastisetae, parasubterminala, and genualae II and III. Scutum much wider than deep, with five setae and expanded sensillae. Eyes present. Cheliceral blades with tricuspid cap. Palpal tarsus with four branched setae and a tarsala. Intercutestrix tryssa (Brennan and Jones), new combination. Figure 18. Euschoengastia tryssa Brennan and Jones, 1961, Acarologia, 3, (2), pp. 189-190, fig. 9. Additional material from Panama has indicated the need for erecting a genus to receive this species which was described from a single Peruvian specimen off Proechimys hendeei. The conspecificity of this and the Pan- amanian form is evident, although as might be expected, the latter exhibits geographic variation and may be a subspecies. Here, as figured, the scutum is somewhat deeper and AM < AL. The dorsal setae are longer. Engorged specimens of both forms are circular in outline. Forty-five specimens off (15) Proechimys semispinosus, all from Canal Zone : Fort Gulick, 9 March 1954, 7 December 1960, 14 and 15 December 1961 ; France Field, 8 September to 21 December 1961 ; Pifia, 29 November and 29 August, 1961, 9 February 1962 ; Summit, 1 December 1960. One off 238 ECTOPARASITES OF PANAMA Oryzomys caliginosus, Cerro Azul (Panama), 26 January 1956. Two off Tylomys watsoni, Pifia (Canal Zone), 7 February 1962, and 4 off Oryzomys capita, Pifia, 20 December 1960. First records for Panama. FIG. 18. Intercutestrix tryssa (Brennan and Jones). Scutum of holotype (Peru), above, and Panamanian form, below. Genus Leptotrombidium Nagayo et al. Leptotrombidium Nagayo et al., 1916, Dobutsugaku Zasshi, 28: 392. Type-species: Trombidium akamushi Brumpt, 1910. Leptotrombidium panamensis (Ewing) Trombicula panamensis Ewing, 1925, Amer. Jour. Trop. Med., 5, (3), pp. 259-260. About 160 specimens from 36 lots. Hosts : Didelphis marsupialis, Mar- moset, robinsoni, Artibeus toltecus (one specimen, Bocas del Toro), Pro- echimys semispinosus, Hoplomys gymnurus, Heteromys australis, Liomys adspersus, Coendou rothschildi, Sigmodon hispidus, Oryzomys bombycinus, 0. caliginosus, 0. capito, Zygodontomys microtinus. Localities: Fort Sherman, Gamboa Road, Summit, Madden Forest, Pedro Miguel River, and Fort Kobbe (Canal Zone) ; Almirante and Rio Changena (Bocas del Toro). Irregular collections from 1954 to 1962 sug- gest that peak activity of this species is during the dry season, January to April, although a few collections were made in June and September. Myxacarus, new genus Type-species : Myxacarus oscillatus, new species. DIAGNOSIS : Intranasal trombiculine larvae with leg segmentation 7-7-7 ; legs without mastisetae and genualae II and III, but with tibiala III. Scutum subquadrate, with five setae and expanded sensillae. Eyes apparently absent. Cheliceral blades with a series of dorsal teeth of which the proximal is enlarged. Palpal tarsus with five branched setae and a tarsala; tibial claw bifurcate. Ventral humeral setae present. Distinguished from Blix n. gen. by armature of cheliceral blades, special- BRENNAN AND YUNKER: CHIGGERS 239 ized setation of legs, bifurcate palpaltibial claw, and ventral humeral setae. Myxacarus oscillatus, new species. Figure 19. DIAGNOSIS : Intranasal habitat. Scutum subquadrate with concave pos- terior margin and PL'S arising from tubercles. Cheliceral blades with series of dorsal teeth. Two genualae I, no genualae II or III. FIG. 19. Myxacarus oscillatus, new species, setae of legs. Scutum. Half of gnathosoma. Specialized DESCRIPTION: Idiosoma. — Broad-ellipsoidal, slightly constricted. Length and width of holotype 610 by 390 u. Eyes not seen. Anus at fourth row of ventral setae. Scutum. — Subquadrate with straight to concave posterior margin, densely punctate. Postero- lateral angles produced and tuberculate. Anterior setae with distinct barbs, posterior setae apparently nude. Sensillae broadlanceolate, with minute setules. Measurements of holotype: AW 47, PW 76, SB 26, ASB 30, PSB 29, AP 57, AM 40, AL 40, PL 40, S 40 n. Gnath- osoma.— Densely punctate. Blades strong, with a large dorsal tooth on apical third, in front of which is a series of four or five smaller teeth. Galeal seta nude. Palpal setae B/N/NNN: tarsus with five branched setae and a tarsala; claw bifurcate. Legs. — Densely punctate. Specialized setae as figured. Non-specialized setae barbed to branched. Empodium long and slender. Body setae. — Dorsal setae with short barbs, 29 to 46 n, arranged 2-6-6-4-2. Ventral setae, 2-2 sternals and three to four ventral num- erals on each side between coxae II and III, plus 30. Postanals similar to dorsals. TYPE MATERIAL: Holotype and a paratype, RML no. 44241, from nasal mucosa of Proechimys semispinosus, Summit (Canal Zone), 20 December 240 ECTOPARASITES OF PANAMA 1961, MARU. Paratypes: 3, same host, Pifia (Canal Zone), 20 December 1960, and Fort Gulick (Canal Zone) , 6 January 1961, EHB ; 11 off Hoplomys gymnurus, Pina, 6 December 1960, EHB; 1 off Oryzomys capita, Pifia, 7 December 1960, EHB; 6 off Dasypus novemcinctus, Paraiso (Canal Zone), 15 February 1962, MARU ; 1 off Metachirus nudicaudatus, Cerro Azul (Pan- ama), 16 March 1961, MARU. Holotype in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Mountain Laboratory, United States National Museum, Chicago Natural History Museum, and the British Museum (Natural History). FIG. 20. Neoschoengastia electron, new species. Scutum and eyes. Specialized setae of legs. Genus Neoschoengastia Ewing Neoschoengastia Ewing, 1929, Man. Ext. Parasites, p. 187. Type-species: Schoengastia americana Hirst, 1921. Neoschoengastia electron, new species. Figure 20. DIAGNOSIS : Three pairs of humeral setae, sternal setae 2-4, three genualae I, parasubterminala absent, coxa III with three setae. DESCRIPTION : Idiosoma. — Broad-ellipsoidal. Eyes 2/2, in a plate. Length and width of holotype, 587 by 400 ft. Anus at sixth row of ventral setae. Scutum. — Wider than long, with sinuous margins, puncta large, whorls of cuticular striations cover posterior two- thirds of scutum. Setae long, with semi-appressed branches. Sensillae missing. Meas- urements of holotype: AW 62, PW 83, SB 44, ASB 24, PSB 20, AP 27, AM 39, AL 57, PL 78, s- /*. Gnathosoma. — Punctate. Blades with tricuspid cap. Palpal setae B/B/NNB; tarsus with five branched setae, a subterminala, and a tarsala; claw trifurcate. Galeal seta branched. Legs. — Punctate. Specialized setae as figured. No parasubterminala. Non-specialized setae moderately branched. Coxa III with three setae. Tarsal claws with tenent hairs. Body setae. — Dorsal setae 50 to 75 /u, arranged approximately 6-9-8-2-7-5-4-4. Ventral setae, 2-4 sternals plus 52, postanals like dorsals. TYPE MATERIAL : Holotype, RML no. 44332, off Electron platyrhynchum, BRENNAN AND YUNKER: CHIGGERS 241 Cacao Plantation Road (Canal Zone), 2 February 1962, MARU. In the collection of the Rocky Mountain Laboratory. Genus Perates Brennan and Dalmat Perates Brennan and Dalmat, 1960, Ann. Ent. Soc. Amer., 53: 186. Type-species: Perates insessus Brennan and Dalmat, 1960. Perates insessus Brennan and Dalmat Perates insessus Brennan and Dalmat, loc. cit., p. 187, fig. 5. Eleven specimens off (3) Pteronotus psilotis, Penonome (Code), 30 January and 8 February 1962. First Panamanian records. FIG. 21. Perissopalla precaria (Brennan and Dalmat). Scutum and eyes. Genus Perissopalla Brennan and White Perissopalla Brennan and White, 1960, Jour. Parasit., 46: 346. Type-species : Perissopalla flagellisetula Brennan and White, 1960. Perissopalla precaria (Brennan and Dalmat) , new combination. Figure 21. Euschoengastia precaria Brennan and Dalmat, 1960, Ann. Ent. Soc. Amer., 53, (2), p. 190, fig. 9. Eighteen specimens off Micronycteris megalotis, Pacora (Panama), 31 October 1960. This species is transferred to the genus Perissopalla Bren- nan and White on the basis of a specimen in the above series that bears sensillae. Previous specimens lacked these structures. First record for Panama. Genus Polylopadium Brennan and Jones Polylopadium Brennan and Jones, 1961, Jour. Parasit., 47: 112. Type-species: Polylopadium kramisi Brennan and Jones, 1961. The discovery of a second species of Polylopadium, P. confirmatum n. sp., permits a revision of the generic diagnosis as follows : 242 ECTOPARASITES OF PANAMA Trombiculine larvae with dorsal pattern of platelets in addition to scutum. Scutum with five short inconspicuously barbed setae, sensillae expanded. Eyes present or absent. Dorsal setae short and barbed, similar to scutals. Cheliceral blades with tricuspid cap. Palpal tibial claw trif urcate. Leg seg- mentation 7-6-6. FIG. 22. Polylopadium confirmatum, new species. Dorsum, showing scutum and platelets. Oblique lighting photomicrograph by N. J. Kramis. KEY TO PANAMANIAN SPECIES With 11 dorsal platelets; eyes absent; genualae II and III absent kramisi Brennan and Jones With 16 dorsal platelets; eyes present; genualae II and III present confirmatum n. sp. Polylopadium confirmatum, new species. Figures 22, 23. DIAGNOSIS : Dorsum with 16 platelets, two genualae I and a genuala II and III, eyes present. DESCRIPTION: Idiosoma. — Ovate. Eyes 2/2, in a plate, posterior pair obsolescent. Dorsum with 16 small platelets arranged as figured. Length and width of holotype, partly engorged, 315 by 235 /*• Anus at seventh row of ventral setae. Scutum. — Trape- BRENNAN AND YUNKER: CHIGGERS 243 zoidal, with anteromedian and posteromedian extensions, as figured. Setae short and barbed. Sensillae oblanceolate, setules conspicuous. Measurements of holotype: AW 46, PW 59, SB 30, ASB 23, PSB 33, AP 21, AM 7, AL 8, PL 10, S 28 M- Gnathosoma. — Cheliceral blade with tricuspid cap and small ventral notch. Palpal setae B/B/BBB; claw trifurcate; tarsus with five branched setae and a tarsala. Galeal seta nude. Legs. — With fine puncta. Specialized setae as figured. Non-specialized setae sparsely branched. Body setae. — Dorsal setae short, barbed, 14 to 18 /*, arranged in the holotype 4-6-6-2-6-4-4-2. Ventral setae, 2-2 sternals plus 38, sternals with few pronounced branches similar to coxal setae, postanals similar to dorsals. FIG. 23. Polylopadium species. Scutum and eyes of P. confirmatum, new species. Sensilla of P. kramisi Brennan and Jones. Specialized setae of legs of P. confirmatum. TYPE MATERIAL: Holotype, RML no. 43719, off Oryzomys caliginosus, near Rio Changena (Bocas del Toro), 21 September 1961. Two paratypes, same host and locality, 26 and 27 September 1961, EHB. Holotype in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Mountain Laboratory and the United States National Museum. Polylopadium kramisi Brennan and Jones. Figure 23. Polylopadium kramisi Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 112-113, figs. 8, 9. Off (4) Liomys adspersus in the Canal Zone: 2 specimens, Curundu, 3 October 1961 ; 8 specimens, Summit, 27 and 28 December 1961 ; 3 specimens, Cacao Plantation, 31 January 1962. Also reported by Brennan and Jones (1961a) from Proechimys semispinosus, Canal Zone, and from Peromyscus sp., Chiriqui Province. A specimen bearing sensillae permits us to illustrate this structure. Genus Pseudoschoengastia Lipovsky Pseudoschoengastia Lipovsky, 1951, Jour. Kans. Ent. Soc., 24: 95. Type-species : Pseudoschoengastia hunger -fordi Lipovsky, 1951. KEY TO PANAMANIAN SPECIES 1. With one genuala I; genualae II and III absent mermeriza n. sp. With two or three genualae I ; genualae II and III present 2 244 ECTOPARASITES OF PANAMA 2. PL'S on scutum; parasubterminala branched; one pair of humeral setae. . .apista n. sp. PL'S off scutum ; parasubterminala nude ; two pairs of humeral setae 3 3. Eyes absent finitima n. sp. Eyes present 4 4. With two genualae I ; idiosomal setae arise from deep, wide pits dasypi n. sp. With three genualae I ; idiosomal setal bases normal 5 5. Sensillae capitate, a small bulb at base; palpal genual and tibial setae branched; AL'S much longer than PL'S bulbifera Brennan Sensillae obovate, no bulb at base; palpal genual tibial and setae nude; AL'S shorter than PL'S 6 6. About 48 short dorsal setae; SB = 18 n abditiva Brennan About 60 longer dorsal setae; SB = 28 /x zona Brennan FIG. 24. Pseudoschoengastia apista, new species, legs. Scutum and eyes. Specialized setae of Pseudoschoengastia abditiva Brennan Pseudoschoengastia abditiva Brennan, 1960, Acarologia, 2, (4), pp. 482-483, fig. 1. Known only from the type series, 4 specimens off Oryzomys capita, Cerro Azul (Panama), 8 February 1956. Pseudoschoengastia apista, new species. Figure 24. DIAGNOSIS : Related to P. inevicta Brennan from which it differs by nude dorsotibial palpal seta, branched parsubterminala, leg segmentation 7-7-7, and tarsala I longer than tarsala II. Distinguished from all other members of the genus by having only one pair of humeral setae. DESCRIPTION : Idiosoma. — Broad-ovate. Length and width of holotype, engorged, 345 by 255 fj.. Eyes large, 2/2, in a plate. Anus at fourth row of ventral setae. Scutum. — As figured, with sinuous margins, moderately punctate. Posterolateral setae on scutum. Sensillae obovate, densely setulose on anterior surface, fewer and larger setules on poste- rior surface, stems barbed nearly to base. Measurements of holotype : AW 46, PW 64, SB 29, ASB 21, PSB 16, AP 28, AM 25, AL 19, PL 30, S 30 jt. Gnathosoma. — Moderately punctate. Cheliceral blades with tricuspid cap and dorsal and ventral tooth. Galeal seta nude, some- times forked. Palpal setae B/B/NNB; tarsus with five branched setae and a tarsala; claw trifurcate. Legs. — Punctate. Segmentation 7-7-7. Specialized setae as figured, para- subterminala forked or branched. Non-specialized setae moderately and conspicuously BRENNAN AND YUNKER: CHIGGERS 245 branched. Body setae. — Dorsal setae short, with conspicuous branches, 24 to 32 /j., ar- ranged 2-8-2-8-2-6-6-4. Ventral setae, 2-2 sternals plus 40, postanals similar to dorsals, two or three ventral humerals on each side between coxae II and III. TYPE MATERIAL : Holotype and 8 paratypes, RML no. 44654, off Dasypus novemcinctus, Paraiso (Canal Zone), 26 March 1962, MARU. Holotype, in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Mountain Laboratory, United States National Museum, British Museum (Natural History), and the Chicago Natural History Museum. FIG. 25. Pseudoschoengastia dasypi, new species. Posterior aspect of sensilla. Scutum and eyes; bases of dorsal setae; sensilla, anterior surface. Specialized setae of legs. Pseudoschoengastia bulbifera Brennan Pseudoschoengastia bulbifera Brennan, 1960, Acarologia, 2, (4), pp. 483-484, fig. 2. About 425 specimens identified from 135 lots. Hosts : REPTILES, Scelo- porus sp. ; MAMMALS, Didelphis marsupialis, Marmosa mexicana, M. robin- soni, Philander opossum, Cryptotis sp., Sturnira ludovici, Saguinus geof- froyi, Proechimys semispinosus, Heteromys australis, H. desmarestianus, Liomys adspersus, Hoplomys gymnurus, Sigmodon hispidus, Oryzomys al- faroi, O. bombycinus, O. callginosus, O. albigularis, O. capito, Oryzomys sp., Peromyscus nudipes, Peromyscus sp., Reithrodontomys sp., Scotinomys xer- ampelinus, Zygodontomys microtinus, Nectomys alfari. Localities: Pina, Miraflores, Summit, and Corte Culebra Road (Canal Zone) ; Cerro Azul (Panama) ; Isla Bastimentos and Rio Changena (Bocas del Toro) ; Cerro Pirre (Darien) ; Boquete Trail, Cerro Punta, Bambito, and El Hato (Chiriqui). See Brennan (1960) for other Panamanian records. In miscellaneous collections made from 1954 to 1962, this species was found in all months except June, July and August. Pseudoschoengastia dasypi, new species. Figure 25. DIAGNOSIS: Separated from other species of the genus by coarsely branched idiosomal setae arising from deep, wide pits. It is related to 246 ECTOPARASITES OF PANAMA P. guatemalensis Brennan from which it differs by wider scutum in propor- tion to depth, more widely separated sensillary bases, and more and larger idiosomal setae. DESCRIPTION: Idiosoma. — Ovate. Length and width of holotype, slightly engorged, 237 by 165 M- Eyes large, 2/2, in a plate. Anus at fourth row of ventral setae. Scu- tum.— Conspicuously punctate, as figured ; extrascutal PL'S coarsely branched setae. Sen- sillate capitate, anterior surface moderately setulose and with small setules sparsely dis- tributed on posterior surface. Measurements of holotype: AW 55, SB 29, ASB 29, PSB 19, AM 32, AL 73, PL 53, S 32 /JL. Gnathosoma, — Conspicuously punctate. Cheliceral blades FIG. 26. Pseudoschoengastia finitima, new species. Scutum. Specialized setae of legs. with tricuspid cap and minute subapical dorsal and ventral teeth. Galeal seta nude. Palpal setate B/B/BNB; tarsus with five branched setae and a tarsala; claw trifurcate. Legs. — Punctate. Leg segmentation 7-6-6. Specialized setae as figured. Non-specialized setae coarsely branched. Body setae. — Dorsal setae coarsely branched, arising from deep, wide pits, 33 to 50 /*, 2-2 numerals plus about 70. Ventral setae, 2-2 sternals, 2-2 ventral numerals, plus about 60. These also arise from wide pits. TYPE MATERIAL : Holotype and 4 paratypes, RML no. 44654, off Dasypus novemcinctus, Paraiso (Canal Zone) , 26 March 1962, MARU. In the collec- tion of the Rocky Mountain Laboratory. All specimens in fair to poor con- dition. Pseudoschoengastia finitima, new species. Figure 26. DIAGNOSIS : Distinguished from all other species of the genus by the ab- sence of eyes, and from P. farneri Lipovsky by the more appressed branches of branched setae, wider scutum in proportion to depth, and smaller tarsalae I and II. DESCRIPTION : Idiosoma. — Broad-ovate, slightly constricted. Length and width of holo- type, engorged, 400 by 260 /j.. Eyes abs?nt. Anus at about fifth row of ventral setae. Scutum. — As figured, PL'S extrascutal, sensillae broken off throughout the series. Meas- urements of holotype: AW 47, SB 25, ASB 21, PSB 10, AM 33, AL 24, PL 42, s- /j.. Gnath- osoma.— Punctate. Cheliceral blades with tricuspid cap, and apparently without addi- tional teeth. Galeal seta nude. Palpal setae B/N/BNB; tarsus with five branched setae BRENNAN AND YUNKER : CHIGGERS 247 and a tarsala; claw trifurcate. Legs. — Leg segmentation 7-6-6. Specialized setae as figured. Non-specialized setae moderately branched. Body setae. — Dorsal setae 32 to 40 fj., 2-2 humerals plus about 50. Ventral setae, 2-2 sternals, 2-2 numerals, plus 60. TYPE MATERIAL: Holotype and 7 paratypes, RML no. 40112, off Het- eromys desmarestianus, Pina (Canal Zone), 7 December 1960. Two para- types, same host and locality, 29 December 1961, EHB. Holotype in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Lab- oratory, United States National Museum, British Museum (Natural His- tory), and the Chicago Natural History Museum. FIG. 27. Pseudoschoengastia mermeriza, new species. Scutum and eyes. Specialized setae of legs. Pseudoschoengastia mermeriza, new species. Figure 27. DIAGNOSIS: Similar to P. aberrans Brennan and Jones (PL'S on deep- narrow scutum, palpal setation, one genuala I, no genualae II and III, and small eyes) but a distinct form with leg segmentation apparently 7-7-6, all branched setae with more appressed and shorter branches and different ar- rangement of dorsal setae. DESCRIPTION : Idiosoma. — Broad-ovate. Length and width, partly engorged, 285 by 206 /j.. Eyes small, 2/2, posterior pair obsolescent. Anus at fourth row of ventral setae. Scutum. — As figured, punctate, deep-narrow. Posterolateral setae on scutum. Sensillae obovate, with large setules, anterior surface completely covered, posterior surface with a denuded median area, stems barbed nearly to their bases. Measurements: AW 41, PW 50, SB 19, ASB 20, PSB 17, AP 23, AM 24, AL 12, PL 31, s 29 /JL. Gnathosoma. — Moderately punc- tate. Cheliceral blades with tricuspid cap and a ventral tooth. Galeal seta nude. Palpal setate B/N/NNB; tarsus with five branched setae and a tarsala; tibal claw trifurcate. Legs. — Punctate. Segmentation 7-7-6. Specialized setae as figured. Non-specialized setae moderately branched. Body setae. — Dorsal setae similar to PL'S with short ap- pressed branches, 19 to 28 /JL arranged 4-4-6-8-2-8-6-4-2. Ventral setae, 2-3 (can be in- terpreted as 2-2 or 2-4) plus 32, postanals similar to dorsals, two ventral humerals on each side between coxae II and III. TYPE MATERIAL: Holotype, RML no. 44508, off Coendou rothschildi, Paraiso (Canal Zone), 26 February 1962, MARU. In the collection of the Rocky Mountain Laboratory. 248 ECTOPARASITES OF PANAMA Pseudoschoengastia zona Brennan Pseudoschoengastia zona Brennan, I960, Acarologia, 2, (4), pp. 490-492, fig. 8. Five specimens off Liomys adspersus, Nuevo Emperador (Canal Zone), 7 August 1961 ; 3 off same host, Summit (Canal Zone) , 27 and 28 December 1961; 1 off Heteromys australis, Cerro Pirre (Darien), 3 February 1961; 1 off Oryzomys caliginosus, Rio Changena (Bocas del Toro), 23 September 1961. In addition, reported by Brennan (1960) from Sigmodon hispidus and Tylomys panamensis. Genus Speleocola Lipovsky Speleocola Lipovsky, 1952, Jour. Kans. Ent. Soc., 25: 132. Type-species : Speleocola tadaridae Lipovsky, 1952. Speleocola secunda Brennan and Jones Speleocola secunda Brennan and Jones, 1960, Acarologia, 2, (4), pp. 509-510, fig. 8. Three specimens off Coendou rothschildi, along Pedro Miguel River (Canal Zone) , 20 March 1962. First record for Panama. Genus Tecomatlana Hoffmann Tecomatlana Hoffmann, 1947, An. Esc. Nac. Cienc. Biol. Mex., 4, (4), p. 451. Type-species : Tecomatlana sandovali Hoffmann, 1947. Tecomatlana sandovali Hoffmann Tecomatlana sandovali Hoffmann, loc. cit., pp. 452-457, figs. 1-6. Six specimens off Peropteryx macrotis, Buena Vista (Colon) , 29 October 1959; 5 off undetermined bats, cave near Quebrada Bonita (Panama), 16 February 1962. First records for Panama. Genus Blankaartia Oudemans Blankaartia Oudemans, 1911, Ent. Ber., 3: 123. Type-species : Trombidium niloticum Tragardh, 1905. KEY TO PANAMANIAN SPECIES 1. Sensillae nude; all setae of palpal tarsus nude marui n. sp. Sensillae branched, at least some setae of palpal tarsus branched 2 2. Palpal tibial claw bifurcate; palpal genual seta nude: galeal seta nude; tarsala I extremely long ( >50 /j.) wetmorei n. sp. Palpal tibial claw trifurcate; palpal genual seta branched; galeal seta forked or branched; tarsala I not unusually long 3 3. Scutum not pentagonal, its posterior margin broadly curved arremonops (Brennan and Jones) Scutum pentagonal 4 4. Posterior angle of scutum acute attenuata (Michener) Posterior angle of scutum not acute 5 5. Length of tarsala I about 20 M alleei (Ewing) Length of tarsala I about 15 // 6 6. Large, heart-shaped species (length, engorged, may exceed 1 mm.) sinnamaryi (Floch and Fauran) Smaller species, not heart-shaped velascoi (Boshell and Kerr) BRENNAN AND YUNKER : CHIGGERS 249 Blankaartia alleei (Ewing) Trombicula alleei Ewing, 1926, Ent. News, 37: 111-112. Ten specimens off Hydrochaeris hydrochaeris, Cerro Pirre (Darien), 7 February 1961. This species was described from a single adult from Barro Colorado Island. For additional information, including larval records from Panama, see Michener (1946) who gives biologic data and shows that the larvae are characteristically parasitic on birds. FIG. 28. Blankaartia marui, new species. Scutum and eyes. Specialized setae of legs. Blankaartia arremonops (Brennan and Jones), new combination. Trombicula arremonops Brennan and Jones, 1961, Jour. Parasit., 47, (2), pp. 114-115, fig. 10. Known only from the holotype off Arremonops conirostris, Cacao Planta- tion (Canal Zone) , 4 May 1955. Blankaartia attenuata (Michener) Trombicula (Megatrombicula) attenuata Michener, 1946, Ann. Ent. Soc. Amer., 39, (3), pp. 440-444, figs. 12-17, 22. The description of this species is based principally on adults of both sexes, found only on floating plants of water lettuce (Pistia stratiotes) , Juan Mina (Canal Zone), September and October 1945. Larvae, obtained by rearing, were briefly described. The holotype is a male, the allotype a female. Hosts are unknown. No additional records. Blankaartia marui, new species. Figure 28. DIAGNOSIS: Distinguished from all other species by the nude sensillae and nude setae of palpal tarsus ; in addition, distinguished from wetmorei n. sp. by the greater number of body setae and much shorter tarsala I. DESCRIPTION : Idiosoma. — Ellipsoidal. Length and width of holotype 885 by 463 /*. 250 ECTOPARASITES OF PANAMA Eyes prominent, 2/2, in a plate, the anterior larger. Anus at fourth row of ventral setae. Gnathosoma. — Densely and conspicuously punctate. Chelicerae long and narrow, blades long, with tricuspid cap. Palpal setae B/N/NNN; tarsus with six nude setae, a subterminala and a tarsala; claw bifurcate. Galeal seta nude. Scutum. — As figured, nearly quadrate, with shallow curving posterior margin, densely punctate. Setae with appressed branches, AM and AL'S considerably removed from anterior margin. Sensillae thread-like, nude. Measurements of holotype: AW 99, PW 100, SB 36, ASB 52, PSB 41, AP 44, AM 40, AL 40, PL 51, s 88 fi. Legs. — Long, conspicuously punctate. Specialized setae as figured. Non-specialized setae long, sparsely branched, some seemingly nude. Body setae. — Dorsal setae with appressed branches, 42 to 55 /u, arranged 2-6-6-6-4-2-2. Ventral setae, 2-2 sternals plus 36. TYPE MATERIAL : Holotype and a paratype, RML no. 40982, off Nycticorax nycticorax, Cerro Azul (Panama) , 29 May 1961 ; one paratype, same collect- ing1 data, MARU. Holotype in the collection of the Rocky Mountain Labo- ratory. Paratypes in the Rocky Mountain Laboratory and the United States National Museum. Blankaartia sinnamaryi (Floch and Fauran) Trombicula (Tragardula) ! sinnamaryi Floch and Fauran, 1956, Arch. Inst. Pasteur Guyane Franc, et Inini, no. 405, pp. 3-7, figs. 1-5. About 200 specimens identified from 47 lots. Hosts : BIRDS, Anhinga anhinga, Aramides cajanea, Ciccaba virgata, Otus guatemalae, Caprimulgus rufus, Nyctidromus albicollis, Trogon massena, Baryphthengus ruficapillus, Bucconidae sp., Monasa morphoeus, Catharus mexicanus, Dendrocincla homochroa, Dumetella carolinensis , Dysithamnus mentalis, Geothlypis semi- ftava, Hylocichla ustulata, Icterus mesomelas, Microbates cinereiventris, Myiarchus ferox, Myiozetetes sp., Oporornis formosus, Piranga rubra, Sclerurus guatemalensis, Sporophila aurita corvina, Taraba major, Xiphorhynchus guttatus; MAMMALS, Phyllostomus hastatus. Localities: Juan Mina, Nuevo Emperador, Cacao Plantation, and Rod- man Naval Base (Canal Zone) ; Madden Airstrip and Pacora (Panama) ; Almirante and Rio Changena (Bocas del Toro) ; Coiba Island. From 1956 to 1962, taken in all months of the year except March, April, May, and July. Blankaartia velascoi (Boshell and Kerr) Trombicula velascoi Boshell and Kerr, 1942, Rev. Acad. Colomb. Cienc. Exacta, Fisico- Quim. y Nat., 5, (17), pp. 9-10 (in reprint). Described from 8 adults from Colombia. The only known larval records of this species are those of Michener (1946) from Panama. Michener also gives interesting biologic findings, e.g., the collecting of 4 larvae from the branches of a mango tree, and lists numerous bird hosts. Blankaartia wetmorei, new species. Figure 29. DIAGNOSIS : Distinguished from all other species of Blankaartia by the extremely long tarsala I (about 60 /t), and from all except marui n. sp. by the nude palpal genual and tibial setae and the bifurcate palpal tibial claw. DESCRIPTION : Idiosoma. — Elliptical, red in life. Length and width of holotype 650 by 425 /u. Eyes 2/2, in a plate. Anus at fourth row of ventral setae. Gnathosoma. — Punc- tate. Chelicerae and cheliceral blades elongate, the latter with tricuspid cap. Galeal seta BRENNAN AND YUNKER : CHIGGERS 251 nude. Palpal setae B/N/NNN : tarsus with six setae (four branched and two apparently nude) a subterminala and a tarsala; claw bifurcate. Scutum. — Densely punctate, with broadly curved posterior angle. Setae with appressed barbs, AM and AL'S posterior to margin. Sensillae branched. Measurements of holotype: AW 93, PW 100, SB 37, ASB 42, PSB 36, AP 35, AM 42, AL 40, PL 57, S 50 /*. Legs. — Punctate. Specialized setae as figured, tarsala I extremely long. Non-specialized setae long and with few branches. Body setae. — Dorsal setae with appressed barbs, 40 to 55 /u, arranged 2-6-6-4-2. Ventral setae, 2-2 sternals plus 16. TYPE MATERIAL : Holotype and 3 paratypes, RML no. 43234, off Nyc- tanassa violacea, Fort Kobbe (Canal Zone), 20 July 1961, Frank Todd, col- lector. Holotype in the collection of the Rocky Mountain Laboratory. Para- types in the Rocky Mountain Laboratory and the United States National Museum. Named for Dr. Alexander Wetmore, who graciously identified all bird hosts. FIG. 29. Blankaartia wetmorei, new species. Scutum and eyes. Specialized setae of legs. Genus Trombicula Berlese (sensu lato) Trombicula Berlese, 1905, Redia, 2: 155. Type-species : Trombicula minor Berlese, 1905. KEY TO PANAMANIAN SPECIES 1. Palpal tibial claw bifurcate; parasubterminala branched dicrura Brennan and Jones Palpal tibial claw trif urcate ; parasubterminala nude 2 2. Mastitarsala III present 3 Mastitarsala III absent 13 3. With five genualae I ; on bats saccopteryx Brennan and Jones With two or three genualae 1 4 4. Galeal seta branched pecari Brennan and Jones Galeal seta nude 5 5. With two genualae 1 6 With three genualae 1 8 6. Eyes absent, palpal femur and genu flattened and with pronounced lateral angles anophthalma Hoffmann 252 ECTOPARASITES OF PANAMA Eyes present ; palpal femur and genu normal 7 7. Dorsal formula begins 2-6 dunni Ewing Dorsal formula begins 2-8 cribanus Brennan and Jones 8(5). All palpal tibial setae nude; genuala III unusually long; on bats vesperuginis Brennan and Jones Not all palpal tibial setae nude ; genuala III of normal length 9 9. Dorsal formula begins 2-4 or 4-4 keenani Brennan and Jones Dorsal formula begins 2-6 10 10. Palpal tarsus with five branched setae ; sensillae with two long branches ; eyes not in a plate ; on bats carmenae Brennan and Jones Palpal tarsus with seven branched setae; sensillae with several branches of moderate length ; eyes in a plate 11 11. Posterior scutal margin sinuous; AL'S considerably behind margin of scutum; on bats soucouyanti n. sp. Posterior scutal margin broadly rounded ; AL'S not set far behind scutal mar- gin ; on rodents 12 12. Palpal laterotibial seta branched; scutum sparsely punctate; scutal setae slender, with semi-appressed branches ; AM < AL chiriquensis Brennan and Jones Palpal laterotibial seta nude; scutum densely punctate; scutal setae thick, with appressed barbs; AM > AL caccabulus Brennan and Jones 13(2). With three pairs of sternal setae liomys Brennan and Jones With two pairs of sternal setae 14 14. Palpal femoral seta nude ; two genualae I tiptoni Brennan and Jones Palpal femoral seta branched, or at least delicately barbed ; three genualae I . . 15 15. Galeal seta branched; eyes absent; palpal femur and genu with prominent lateral angles; on bats tibbettsi Brennan and White Galeal seta nude ; eyes present ; palpal femur and genu without lateral angles . 16 16. With a single pair of eyes 17 With two pairs of eyes in plates 18 17. Sensillae nude; posterior margin of scutum nearly straight; palpal genual seta nude; on bats monops Brennan and Jones Sensillae branched; posterior margin of scutum sharply angulate; palpal genual seta branched; on rodents punctata Boshell and Kerr 18(16). Palpal ventrotibial seta branched; posterior scutal margin broadly rounded; AL'S set in anterolateral angles dasyproctae Ewing Palpal ventrotibial seta nude; posterior scutal margin sinuous; AL'S set be- hind margin 19 19. Palpal tarsala extremely long; on tree-frequenting animals longicalcar Brennan and Jones Palpal tarsala of normal length; on various animals, .manueli Brennan and Jones Trombicula anophthalma Hoffmann Trombicula anophthalma Hoffmann, 1960, Ciencia (Mex.), 20, (3-4), pp. 102-103, figs. 11-13. Euschoengastia anops Brennan and Jones, 1960, Acarologia, 2, (4) , pp. 498-500, fig. 2. New synonymy. Two specimens identified off Pteronotus parnellii, Chilibrillo Caves, Chilibre (Panama), 2 August 1960. First Panamanian record. Trombicula caccabulus Brennan and Jones Trombicula caccabulus Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 115-116, fig. 11. Forty-seven specimens identified from 13 lots. Hosts: Peromyscus BRENNAN AND YUNKER : CHIGGERS 253 nudipes, Scotinomys teguina. Localities: Bambito and Cerro Punta (Chiriqui). Active during the dry season, January to May. In addition, Brennan and Jones (1961a) report T. caccabulus from Oryzomys fulvescens and Reithrodontomys mexicanus. Trombicula carmenae Brennan and Jones Trombicula carmenae Brennan and Jones, 1960, Acarologia, 2, (4), p. 513, fig. 10. Ten specimens off Phyllostomus hastatus, Pacora (Panama), 6 June 1961; 1 off Sturnira ludovici, Bambito (Chiriqui), 24 April 1961; 1 off S. ludovici and 4 off Artibeus jamaicencis, Rio Changena (Bocas del Toro), 19 September 1961. First records for Panama. Trombicula chiriquensis Brennan and Jones Trombicula chiriquensis Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 116-117, fig. 12. Two specimens off Scotinomys teguina, Cerro Punta (Chiriqui), 26 and 28 April 1961. Brennan and Jones (1961a) also record this species from Peromyscus sp. and Reithrodontomys sp. Trombicula cribanus Brennan and Jones Trombicula cribanus Brennan and Jones, 1961, loc. cit., pp. 117-118, fig. 13. Known only from the holotype off Proechimys semispinosus, Cerro Azul (Panama), 24 January 1956. Trombicula dasyproctae Ewing Trombicula dasyproctae Ewing, 1937, Proc. Biol. Soc. Wash., 50: 172. In spite of additional examinations of the type host-species from the type locality, this chigger is still known only from the type series, 8 specimens off Dasyprocta punctata, Capira (Panama), 28 August 1931. Trombicula dicrura Brennan and Jones Trombicula dicrura Brennan and Jones, 1961, loc. cit., pp. 118-119, fig. 14. About 125 specimens identified from 32 lots. Hosts: Coendou roth- schildi, Heteromys desmarestianus, Reithrodontomys creper, R. mexicanus, Sciurus granatensis, Peromyscus nudipes, Scotinomys teguina, S. xeram- pelinus, and mouse. Localities : Pedro Miguel River (Canal Zone) ; Cerro Punta and Boquete Trail (Chiriqui). Taken in the dry season, January to May. For additional Panamanian records, see Brennan and Jones (1961a) . Trombicula dunni Ewing Trombicula dunni Ewing, 1931, Proc. U. S. Nat. Mus., 80: 12-13, fig. 2. Trombicula (Trombicula) agutii Floch and Fauran, 1957, Arch. Inst. Pasteur Guyane Franc., et Inini, no. 426, pp. 4-6, fig. 2. New synonymy. Trombicula paragoga Brennan and Jones, 1960, Acarologia, 2, (4), pp. 524-527, fig. 18. New synonymy. About 900 specimens identified from 165 lots. Hosts: BIRDS, Odonto- 254 ECTOPARASITES OF PANAMA phorus erythrops, Leptotila cassinii, Neomorphus geoffroyi salvini, Micro- bates cinereiventris; MAMMALS, Didelphis marsupialis, Marmosa robinsoni, Metachirus nudicaudatus, Philander opossum, Vampyressa pusilla, Dasypus novemcinctus, Bradypus infuscatus, Proechimys semispinosus, Hoplomys gymnurus, Heteromys desmarestianus, H. australis, Dasyprocta punctata, Sciurus granatensis, Microsciurus alfari, Sigmodon hispidus, Oryzomys caliginosus, O. capita, Peromyscus flavidus, P. nudipes, Sylvilagus brasilien- sis, Nasua nasua, Felis pardalis, Tayassu tajacu. Localities : Barro Colorado Island, Fort Davis, Fort Gulick, Fort Sher- man, France Field, Galeta Point, Pina, Paraiso, Summit, Madden For- est, Cacao Plantation, Miraflores, Fort Kobbe, Quarry Heights, and Corte Culebra Road (Canal Zone) ; Cerro Azul and Cerro Campana (Panama) ; Cerro Pirre (Darien) ; Rio Changena, etc. (Bocas del Toro) ; Camp Pital, etc. (Chiriqui) ; Cerro Hoya (Los Santos) . Active in the dry season. From 1953 to 1963, 95 percent of all collections made were from December to April, the remaining 5 percent sporadically during June, September, October, and November. T. dunni is a very common polymorphic species parasitizing a wide va- riety of vertebrates in the American tropics. At least three forms are ap- parent, but because of intergradation observed in large series and the rela- tive absence of biologic data, any consideration of this taxon other than in terms of population variants is not feasible. Principal differences among the forms are in length, and structure and number of idiosomal setae. Bren- nan and Jones (1960) redescribed the species, included new synonymy and designated a lectotype. Trombicula keenani Brennan and Jones Trombicula keenani Brennan and Jones, 1961, Jour. Parasit., 47, (1), pp. 119-120, fig. 15. One hundred and thirty-five specimens identified from 38 lots. Hosts : Didelphis marsupialis, Philander opossum, Heteromys australis, H. des- marestianus, Proechimys semispinosus, Coendou mexicanus, Peromyscus nudipes, Reithrodontomys creper, R. mexicanus, Scotinomys teguina, Ory- zomys capito. Localities: Pina (Canal Zone) ; Cerro Pirre (Darien) ; Boquete Trail, Cerro Punta, Bambito, and El Hato (Chiriqui). From 1954 to 1962, taken principally during the dry season, November to May. In addition, Brennan and Jones (1961a) report T. keenani from Hoplomys gymnurus and Sciurus granatensis. Trombicula liomys Brennan and Jones Trombicula liomys Brennan and Jones, 1961, loc. cit., p. 121, fig. 16. Two specimens off Liomys adspersus, Corte Culebra Road (Canal Zone) , 28 September 1961. Described from 4 specimens off same host, Curundu (Canal Zone) , 8 July 1954 and Corozal (Canal Zone) , 21 October 1954. No other records. BRENNAN AND YUNKER: CHIGGERS 255 Trombicula longicalcar Brennan and Jones Trombicula longicalcar Brennan and Jones, 1960, Acarologia, 2, (4), pp. 517-518, fig. 13. Five specimens off Didelphis marsupialis, Summit (Canal Zone) , 23 Jan- uary 1962 ; 17 off (4) Coendou rothschildi, along Pedro Miguel River (Canal Zone), 21 February to 20 March 1962; 9 off unidentified bat, Las Palmitas (Los Santos), 27 January 1962; 6 off Vampyrum spectrum, Rio Changena (Bocas del Toro), 23 September 1961. First records for Panama. Trombicula manueli Brennan and Jones Trombicula manueli Brennan and Jones, 1960, loc. cit., pp. 520-522, fig. 15. Three specimens off Marmosa robinsoni, Fort Gulick (Canal Zone), 24 March 1961; 2 off Didelphis marsupialis, Pina (Canal Zone), 28 December 1961; 12 off Sciurus granatensis, Cerro Punta (Chiriqui), 2 May 1961. First Panamanian records. The form from the Chiriqui highlands is a variant and may be a sub- species. It differs from the typical form in having a branched palpal ventro- tibial seta and more dorsal setae. Trombicula monops Brennan and Jones Trombicula monops Brennan and Jones, 1960, loc. cit., pp. 524-525, fig. 17. One specimen off Myotis nigricans, from a cave, Cerro Punta (Chiriqui) , 3 May 1961; 3 off undetermined bat, Penonome (Code), 30 January 1962; 1 off Pteronotus psilotis, Penonome, 8 February 1962. First Panamanian records. Trombicula pecari Brennan and Jones Trombicula pscari Brennan and Jones, 1960, loc. cit., pp. 527-528, fig. 19. No additional records since described from 21 specimens off Tayassu tajacu, Chiriqui Province, no date, and 1 specimen (not of the type series) off Bradypus griseus (=B. infuscatus) , Chilibre (Panama), no date. Trombicula punctata Boshell and Kerr Trombicula (s.l.) punctata Boshell and Kerr, 1942, Rev. Acad. Colomb. Cienc. Exacta, Fisico-Quim. y Nat., 5, (17), pp. 15-16 (in reprint), figs. 23, 24. Twenty-one specimens off Hoplomys gymnurus, Pifia (Canal Zone), 6 December 1960; 1 off same host, Rio Changena (Bocas del Toro), 23 Sep- tember 1961. First records for Panama. Fauran (1959) described a very closely related species, tuberculata, which in all likelihood is a form of punctata. On the basis of a pentagonal scutum with a very sharp posterior angle, a single pair of eyes, and body setae arising from tubercles, he also erected a subgenus, Boshkerria — type- species, Trombicula (Boshkerria) tuberculata — to receive these two forms. Trombicula saccopteryx Brennan and Jones Trombicula saccopteryx Brennan and Jones, 1960, loc. cit., pp. 530-531, fig. 21. 256 ECTOPARASITES OF PANAMA Ten specimens off Saccopteryx bilineata, Pacora (Panama), 24 April 1957 ; 27 specimens, same host and locality, 31 October 1960 ; 13 off same host, Bocas del Toro Province, 24 January 1960; 10 off same host, Cacao Plantation (Canal Zone) , 5 February 1962. First Panamanian records. Trombicula soucouyanti, new species. Figure 30. DIAGNOSIS : Superficially resembles T. manueli Brennan and Jones, a com- mon Caribbean species not known from bats, from which it is separated by a mastitarsala III and branched palpal dorso- and ventrotibial setae. FIG. 30. Trombicula soucouyanti, new species. Scutum and eyes. Specialized setae of legs. DESCRIPTION : Idiosoma. — Broad-ovate. Length and width of holotype, engorged, 635 by 450 p.. Eyes 2/2, in an obscure plate, the posterior pair obsolescent. Anus at about fourth row of ventral setae. Scutum. — As figured, densely punctate with sinuous mar- gins. Sensillae with several long branches. Measurements of holotype : AW 72, PW 84, SB 30, ASB 33, PSB 19, AP 24, AM 53, AL 33, PL 75, s 73 /*• Gnathosoma. — Densely punctate. Cheliceral blades with tricuspid cap. Galeal seta nude. Palpal setae B/B/BNB (latero- tibial may be forked) ; tarsus with seven branched setae, a subterminala, and a tarsala; claw trif urcate. Legs. — Densely punctate. Specialized setae as figured. Non-specialized setae with long conspicuous branches. Body setae. — Dorsal setae similar to scutals, with semi-appressed branches, 44 to 66 n, arranged 2-6-6-4-2-2-2. Ventral setae, 2-2 plus 20 to 26. TYPE MATERIAL : Holotype and 7 paratypes, RML no. 40711, off Sturnira ludovici, Cerro Punta (Chiriqui), 24 April 1961, MARU. Holotype in the collection of the Rocky Mountain Laboratory. Paratypes in the Rocky Mountain Laboratory, United States National Museum, British Museum (Natural History) , and the Chicago Natural History Museum. ADDITIONAL MATERIAL: Two specimens from Myotis sp., same general locality as type series, 5 May 1961, obviously conspecific, but with posterior area of scutum less densely punctate, anterior angles of scutum less abrupt, and reduced AP (17 /*) . BRENNAN AND YUNKER: CHIGGERS 257 Trombicula tibbettsi Brennan and White Trombicula tibbettsi Brennan and White, 1960, Jour. Parasit., 46, (3), pp. 348-350, fig. 3. A single specimen off Pteronotus suapurensis, Chilibrillo Caves, Chilibre (Panama), 2 August 1960. First Panamanian record. Trombicula tiptoni Brennan and Jones Trombicula tiptoni Brennan and Jones, 1961, Jour. Parasit., 47, (1), p. 122, fig. 17. One specimen off Peromyscus nudipes, Boquete Trail (Chiriqui), 3 May 1961. Otherwise known only from the holotype, off the same host species from the same general locality, 12 February 1960. Trombicula vesperuginis Brennan and Jones Trombicula vesperuginis Brennan and Jones, 1960, Acarologia, 2, (4) pp. 533-535, fig 23. One specimen off Carollia perspicillata, Sardanillo Caves, Summit (Ca- nal Zone), 12 August 1961 ; 11 off same host, Barro Colorado Island, (Canal Zone), 13 February 1957; 1 off Glossophaga soricina, Rio Changena (Bocas del Toro) , 17 September 1961 ; 3 off Vampyrum spectrum, Rio Changena, 23 September 1961 ; 10 off G. soricina, Cerro Hoya (Los Santos) , 10 February 1962 ; 32 off (7) unidentified bats, Los Santos Province, 16 February 1962. Genus Vanidicus Brennan and Jones Vanidicus Brennan and Jones, 1961, Jour. Parasit., 47, (1), p. 123. Type-species: Vanidicus tricosus Brennan and Jones, 1961. Vanidicus tricosus Brennan and Jones Vanidicus tricosus Brennan and Jones, 1961, loc. cit., p. 123, fig. 18. One specimen off Heteromys desmarestianus , Pina (Canal Zone), 7 De- cember 1960. Described from 4 specimens off Liomys adspersus, Curundu and Summit Garden (Canal Zone), 8 July and 21 September 1954. Genus Vergrandia Yunker and Jones Vergrandia Yunker and Jones, 1961, Jour. Parasit., 47, (6), p. 996. Type-species : Vergrandia galei Yunker and Jones, 1961. Vergrandia galei Yunker and Jones Vergrandia galei Yunker and Jones, 1961, loc. cit., pp. 997-998, pi. II. Described from 3 specimens off Pteronotus parnellii ( = Chilonycteris rubiginosa), Chilibrillo Caves, Chilibre (Panama), 2 August 1960. No ad- ditional records. Abstract Seventy-six species of chiggers (16 new), distributed among 29 genera (5 new) are recorded. New genera are: Blix, Crotonasis, Inter cutestrix, Myxacarus, and Vargatula. New species and their type hosts are: Vargatula hispida off Dasypus novemcinctus; Euschoengastia belgicae off Heteromys desmarestianus; Euschoengastia enhebra off Tylomys watsoni; Pseudoschoengastia apista off Dasypus novemcinctus; Pseudoschoen- gastia dasypi off Dasypus novemcinctus; Pseudoschoengastia finitima off Heteromys 258 ECTOPARASITES OF PANAMA desmarestianus ; Pseudoschoengastia mermeriza off Coendou rothschildi, Neoschoengastia electron off Electron platyrhynchum ; Polylopadium confirmatum off Oryzomys caligi- nosus; Blix cabassoi, intranasal, off Cabassous centralis; Myxacarus oscillatus, intra- nasal, off Proechimys semispinosus ; Alexfainia munozi, intranasal, off Pteronotus psilotis; Crotonasis fissa, intranasal, off Liomys adspersus; Blankaartia marui off Nycticorax nycticorax; Blankaartia wetmorei off Nyctanassa violacea; Trotnbicula soucouyanti off Sturnira ludovici. Keys to the genera and species and a classified host list are given. Euschoengastia anops Brennan and Jones is synonymized under Trombicula anoph- thalma Hoffmann; Trombicula agutii Floch and Fauran and T. paragoga Brennan and Jones are synonymized under T. dunni Ewing. New combinations are the transfer of Trombicula arremonops Brennan and Jones to Blankaartia, Euschoengastia precaria Brennan and Dalmat to Perissopalla, and Eu- schoengastia tryssa Brennan and Jones to Intercutestrix. The description of the genus Polylopadium Brennan and Jones has been expanded and additional descriptions and/or illustrations are given for Polylopadium kramisi Brennan and Jones and Perissopalla precaria (Brennan and Dalmat). HOST-PARASITE LIST Class REPTILIA Order SQUAMATA Suborder IGUANIDAE Ameiva bifrontata Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Ameiva festiva Eutrombicula goeldii (Oudemans) Ameiva undulata Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Anolis sp. Eutrombicula alfreddugesi (Oudemans) Sceloporus sp. Eutrombicula alfreddugesi (Oudemans) Pseudoschoengastia bulbifera Brennan Spilotes pullatus Eutrombicula alfreddugesi (Oudemans) lizards Eutrombicula alfreddugesi (Oudemans) Suborder SERPENTES Oxybelis sp. Eutrombicula alfreddugesi (Oudemans) Pseustes poecilonotus Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Class AVES Order PELECANIFORMES Anhinga anhinga Blankaartia sinnamaryi (Floch and Fauran) Order CICONIIFORMES Nyctanassa violacea Blankaartia wetmorei n. sp. Nycticorax nycticorax Blankaartia marui n. sp. Order FALCONIFORMES Caracara plancus Eutrombicula batatas (Linnaeus) Hypomorphnus urubitinga Eutrombicula batatas (Linnaeus) Order GALLIFORMES Gallus gallus Eutrombicula batatas (Linnaeus) Odontophorus erythrops Eutrombicula alfreddugesi (Oudemans) Odontacarus fieldi Brennan and Jones Trombicula dunni Ewing Order GRUIFORMES Aramides cajanea Blankaartia sinnamaryi (Floch and Fauran) BRENNAN AND YUNKER I CHIGGERS 259 Order COLUMBIFORMES Leptotila cassinii Trombicula dunni Ewing Order CucuLiFORMES Crotophaga ani Eutrombicula batatas (Linnaeus) Neomorphus geoffroyi salvini Eutrombicula goeldii (Oudemans) Odontacarus fieldi B. & J. Trombicula dunni Ewing Order STRIGIFORMES Ciccaba virgata Blankaartia sinnamaryi (Floch and Fauran) Otus guatemalae Blankaartia sinnamaryi (Floch and Fauran) Order CAPRIMULGIFORMES Caprimulgus rufus Blankaartia sinnamaryi (Floch and Fauran) Nyctidromus albicollis Eutrombicula batatas (Linnaeus) Blankaartia sinnamaryi (Floch and Fauran) Order TROGONIFORMES Trogon massena Blankaartia sinnamaryi (Floch and Fauran) Order CORACIIFORMES Electron platyrhynchum Neoschoengastia electron n. sp. Baryphthengus ruficapillus Blankaartia sinnamaryi (Floch and Fauran) Momotus momota Eutrombicula goeldii (Oudemans) Order PICIFORMES Bucconidae, undet. sp. Blankaartia sinnamaryi (Floch and Fauran) Malacoptila panamensis Eutrombicula goeldii (Oudemans) Monasa morphoeus Blankaartia sinnamaryi (Floch and Fauran) Order PASSERIFORMES Dendrocincla homochroa Blankaartia sinnamaryi (Floch and Fauran) Xiphorhynchus guttatus Blankaartia sinnamaryi (Floch and Fauran) Sclerurus guatemalensis Blankaartia sinnamaryi (Floch and Fauran) Taraba major Eutrombicula alfreddugesi (Oudemans) Blankaartia sinnamaryi (Floch and Fauran) l)\ sithamnus mentalis Blankaartia sinnamaryi (Floch and Fauran) Myiozetetes sp. Eutrombicula batatas (Linnaeus) Blankaartia sinnamaryi (Floch and Fauran) Myiarchus ferox Blankaartia sinnamaryi (Floch and Fauran) Cyphorhinus aradus phaeocephalus Eutrombicula goeldii (Oudemans) Dumetella carolinensis Blankaartia sinnamaryi (Floch and Fauran) Hylocichla ustulata Blankaartia sinnamaryi (Floch and Fauran) Catharus mexicanus Eutrombicula batatas (Linnaeus) Blankaartia sinnamaryi (Floch and Fauran) Microbates cinereiventris Trombicula dunni Ewing Blankaartia sinnamaryi (Floch and Fauran) Oporornis formosus Blankaartia sinnamaryi (Floch and Fauran) Geothlypis semiflava Blankaartia sinnamaryi (Floch and Fauran) Icterus mesomelas Blankaartia sinnamaryi (Floch and Fauran) Piranga rubra Blankaartia sinnamaryi (Floch and Fauran) Sporophila aurita corvina Blankaartia sinnamaryi (Floch and Fauran) Arremonops conirostris Blankaartia arremonops (B. & J.) 260 ECTOPARASITES OF PANAMA Class MAMMALIA Order MARSUPIALIA Family Didelphidae Marmosa mexicana Eutrombicula goeldii (Oudemans) Pseudoschoengastia bulbifera Brennan Marmosa robinsoni Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Leptotrombidium panamensis (Ewing) Pseudoschoengastia bulbifera Brennan Trombicula dunni Ewing " manueli B. & J. Philander opossum Crotiscus desdentatus (Boshell and Kerr) Euschoengastia nunezi (Hoffmann) Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Pseudoschoengastia bulbifera Brennan Trombicula dunni Ewing keenani B. & J. Metachirus nudicaudatus Eutrombicula goeldii (Oudemans) Myxacarus oscillatus n. sp. Trombicula dunni Ewing Didelphis marsupialis Ascoschoengastia dyscrita B. & J. Crotiscus desdentatus (Boshell and Kerr) Euschoengastia megastyrax B. & J. nunezi (Hoffmann) tragulata B. & J. Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Hoffmannina handleyi B. & J. Leptotrombidium panamensis (Ewing) Odontacarus fieldi B. & J. Pseudoschoengastia bulbifera Brennan Sasacarus furmani (Hoffmann) Trombicula dunni Ewing keenani B. & J. longicalcar B. & J. manueli B. & J. Chironectes minimus Doloisia (Kymocta) chironectes Yunker and Brennan Order INSECTIVORA Family Soricidae Cryptotis sp. Pseudoschoengastia bulbifera Brennan Order CHIROPTERA Family Emballonuridae Saccopteryx bilineata B earner ella acutascuta Brennan Euschoengastia desmodus Brennan and Dalmat Trombicula saccopteryx B. & J. Peropteryx macrotis Tecomatlana sandovali Hoffmann Family Phyllostomidae Pteronotus parnellii Alexfainia chilonycteris Yunker and Jones Trombicula anophthalma Hoffmann Vergrandia galei Yunker and Jones Pteronotus psilotis Alexfainia munozi n. sp. Perates insessus Brennan and Dalmat Trombicula monops B. & J. Pteronotus suapurensis Trombicula tibbettsi Brennan and White Micronycteris megalotis Beamerella acutascuta Brennan Euschoengastia desmodus Brennan and Dalmat Perissopalla precaria (Brennan and Dalmat) Phyllostomus hastatus Blankaartia sinnamaryi (Floch and Fauran) Trombicula carmenae B. & J. Vampyrum spectrum Trombicula longicalcar B. & J. vesperuginis B. & J. Glossophaga soricina Euschoengastia desmodus Brennan and Dalmat Trombicula vesperuginis B. & J. Carollia castanea Euschoengastia desmodus Brennan and Dalmat Carollia perspicillata Alexfainia chilonycteris Yunker and Jones Beamerella acutascuta Brennan Euschoengastia megastyrax B. & J. Trombicula vesperuginis B. & J. Carollia subrufa Euschoengastia colombiae (Boshell and Kerr) BRENNAN AND YUNKER I CHIGGERS 261 Euschoengastia desmodus Brennan and Dalmat Sturnira ludovici Pseudoschoengastia bulbifera Brennan Trombicula carmenae B. & J. soucouyanti n. sp. Vampyressa pusilla Trombicula dunni Ewing Artibeus toltecus Leptotrombidium panamensis (Ewing) Artibeus jamaicensis Trombicula carmenae B. & J. Family Vespertilionidae Myotis nigricans Trombicula monops B. & J. Myotis sp. Trombicula soucouyanti n. sp. Family unknown Alexfainia munozi n. sp. Euschoengastia desmodus Brennan and Dalmat Euschoengastia megastyrax B. & J. Eutrombicula goeldii (Oudemans) Tecomatlana sandovali Hoffmann Trombicula longicalcar B. & J. monops B. & J. vesperuginis B. & J. Order PRIMATES Family Cebidae Aotus trivirgatus Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Family Callithricidae Saguinus geoffroyi Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Pseudoschoengastia bulbifera Brennan Family Hominidae Homo sapiens Eutrombicula batatas (Linnaeus) Order EDENTATA Family Bradypodidae Bradypus infuscatus Trombicula pecari B. & J. dunni Ewing Family Dasypodidae Cabassous centralis Blix cabassoi n. sp. Dasypus novemcinctus Aniatrus bifax B. & J. Eutrombicula goeldii (Oudemans) Myxacarus oscillatus n. sp. Pseudoschoengastia apista n. sp. dasypi n. sp. Trombicula dunni Ewing Vargatula hispida n. sp. Order LAGOMORPHA Family Leporidae Sylvilagus brasiliensis Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Trombicula dunni Ewing Order RODENTIA Family Sciuridae Sciurus granatensis Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Odontacarus fieldi B. & J. Trombicula dicrura B. & J. dunni Ewing keenani B. & J. manueli B. & J. Microsciurus alfari Eutrombicula goeldii (Oudemans) Trombicula dunni Ewing Family Heteromyidae Liomys adspersus Ascoschoengastia dyscrita B. & J. Crotonasis fissa n. sp. Eutrombicula goeldii (Oudemans) Leptotrombidium panamensis (Ewing) Odontacarus fieldi B. & J. Polylopadium kramisi B. & J. Pseudoschoengastia bulbifera Brennan zona Brennan Trombicula liomys B. & J. Vanidicus tricosus B. & J. Heteromys australis Eutrombicula goeldii (Oudemans) Leptotrombidium panamensis (Ewing) Pseudoschoengastia bulbifera Brennan zona Brennan Trombicula dunni Ewing keenani B. & J. Heteromys desmarestianus Ascoschoengastia dyscrita B. & J. Crotiscus desdentatus (Boshell and Kerr) Euschoengastia belgicae n. sp. Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Hoffmannina handleyi B. & J. 262 ECTOPARASITES OF PANAMA Doloisia (Kymocta) teratarsalis Yunker and Brennan Pseudoschoengastia bulbifera Brennan finitima n. sp. Sasacarus furmani (Hoffmann) Trombicula dicrura B. & J. " dunni Ewing " keenani B. & J. Vanidicus tricosus B. & J. Family Cricetidae Oryzomys albigularis Pseudoschoengastia bulbifera Brennan Oryzomys alfaroi Eutrombicula alfreddugesi (Oudemans) Pseudoschoengastia bulbifera Brennan Oryzomys bombycinus Leptotrombidium panamensis (Ewing) Pseudoschoengastia bulbifera Brennan Oryzomys caliginosus Ascoschoengastia dyscrita B. & J. Eutrombicula goeldii (Oudemans) Intercutestrix tryssa (B. & J.) Leptotrombidium panamensis (Ewing) Polylopadium confirmatum n. sp. Pseudoschoengastia bulbifera Brennan zona Brennan Trombicula dunni Ewing Oryzomys capito Ascoschoengastia dyscrita B. & J. Euschoengastia cunctata B. & J. Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Intercutestrix tryssa (B. & J.) Myxacarus oscillatus n. sp. Pseudoschoengastia abditiva Brennan bulbifera Brennan Trombicula dunni Ewing keenani B. & J. Leptotrombidium, panamensis (Ewing) Oryzomys fulvescens Trombicula caccabulus B. & J. Oryzomys sp. Eutrombicula alfreddugesi (Oudemans) Pseudoschoengastia bulbifera Brennan Nectomys alfari Crotiscus desdentatus (Boshell and Kerr) Eutrombicula goeldii (Oudemans) Pseudoschoengastia bulbifera Brennan Tylomys watsoni Ascoschoengastia dyscrita B. & J. Euschoengastia enhebra n. sp. Intercutestrix tryssa (B. & J.) Kymocta teratarsalis Yunker and Brennan Pseudoschoengastia zona Brennan Sasacarus furmani (Hoffmann) Reithrodontomys creper Hoffmannina suriana (Hoffmann) Trombicula dicrura B. & J. keenani B. & J. Reithrodontomys mexicanus Eutrombicula alfreddugesi (Oudemans) Hoffmannina handleyi B. & J. Trombicula caccabulus B. & J. dicrura B. & J. keenani B. & J. Reithrodontomys sumichrasti Hoffmannina suriana (Hoffmann) Reithrodontomys sp. Eutrombicula batatas (Linnaeus) Hoffmannina handleyi B. & J. Pseudoschoengastia bulbifera Brennan Trombicula chiriquensis B. & J. Peromyscus flavidus Eutrombicula alfreddugesi (Oudemans) Trombicula dunni Ewing Peromyscus nudipes Cordiseta mexicana (Hoffmann) Euschoengastia libertatis Brennan and Dalmat spissa B. & J. Eutrombicula alfreddugesi (Oudemans) Hoffmannina handleyi B. & J. suriana ( Hoffman ) Pseudoschoengastia bulbifera Brennan Trombicula caccabulus B. & J. dicrura B. & J. dunni Ewing keenani B. & J. tiptoni B. & J. Peromyscus sp. Polylopadium kramisi B. & J. Pseudoschoengastia bulbifera Brennan Trombicula chiriquensis B. & J. Zygodontomys microtinus Eutrombicula alfreddugesi (Oudemans) batatas (Linnaeus) goeldii (Oudemans) Leptotrombidium panamensis (Ewing) Odontacarus fieldi B. & J. Pseudoschoengastia bulbifera Brennan Scotinomys teguina Cordiseta mexicana (Hoffmann) Eutrombicula alfreddugesi (Oudemans) Hoffmannina handleyi B. & J. Trombicula caccabulus B. & J. chiriquensis B. & J. dicrura B. & J. " keenani B. & J. BRENNAN AND YUNKER : CHIGGERS 263 Scotinomys xerampelinus Eutrombicula alfreddugesi (Oudemans) Hoffmannina suriana (Hoffmann) Pseudoschoengastia bulbifera Brennan Trombicula dicrura B. & J. Sigmodon hispidus Ascoschoengastia dyscrita B. & J. Crotiscus desdentatus (Boshell and Kerr) Eutrombicula alfreddugesi (Oudemans) " batatas (Linnaeus) " goeldii (Oudemans) Intercutestrix tryssa (B. & J.) Leptotrombidium panamensis (Ewing) Odontacarus fieldi B. & J. Pseudoschoengastia bulbifera Brennan zona Brennan Trombicula dunni Ewing Family Muridae Rattus rattus Eutrombicula goeldii (Oudemans) Family Erethizontidae Coendou mexicanus Trombicula keenani B. & J. Coendou rothschildi Euschoengastia tragulata B. & J. Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Leptotrombidium panamensis (Ewing) Pseudoschoengastia mermeriza n. sp. Speleocola secunda B. & J. Trombicula dicrura B. & J. longicalcar B. & J. Family Hydrochaeridae Hydrochaeris hydrochaeris Blankaartia alleei (Ewing) Eutrombicula alfreddugesi (Oudemans) batatas (Linnaeus) Family Dasyproctidae Agouti paca Eutrombicula alfreddugesi (Oudemans) Dasyprocta punctata Eutrombicula goeldii (Oudemans) Trombicula dasyproctae Ewing dunni Ewing Proechimys semispinosus Ascoschoengastia dyscrita B. & J. Crotiscus desdentatus (Boshell and Kerr) Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Intercutestrix tryssa (B. & J.) Leptotrombidium panamensis (Ewing) Myxacarus oscillatus n. sp. Odontacarus chiapanensis (Hoffmann) fieldi B. & J. Polylopadium kramisi B. & J. Pseudoschoengastia bulbifera Brennan Sasacarus furmani (Hoffmann) Trombicula cribanus B. & J. dunni Ewing keenani B. & J. Hoplomys gymnurus Ascoschoengastia dyscrita B. & J. Crotiscus desdentatus (Boshell and Kerr) Eutrombicula goeldii (Oudemans) Doloisia (Kymocta) teratarsalis Yunker and Brennan Leptotrombidium panamensis (Ewing) Myxacarus oscillatus n. sp. Pseudoschoengastia bulbifera Brennan Trombicula dunni Ewing keenani B. & J. punctata Boshell and Kerr mouse Eutrombicula alfreddugesi (Oudemans) Hoffmannina handleyi B. & J. Trombicula dicrura B. & J. Order CARNIVORA Family Procyonidae Nasua nasua Euschoengastia tragulata B. & J. Eutrombicula alfreddugesi (Oudemans) goeldii (Oudemans) Trombicula dunni Ewing Family Felidae Felis pardalis Eutrombicula goeldii (Oudemans) Odontacarus fieldi B. & J. Trombicula dunni Ewing Order ARTIODACTYLA Family Tayassuidae Tayassu tajacu Eutrombicula alfreddugesi (Oudemans) Trombicula dunni Ewing pecari B. & J. Family Cervidae Mazama americana Eutrombicula batatas (Linnaeus) References BERLESE, A. 1905. Acari nuovi. Manipulus IV. (Acaridi Giava). Redia, 2: 154-176, pis. XV- XVII. BOSHELL, J., AND KERR, J. A. 1942. Veinticinco especies nuevas de Trombidiideos en Colombia. Rev. Acad. Colomb. Cienc. Exacta, Fisico-Quim. y Nat., 5: 110-127 (3-38 in reprint) pis. VII. BRENNAN, J. M. 1958. Beamerella and Kaaia, two new genera of chiggers from South America. (Acarina, Trombiculidae). Jour. Kans. Ent. Soc., 31: 71-75, 2 text figs. 1960. Eight new species of Pseudoschoengastia from Mexico and Panama with a re- vised key to species (Acarina, Trombiculidae). Acarologia, 2: 480-492, 8 text figs. BRENNAN, J. M., AND DALMAT, H. T. 1960. Chiggers of Guatemala (Acarina, Trombiculidae). Ann. Ent. Soc. Amer. 53: 183-191, 9 text figs. BRENNAN, J. M., AND JONES, E. K. 1959. Keys to the chiggers of North America with synonymic notes and descriptions of two new genera (Acarina, Trombiculidae). Ann. Ent. Soc. Amer. 52: 7-16, 3 text figs. 1960. Chiggers of Trinidad, B. W. I. (Acarina, Trombiculidae). Acarologia, 2: 494- 539, 23 text figs. 1961a. New genera and species of chiggers from Panama (Acarina, Trombiculidae). Jour. Parasit., 47: 105-124, 18 text figs. 1961b. Chiggers of Peru (Acarina: Trombiculidae). Acarologia, 3: 172-205, 18 text figs. BRENNAN, J. M., AND WHITE, J. S. 1960. New records and descriptions of chiggers (Acarina, Trombiculidae) on bats in Alabama. Jour. Parasit., 46: 346-350, 3 text figs. EWING, H. E. 1925. A contribution to our knowledge of the taxonomy of chiggers including the de- scriptions of a new genus, six new species and a new variety. Amer. Jour. Trop. Med., 5: 251-265, 4 text figs. 1926. Key to the known adult Trombiculas (adults of chiggers) of the New World with descriptions of two species (Acarina, Trombidoidea). Ent. News, 37: 111- 113. 264 BRENNAN AND YUNKER I CHIGGERS 265 1929a. Four new species of chiggers (Acarina, Trombidiidae ) . Ent. News, 40: 294- 297. 1929b. A manual of external parasites. Springfield, Illinois, Charles C. Thomas. xiv + 225 pp., 96 text figs. 1931. A catalogue of the Trombiculinae or chigger mites of the New World, with new genera and species and a key to the genera. Proc. U. S. Nat. Mus., 80: 1-19, 3 pis. 1933. Three new chigger mites of the genus Trombicula from Panama with a key to the known adults of Trombicula of the New World. Proc. U. S. Nat. Mus., 82: 1-6 3 text figs. 1937. New species of mites of the subfamily Trombiculinae with a key to the New World larvae of the akamushi group of the genus Trombicula. Proc. Biol. Soc. Wash., 50: 167-173. 1938. A key to the genera of chiggers (mite larvae of the subfamily Trombiculinae) with descriptions of new genera and species. Jour. Wash. Acad. Sci., 28: 288-295, 1 text fig. 1944. Notes on the taxonomy of the trombiculid mites. Proc. Biol. Soc. Wash., 57: 101-104. 1946. Notes on the taxonomy of three genera of trombiculid mites (chigger mites), together with the description of a new genus. Proc. Biol. Soc. Wash., 59: 69-71, 1 text fig. FAIRCHILD, G. B. 1943. An annotated list of the blood-sucking insects, ticks and mites known from Panama. Amer. Jour. Trop. Med., 23: 569-591. FAURAN, P. 1959. Nouveaux Trombiculidae de Guyane franchise. Arch. Inst. Pasteur Guyane franc., et Inini, no. 457, pp. 3-15, 4 figs. 1960. Description de quatre nouvelles especes et d'une nouvelle sous-espece de Trom- biculides de Guyane franchise. Ibidem, no. 459, pp. 2-20, figs. FLOCH, H., AND FAURAN, P. 1956. Sur deux especes du genre Trombicula (Acariens, Trombiculides) nouvelles pour la faune de la Guyane Franchise: Trombicula alfreddugesi (Oudemans, 1910) et Trombicula sinnamaryi n. sp. Ibidem, no. 405, pp. 1-7, 5 figs. 1957. Description de deux nouvelles especes du genre "Trombicula" (Acariens, trombiculides) de Guyane franchise. HOFFMANN, A. 1944. Neoschoengastia nunezi n. sp. (Acarina: Trombidiidae). Rev. Inst. Salub. Enferm. Trop., 5: 221-225, 4 text figs. 1947. Un nuevo genero de trombidido mexicano. An Esc. Nac. Cien. Biol. Mex., 4: 451-457, 6 text figs. 1948. Dos especies nuevos de trombiculidos mexicanos. Rev. Inst. Salub. Enferm. Trop., 9: 177-189, pis. 1-4. 1954. Contribuciones al conocimiento de los trombiculidos mexicanos (Acarina, Trombiculidae). (6a. parte.) An. Esc. Nac. Cien. Biol. Mex., 8: 17-30, 20 text figs. 1960. Contribuciones al conocimiento de los trombiculidos mexicanos (Acarina : Trom- bicul.) (8a. parte.) Ciencia (Mexico), 20: 99-105, 18 text figs. LINNAEUS, C. 1758. Systema Naturae (10th ed.). Holmiae, L. Salvii. 1, ii + 824 pp. [Genus 235, species 22, page 617]. LIPOVSKY, L. J. 1951. A new genus of Walchiinae (Acarina, Trombiculidae). Jour. Kans. Ent. Soc., 24:95-102, 3 pis. 266 ECTOPARASITES OF PANAMA 1952. A new genus and species of chigger mite. (Acarina: Trombiculidae). Ibidem, 25: 132-137, figs. 1-12. MlCHENER, C. D. 1946. The taxonomy and bionomics of a new subgenus of chigger mites (Acarina, Trombiculinae). Ann. Ent. Soc. Amer., 39: 431-445, 23 figs., 2 tables. NAGAYO, M., MIYAGAWA, Y., MITAMURA, T., AND IMAMURA, A. 1916. [On Trombidium and the akamushi.] Dobutsugaku Zasshi, 28: 300-313, 379- 394. OUDEMANS, A. C. 1910. Acarologische Aanteekeningen XXXIII. Ent. Ber., 3: 83-90. 1911. Acarologische Aanteekeningen XXXV. Ibidem, 3: 118-126. TRAGARDH, I. 1905. Acariden aus Agypten und dem Sudan. Zugleich ein Beitrag sur Kenntnis der Entwichlungsgeschichte der Gattungen Phytoptipalpus, Pimeliaphilus, Ptery- gosoma und Podapolipus. Results Swed. Zool. Exped. to Egypt and the White Nile, 1901, under the direction of L. A. Jagerskiold. Uppsala University. Pt. II. 124 pp. + I-IV, A-H, index (A-B), 6 pis. WHARTON, G. W., AND FULLER, H. S. 1952. A manual of the chiggers. The biology, classification, distribution, and im- portance to man of the larvae of the family Trombiculidae (Acarina). Mem. Ent. Soc. Wash., no. 4, 185 pp., 17 text figs., 4 tables. YUNKER, C. E., AND BRENNAN, J. M. 1962. Endoparasitic chiggers: II. Rediscovery of Doloisia synoti Oudemans, 1910, with descriptions of a new subgenus and two new species (Acarina: Trombicu- lidae). Acarologia, 4, (4) : 570-576, 3 text figs. YUNKER, C. E., AND JONES, E. K. 1961. Endoparasitic chiggers: I. Chiroptera, a new host order for intra-nasal chig- gers, with descriptions of two new genera and species (Acarina: Trombiculidae). Jour. Parasit., 47, (6), pp. 995-1000, 2 pis. Mallophaga of the Mammals of Panama K. C. EMERSON1 Mallophaga are obligatory external parasites and host-specific; there- fore, their distribution is dependent entirely upon distribution of the hosts. Some species of Mallophaga are restricted to a single host species, and others are restricted to a host genus or closely related genera. With few excep- tions, the same species of Mallophaga are found on a mammal species throughout its range without regard to host subspecies. In addition, many groups of mammals do not harbor Mallophaga. These factors have been considered in the compilation of this list. Species collected in Panama are marked with an asterisk. Species are also listed which should be found in Panama, but to date have not been col- lected in that country. If a host is found in Costa Rica, Panama and Colom- bia, and a parasite has been recorded from Costa Rica or Colombia, it can be found in Panama with adequate collecting. Specimens examined are in collections of the author, United States Na- tional Museum, Gorgas Memorial Laboratory, and the Chicago Natural His- tory Museum. Collections were made by John H. Paine, Alexander Wetmore, Robert M. Altman, Charles M. Keenan, Vernon J. Tipton, Eustorgio Mendez, Phyllis T. Johnson, Lloyd E. Rozeboom, William H. Komp, Lawrence H. Dunn, and others not recorded. Detailed taxonomic discussions are not included since these are available in Os Malofagos de Mamiferos published by the late Dr. F. L. Werneck in 1948 and 1950 (see References). Family Trimenoponidae Cummingsia intermedia Werneck Cummingsia intermedia Werneck, 1937, Mem. Inst. Oswaldo Cruz, 32: 70, figs. 1-6. This species is found on hosts of the genus Marmosa. Cummingsia peramydis Ferris Cummingsia peramydis Ferris, 1922, Parasitology, 14: 85, figs. 2d, 3e, 4e, 8. This species is found on hosts of the genus Monodelphis. 1 Stillwater, Oklahoma, Research Associate, United States National Museum. 267 268 ECTOPARASITES OF PANAMA * Harrisonia uncinata Ferris Harrisonia uncinata Ferris, 1922, loc. cit., p. 81, figs. 2c, 3c, 4d, 6. Collected from Proechimys semispinosus at Gamboa, Fort Clayton, Fort Davis, Fort Kobbe, Madden Dam, Summit, Cocoli, Camp Pifia, and Curundu (Canal Zone) ; Almirante (Bocas del Toro) ; and Cerro Azul (Panama). * Trimenopon hispidum (Burmeister) Gyropus hispidus Burmeister, 1838, Handb. Ent., 2: 443. Collected from laboratory guinea pigs, Cavia porcellus, at Corozal (Canal Zone) and Panama City. Family Boopidae * Heterodoxus spiniger (Enderlein) Menopon spiniger Enderlein, 1909, Denkschr. Med. Nat. Ges., Jena, 14: 80, pi. 8, figs. 4-5. Collected off dogs in Panama City. Family Gyropidae * Gliricola panamensis Werneck Gliricola panamensis Werneck, 1944, Rev. Brasil. Biol., 4: 391, figs. 1-4. Collected from Proechimys semispinosus at Madden Dam, Camp Pina, Fort Kobbe, Gamboa, Fort Clayton, Fort Davis, Curundu, and Cocoli (Canal Zone) ; and Almirante (Bocas del Toro) . Previously recorded from Bugaba (Chiriqui) off "Proechimys centralis chiriquinus" (P. semispinosus chiri- quinus) by Werneck (1948, p. 46) and from the type locality, Camp Pital (Chiriqui) off Sigmodon hispidus chiriquensis which is probably a contam- ination from P. semispinosus. * Gliricola porcelli (Schrank) Pediculus porcelli Schrank, 1781, Enum. Ins. Austr. Indig., p. 500, pi. 1, fig. 1. Collected from laboratory guinea pigs, Cavia porcellus, at Corozal (Canal Zone) and Panama City. * Gliricola sp. (near panamensis) Collected from Hoplomys gymnurus at Almirante and Cayo Agua (Bocas del Toro) , and Tacarcuna (Darien) .. Gyropus aotophilus Ewing Gyropus aotophilus Ewing, 1924, Proc. U. S. Nat. Mus., 63: 23, fig. 1. This species is found on hosts of the genus Aotus, and has been collected off A. trivirgatus elsewhere. * Gyropus ovalis Burmeister Gyropus ovalis Burmeister, 1838, Handb. Ent., 2: 443. Collected from laboratory guinea pigs, Cavia porcellus, at Corozal (Canal Zone) and Panama City. EMERSON: MALLOPHAGA 269 * Gyropus setif er Ewing Gyropus setif er Ewing, 1924, loc. cit., p. 22, fig. 10. Collected off Proechimys semispinosus at Fort Clayton, Fort Davis, Fort Kobbe, Cocoli, Madden Dam, Gamboa, Camp Pifia, Summit, Fort Gulick and Curundu (Canal Zone) ; and Almirante (Bocas del Toro). Collected off Hoplomys gymnurus at Tacarcuna (Darien), Curundu (Canal Zone), Al- mirante (Bocas del Toro), Salinas Place, Aguadulce (Code), and Cerro Azul (Panama). * Macrogyropus costalimai (Werneck) Heterogyropus costalimai Werneck, 1931, Bol. Biol. Fac. Med. Sao Paulo, 18: 21, figs. 1-3. A slide in the E. W. Stafford Collection has one specimen from Agouti paca in Panama. No specific locality is given. * Macrogyropus dicotylis (Macalister) Gyropus dicotylis Macalister, 1869, Proc. Zool. Soc. London, 1869: 420, fig. Collected from Tayassu tajacu at Gamboa (Canal Zone). It was re- corded by Werneck (1948, p. 89) off Tayassu angulatus bangsi (T. tajacu bangsi) from Porto Belo (Colon). Family Trichodectidae * Bovicola bovis (Linnaeus) Pediculus bovis Linnaeus, 1758, Syst. Nat., ed. 10, p. 611. This species has been collected off cows, in dairy herd in the Canal Zone. Bovicola caprae (Gurlt) Trichodectes caprae Gurlt, 1843, Mag. ges. Thierheilk., 9: 3, pi. 1, fig. 2. This species should be found on the goat in Panama, as it is world-wide in distribution. * Bovicola equi (Denny) Trichodectes equi Denny, 1842, Mon. Anopl. Brit., pp. 61, 191, pi. 17, fig. 7. Prior to 1941, this species was common on horses and mules maintained by the United States Army in the Canal Zone. * Bovicola limbatus (Gervais) Trichodectes limbatus Gervais, 1844, Walck. Hist. Nat. Ins., Apteres, 3: 313, pi. 48, fig. 4. Recorded by Werneck (1950, p. 62) from Capra hircus, "Mona Island" (Mono Island, Darien) . Bovicola ovis (Schrank) Pediculus ovis Schrank, 1781, Enum. Ins. Austr. Indig., p. 502, pi. 1, figs. 8-9. This is a common parasite of sheep and is world-wide in distribution. 270 ECTOPARASITES OF PANAMA Cebidicola semiarmatus (Neumann) Trichodectes semiarmatus Neumann, 1913, Arch. Parasit., 15: 611, fig. 5. This species is found on hosts of the genus Alouatta. * Eutrichophilus maximus Bedford Eutrichophilus maximus Bedford, 1939, Onderstepoort Jour. Vet. Sci., 12: 118, fig. 24. This species was described from a specimen off Coendou rothschildi taken at Gamboa (Canal Zone) . Eutrichophilus mexicanus (Rudow) Trichodectes mexicanus Rudow, 1866, Zeitschr. ges. Naturw., 27: 109, pi. 5, fig. 1. This species is found on Coendou mexicanus. Felicola felis (Werneck) Trichodectes felis Werneck, 1934, Mem. Inst. Oswaldo Cruz, 28: 282, figs. 11-14. This species has been reported from Felis concolor, Felis pardalis and Felis yagouaroundi which are found in Panama, and some other hosts of the genus Felis found elsewhere. Specimens are rarely collected, so an adequate evaluation of distribution on the hosts has not been made. * Felicola subrostratus (Burmeister) Trichodectes subrostratus Burmeister, 1838, Handb. Ent., 2: 436. This species was collected off the domestic cat in the Canal Zone and Panama City. * Geomydoecus species A large series from Macrogeomys cavator, taken at Casa Pitty, Cerro Punta (Chiriqui), represents a new species which will be described by Dr. Ronald Ward in his revision of the genus. Lymeon cummingsi Eichler Lymeon cummingsi Eichler, 1943, Mitt. Deutsch. Ent. Ges., 11: 111. This species is probably found on Bradypus infuscatus. Lymeon gastrodes (Cummings) Trichodectes gastrodes Cummings, 1916, Ann. Mag. Nat. Hist., (8), 17: 94, figs. 2-4. This species is probably found oriCholoepus hoffmanni. ? Neotrichodectes minutus (Paine) Trichodectes minutus Paine, 1912, Ent. News, 23: 439, pi. 20, fig. 4. This species is found on Mustela frenata. A specimen collected at La Amenaza, Bambito (Chiriqui) is probably this species. * Neotrichodectes pallidus (Piaget) Trichodectes pallidus Piaget, 1880, Les Pediculines, p. 405, pi. 42, fig. 9. Collected from Nasua nasua at Almirante (Bocas del Toro). EMERSON: MALLOPHAGA 271 Trichodectes barbarae Neumann Trichodectes barbarae Neumann, 1913, Arch. Parasit., 15: 616, fig. 9. This species is found on Eira barbara. * Trichodectes canis (De Geer) Ricinus canis De Geer, 1778, Mem. Hist. Ins., 7: 81, pi. 4, fig. 16. Found on dogs in the Canal Zone. Trichodectes fallax Werneck Trichodectes fallax Werneck, 1948, Malof . Mamif ., 1 : 122, figs. 159-165. This species is found on Procyon cancrivorus. * Trichodectes galictidis Werneck Trichodectes galictidis Werneck, 1934, Mem. Inst. Oswaldo Cruz, 28: 161, figs. 1-5. Recorded by Werneck (1948, p. 119) off Galictis allamandi (=."Grisonia canaster") from Pacora (Panama). * Trichodectes octomaculatus Paine Trichodectes octomaculatus Paine, 1912, Ent. News, 23: 438, pi. 20, fig. 1. Collected off Procyon lotor at Finca de P. H., Cerro Punta (Chiriqui) and from Panama without specific locality. * Trichodectes potus Werneck Trichodectes potus Werneck, 1934, Mem. Inst. Oswaldo Cruz, 28: 171, figs. 7-10. Collected off Potos flavus chiriquensis, at Camp Pina (Canal Zone) . Tricholipeurus albimarginatus (Werneck) Trichodectes albimarginatus Werneck, 1936, Mem. Inst. Oswaldo Cruz, 31 : 570, figs. 205-212. This species is found on hosts of the genus Mazama. Tricholipeurus lipeuroides (Megnin) Trichodectes lipeuroides Megnin, 1884, Naturaliste, no. 62, p. 494. This species is found on Odocoileus virginianus. Tricholipeurus parallelus (Osborn) Trichodectes parallelus Osborn, 1896, Bull. U. S. Bur. Ent, (n.s.), 5: 240, fig. 148. This species is found on Odocoileus virginianus. Suricatoecus quadraticeps (Chapman) Trichodectes quadraticeps Chapman, 1897, Ent. News, 8: 185, pi. 9, fig. This species is found on Urocyon cinereoargenteus. References WERNECK, F. L. 1948. Os Malofagos de Mamiferos. Parte 1 : Amblycera e Ischnocera (Philopteridae e parte de Trichodectidae) . Revista Brasileira de Biologia. Rio de Janeiro. 243 pp., 431 text figs. 1950. Idem. Parte II : Ischnocera (continuacao de Trichodectidae) e Rhyncophthirinae. Institute Oswaldo Cruz. Rio de Janeiro. 5+ 207 pp. 316 text figs. 272 Checklist of the Sucking Lice of Panama (Anoplura) RUPERT L. WENZEL l AND PHYLLIS T. JOHNSON 2 Fairchild (1943, p. 581) reported four species of sucking lice from Pan- ama. Since then, five others have been recorded. Eighteen are listed in the present paper, based largely on specimens collected by Lt. Colonel Robert M. Altman, Major Vernon J. Tipton, Charles M. Keenan, and other personnel of the Environmental Health Group, Dr. Conrad E. Yunker, Dr. Charles O. Handley, and personnel of the Gorgas Memorial Laboratory. The junior author, who received this material for identification and study, described two new species in an earlier paper (Johnson, 1962) on the Genus Fahren- holzia. Unfortunately, other commitments made it impossible for her to study all of the specimens collected or to prepare a critical review of the Panamanian species. The following treatment is, therefore, incomplete. Additional species will undoubtedly be obtained through further collect- ing from both wild and domesticated animals. For example, Enderleinellus hondurensis Werneck, which has been taken on various species of Sciurus in Colombia, Honduras, and Mexico, will probably be found in Panama. Likewise, Linognathus pedalis (Osborn) and L. setosus (van Olfers) may occur in Panama on dogs, and Haematopinus asini on horses. The junior author is responsible for identifications of the species pre- ceded by an asterisk ( * ) . The senior author collated the data and informa- tion on other species and prepared the manuscript. Original citations and selected references are given, but one should refer to Ferris (1919-35, 1951) and Johnson (1962) for additional synonymies, references, discussions, and more comprehensive keys. 1 Chicago Natural History Museum. 2 Division of Biological Sciences, University of California, Irvine. 273 274 ECTOPARASITES OF PANAMA Family Haematopinidae Haematopinus eurysternus (Nitsch) Pediculus eurysternus Nitsch, 1818, Germar's Mag. Ent., 3: 305. Haematopinus eurysternus Ferris, 1933, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (6), p. 448, figs. 263, 264; 1951, Mem. Pac. Coast Ent. Soc., 1: 88, figs. 39, 40. Lice from cattle in Code Province were identified as H. eurysternus by Fairchild (1943, p. 581). They may actually have been H. quadripertusus Fahrenholz. * Haematopinus suis (Linnaeus) Pediculus suis Linnaeus, 1758, Syst. Nat., 10th ed., p. 611. Haematopinus suis Ferris, 1933, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (6), p. 425, figs. 252A, 253A, 254; 1951, Mem. Pac. Coast Ent. Soc., 1: 91, figs. 41, 42. FromSus scrofa: 1 male, 2 females, Santa Rita, Anton (Code) , 14 March 1961; 3 males, 3 nymphs, Rio Indio, Arraijan (Panama), 13 June 1960, V. Barria, collector; 2 males, 1 female, and 1 nymph, Guanico (Los San- tos), 25 February 1962 (no. 004) and 2 males, 3 females, and 4 nymphs, same data (no. 007). Family Hoplopleuridae Subfamily Enderleinellinae * Enderleinellus sp. (longiceps group) One female from Sciurus granatensis, Cacao Plantation, Gamboa (Canal Zone), 26 April 1957, R. M. Altman, collector. Subfamily Hoplopleurinae *Hoplopleura hesperomydis (Osborn) Haematopinus hesperomydis Osborn, Bull. Div. Ent., U. S. Dept. Agric., (old series), 7: 26, fig. 14. Hoplopleura hesperomydis Ferris, 1921, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (2), p. 70, figs. 38, 39; 1951, Mem. Pac. Coast Ent. Soc., 1: 136. From Peromyscus nudipes: 18 males and 4 females, Lava Flow, El Hato (Chiriqui), 5000 feet elevation, 7-9 January, 1961; 13 females, Martinz Dairy, Cerro Punta (Chiriqui), 5800 feet elevation, 7-8 January 1961. From Scotinomys teguina, 1 female, Martinz Dairy, 8 January 1961. From Reithrodontomys sp., 1 female, Lava Flow, El Hato, 7-9 January 1961. Col- lected by C. M. Keenan and V. J. Tipton. Additional specimens, probably of this species, from Scotinomys and Peromyscus, have not yet been studied. * Hoplopleura hirsuta Ferris Hoplopleura hirsuta Ferris, 1916, Psyche, 23: 122, figs. 8, 9A, 10, 11B; 1921, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (2), p. 117, figs. 75, 76; 1951, Mem. Pac. Coast Ent. Soc., 1: 137. From Sigmodon hispidus: 1 nymph, probably this species, Camp Pina (Canal Zone), 30 August 1960; 2 males, 1 female, Curundu (Canal Zone), 9 December 1959; 1 male, 2 females, and 4 nymphs, Almirante (Bocas del WENZEL AND JOHNSON — ANOPLURA 275 Toro) , 26 and 29 January, 1960, collected by C. M. Keenan and V. J. Tipton ; 1 male, 1 female, Cocoli (Canal Zone) , 10 January 1957, by R. M. Altman. * Hoplopleura oryzomydis Pratt and Lane Hoplopleura oryzomydis Pratt and Lane, 1951, Jour. Parasit., 37: 141, figs. 1-3, 5. Ferris, 1951, Mam. Pac. Coast Ent. Soc., 1: 139. From Oryzomys caliginosus, 1 male, 1 female, El Valle (Code), March 1957, R. M. Altman, collector. * Hoplopleura nesoryzomydis Ferris Hoplopleura nesoryzomydis Ferris, 1921, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (2), p. 90, fig. 53A; 1951, Mem. Pac. Coast Ent. Soc., 1: 138. From "rat" (Altman K-14) , Barro Colorado Island (Canal Zone) , 4 No- vember 1956, C. Koford, collector. * Pterophthirus audax (Ferris) Hoplopleura audax Ferris, 1921, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2 (2), p. 125, figs. 82, 83. Pterophthirus audax (Ferris), 1951, Mem. Pac. Coast Ent. Soc., 1: 147. From Hoplomys gymnurus, 1 male and 1 female, Cerro Azul (Panama), 29 January 1958, R. M. Altman, collector. Subfamily Polyplacinae * Fahrenholzia fairchildi Johnson Fahrenholzia fairchildi Johnson, 1962, Ann. Ent. Soc. Amer., 55, (4), p. 419, figs. 18- 21, 32, 39. The original description of this species was based on specimens from Panama and Guatemala. The holotype, allotype, and 18 paratypes are from Heteromys desmarestiamis, Santa Fe (Veraguas). Other paratypes from the same host in Panama are from Fort Kobbe and Summit Road (Canal Zone). Numerous additional specimens were collected from the type host at Cerro Punta (Chiriqui) and from Liomys adspersus at Guanico (Los Santos). ^Fahrenholzia hertigi Johnson Fahrenholzia hertigi Johnson, loc. cit., p. 421, figs. 22, 23, 34, 40. The holotype female and 6 paratype females of this species are from Heteromys desmarestianus, Cerro Azul (Panama). An additional female paratype was taken on Zygodontomys microtinus (=cherriei) , Camp Pina (Canal Zone). * Neohaematopinus sp. (sciurinus group) From Sciurus granatensis: 1 male, 7 January 1961, and 2 females, 2 May 1961, and 2 males, 26 April, 1961, Martinz Dairy, near Cerro Punta (Chiri- qui) ; 2 males, 10 females, and 4 nymphs, Cerro Hoya (Los Santos) , 10 Feb- ruary 1962. From S. variegatoides : 5 males, 13 females, Guanico (Los Santos), February 1962. 276 ECTOPARASITES OF PANAMA * Polyplax spinulosa (Burmeister) Pediculus spinulosus Burmeister, 1839, Gen. Insect., Rhynchota, no. 8. Polyplax spinulosa Ferris, 1923, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (4), p. 187, figs. 119, 120A, D, F, H; 1951, Mem. Pac. Coast Ent., Soc., 1: 211, figs. 90, 91. From Rattus rattus, 6 males, 6 females, and 5 nymphs, Guanico (Los Santos), 5 February 1961, C. 0. Handley, collector. Family Linognathidae Solenopotes binipilosus (Fahrenholz) Linognathus binipilosus Fahrenholz, 1916, Arch. Naturg., A, 81, Heft 11, p. 11, plate, figs. 11-13. Solenopotes binipilosus Ferris, 1932, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (5), p. 131, figs. 245, 246; 1951, Mem. Pac. Coast Ent. Soc., 1: 252. Johnson, 1958, Proc. U. S. Nat. Mus., 108: 49. Recorded by Ferris (1932) from Odocoileus chiriquiensis (sic!) from the Canal Zone, L. H. Dunn, collector. Ferris (loc. cit.) regarded Linogna- thus panamensis Ewing as a synonym. Solenopotes panamensis (Ewing) Linognathus panamensis Ewing, 1927, Proc. Ent. Soc. Wash., 29: 119. Solenopotes panamensis Johnson, 1958, loc. cit., pp. 48—49. Ferris (1932, p. 131) placed L. panamensis Ewing as a synonym of S. bipinilosus (Fahrenholz). Johnson (loc. cit.), after re-examining the types of panamensis, concluded that they represented a distinct species and gave characters for separating the females from those of bipinilosus. The male of panamensis is unknown. Johnson also pointed out that there is some doubt about the true host of panamensis, since the type specimens were "from Odocoileus chiriquensis [Odocoileus virginianus chiriquensis'] (ori- gin, Panama) which died at the National Zoological Park. . . ." [Washing- ton, B.C.], and "it is highly possible that the occurrence of panamensis on Odocoileus virginianus chiriquensis was accidental, the true host being some other ungulate with which the deer had come in contact while in the zoo." Family Pediculidae Pediculus atelophilus Ewing Pediculus (Parapediculus) atelophilus Ewing, 1926, Proc. U. S. Nat. Mus., 68: 9, figs. 4A, 5; 1938, Jour. Parasit., 24: 26, figs. 2, 5b, 6b. Johnson, 1958, Proc. U. S. Nat. Mus., 108: 44. Pediculus atelophilus Ferris, 1951, Mem. Pac. Coast Ent. Soc., 1: 273. Pediculus mjobergi Ferris (part.), 1935, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2: (8), pp. 588-598, figs. 318E, 319E, 322F, 3231, 328-332. Reported by Ewing (1938) from Ateles dariensis from Panama and by Hinman (1931, p. 488) from material determined by Ewing from Ateles geoffroyi from Panama. The nomenclatorial status of P. atelophilus, P. chapini (see below) and P. lobatus Fahrenholz, 1916, is still in doubt. In 1935, Ferris placed these WENZEL AND JOHNSON — ANOPLURA 277 names as synonyms of P. mjobergi Ferris, 1916 (= P. affinis Mjoberg, 1910 nee Burmeister, 1839). However, in 1951, though he still presumed them to be conspecific, he treated them separately and assigned to atelophilus specimens "from Ateles dariensis, Cebus capuchinus, and Alouatta palliata, all from Panama from monkeys in captivity." He left the question of syn- onymy open until the types of affinis Mjoberg could be examined, though in his key he employed the name mjobergi to cover all of the forms involved. Pediculus chapini Ewing Pediculus (Parapediculus) chapini Ewing, 1926, Proc. U. S. Nat. Mus., 68: 13, figs. 2, 4b, 5 and pi. 1, figs. 3, 4; 1938, Jour. Parasit., 24: 28, figs. 4a, 5c, 6c. Johnson, 1958, loc. cit., p. 44. Pediculus mjobergi Ferris (part.), 1935, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (8), pp. 588-598. Ewing (1938) reported this species from Panama from Cebus capuchi- nus. Ferris (see above) considered chapini to be a synonym of mjobergi Ferris. Johnson (loc. cit.) , considered that chapini "probably is the same as Pediculus atelophilus Ewing, 1926. Pediculus humanus Linnaeus Pediculus humanus Linnaeus, 1758, Syst. Nat., 10th ed., p. 610. Ferris, 1935, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (8), pp. 17-62, plates 1-3, text figs. 306-327; 1951, Mem. Pac. Coast Ent. Soc., 1: 261-271, figs. 116, 117, 118, 119. Phthirus pubis (Linnaeus) Pediculus pubis Linnaeus, 1758, loc. cit., p. 611. Phthirus pubis Ferris, 1935, Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (8) , p. 603, figs. 335-337; 1951, Mem. Pac. Coast. Ent. Soc., 1: 281, figs. 122-124. Fairchild (loc. cit.) reported that physicians had observed head, body, and crab lice on humans in Panama. Because the lice of Panama are incompletely known, the following key should be used with caution, preferably in conjunction with the works cited above. KEY TO PANAMANIAN ANOPLURA [Largely adapted from Ferris, 1951] 1. Paratergal plates absent, abdomen entirely membranous except for plates asso- ciated with terminal and genital segments and rarely tergal plates in male (Linognathidae) 2 Paratergal plates on at least some abdominal segments 3 2. Apical lobes of female abdomen with a short, slender terminal portion Solenopotes panamensis (Ewing) Apical lobes of female abdomen gradually constricted into long, tapering lobes Solenopotes binipilosus (Fahrenholz) 3. Venter of abdominal segment II with a pair of small, strongly sclerotized plates Enderleinellus Fahrenholz Without such plates 4 278 ECTOPARASITES OF PANAMA 4. Paratergal plates free, at least in part, from the body and not simply forming a cap over the lateral lobes of the abdomen (Hoplopleuridae) 5 Paratergal plates never with any part free from the body wall 12 5. Second abdominal segment with its sternal plate extended laterally on each side to articulate with the corresponding paratergal plate (Hoplopleurinae) 6 Second abdominal segment with its sternal plate never thus extended laterally (Polyplacinae) 10 6. Paratergal plates of second abdominal segment each prolonged into a blade-like process which projects from the body wall Pterophthirus audax (Ferris) Paratergal plates not thus (Hoplopleura) 7 7. Ventral lobe of paratergal plates of third abdominal segment divided into two apically rounded, equal lobes 8 Not thus 9 8. Ventral lobe of paratergal plates of sixth abdominal segment divided into two equal lobes Hoplopleura oryzomydis Pratt and Lane Ventral lobe of paratergal plates of sixth abdominal segment forming a single, acute point Hoplopleura nesoryzomydis Ferris 9. Posterior apex of thoracic sternal plate sharply pointed, no setae laterally off the tergal and sternal abdominal plates Koplopleura hesperomydis (Osborn) Posterior apex of thoracic sternal plate rounded, many setae present laterally off the abdominal plates Hoplopleura hirsuta Ferris At least one of the lobes apically acute Hoplopleura hirsuta Ferris 10. Paratergal plates of second abdominal segment definitely divided longitudinally into two plates, one of which is flat on the dorsum and one on the venter, the ventral portion with a flat, raised apical free point (Fahrenholzia) 11 Paratergal plates of second abdominal segment not thus divided Polyplax spinulosa (Burmeister) 11. Antennal segments 3-5 completely coalesced; thorax not broader than head; thoracic sternal plate with a small detached portion. .Fahrenholzia hertigi Johnson Antennal segments not completely coalesced ; thorax obviously broader than head ; sternal plate of thorax without a detached apical portion Fahrenholzia fairchildi Johnson 12. Eyes conspicuous as a pair of distinct lenses accompanied by pigmentation (visible in uncleared specimens) ; derm of dorsum of abdomen not wrinkled (Pediculidae) 14 Eyes not visible, though represented by vestiges; derm of dorsum of abdomen wrinkled (Haematopinidae) 13 13. Head very short and broad, length only vary slightly shorter than width; penis of male strongly asymmetrical, strongly sclerotized and forming a large hook or barb at its base Haematopinus eurysternus (Nitsch) Head elongate, about as long as broad ; penis V- or Y-shaped Haematopinus suis (Linnaeus) 14. All legs of essentially the same size, claws slender; thorax with a distinct notal pit, sternal plate sclerotized, though its margins are not free from body; ab- domen narrower at base than at middle; spiracle enclosed within borders of paratergal plates (Pediculus) . . : 15 Anterior legs slender, with slender claws, middle and posterior legs very large and stout, with stout claws ; thorax very wide occupying the greater part of the body, without notal pit or sternal plate ; abdomen widest basally, tapering api- cally, with prominent segmentally arranged sclerotized lateral tubercles, the spiracles borne slightly removed from the base of the tubercles Phthirus pubis (Linnaeus) 15. Paratergal plates throughout clearly without evidence of lateral lobes Pediculus humanus Linnaeus Paratergal plates of fifth to sixth abdominal segments bearing strong lateral lobes, both dorsally and ventrally Pediculus mjobergi Ferris (? = P. atelophilus Ewing, P. chapini Ewing) References EWING, H. E. 1938. The sucking lice of American monkeys. Jour. Parasit., 24: 13-33, 6 figs. FAIRCHILD, G. B. 1943. An annotated list of the bloodsucking insects, ticks and mites known from Panama. Amer. Jour. Trop. Med., 23, (6), pp. 569-591. FERRIS, G. F. 1916. A catalogue and host list of the Anoplura. Proc. Calif. Acad. Sci., (4), 6: 129-213. 1919-1935. Contributions toward a monograph of the sucking lice. Stanford Univ. Publ., Univ. Ser., Biol. Sci., 2, (1-8), 634 pp., 388 text figs., 3 plates. [Part 1 (1919), pp. 1-52, figs. 1-32. Part 2 ( 1921), pp. 52-134, figs. 33-89. Part 3 (1922), pp. 135-178, figs. 90-118. Part 4 (1923), pp. 179-270, figs. 119-172. Part 5 (1932), pp. 271-414, figs. 173-251. Part 6 (1933), pp. 415-470, figs. 252-276. Part 7 (1934), pp. 471-526, figs. 277-305. Part 8 (1935), pp. 527-620, figs. 306- 388, pis. 1-3. Index of hosts, pp. 623-628. Index of parasites, pp. 629-634.] 1951. The sucking lice. Mem. Pac. Coast Ent. Soc., 1. ix + 320 pp., 124 figs. HINMAN, E. H. 1931. Pediculus (Parapediculus) atelophilus Ewing 1926 from the red spider monkey, Ateles geoffroyi. Parasitology, 23: 488-491, 2 figs. JOHNSON, P. T. 1958. Type specimens of lice (Order Anoplura) in the United States National Museum. Proc. U. S. Nat. Mus., 108: 39-49. 1962. The species of Fahrenholzia Kellogg and Ferris from spiny pocket mice. Ann. Ent. Soc. Amer., 55, (4), pp. 415-428, 8 pis. 279 New Species of the Genus Amblyopinus Solsky from Panama and Mexico (Coleoptera: Staphylinidae) ALFREDO BARRERA This paper includes descriptions of Amblyopinus tiptoni n. sp. from Northern Panama, A. isabelae n. sp. from Southern Mexico, and records of A. emarginatus Seevers, 1954, from Northern Panama. These species are interesting additions to the primitive "Andean" elements of the highlands of Southern Mexico and Central America. All were found in association with cricetine rodents, like the two closely allied species previously described from that area, namely, A. schmidti Seevers, 1944, and A. bolivari Bar- rera, Machado and Muriiz, 1960. The single male of A. isabelae n. sp. which represents the second species known from Mexico, was collected some years ago by Professor Dionisio Pelaez and myself at Omiltemi, Guerrero, Mexico (Barrera, 1958). I am very much indebted to Dr. Candido Bolivar for the gift of this specimen which had previously been deposited in his laboratory at the Escuela Na- cional de Ciencias Biologicas, Mexico, D.F. The Panamanian species constitute the first records of the genus for that country. The description of A. tiptoni n. sp. is based on a large number of specimens collected in Chiriqui Province near the Costa Rican border. I am very grateful to Lt. Col. Vernon J. Tipton, formerly Chief, Environmen- tal Health Branch, Preventive Medicine Division, United States Army Carib- bean, Canal Zone, for making the Panamanian collection available to me. It is difficult to even attempt to assess my debt to Mr. Carlos Machado, Biologist, now at the Departmento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil. Mr. Machado has been extremely kind in devoting his valuable time to checking specimens for me. His criticism about the des- ignation of abdominal sterna is greatly appreciated. 1 Laboratorio de Biologia Economica y Experimental Escuela Nacional de Cien- cias Biologicas Institute Politecnico Nacional Mexico, D.F. 281 282 ECTOPARASITES OF PANAMA The abdomen of Amblyopinus is of the haplogastrous type with a com- plete second sternum ; thus, the first externally visible sternum is actually the second. When Barrera et al. (1960) described A. bolivari, they named the abdominal sterna following a simple ordinal criterion. However, in this paper, descriptions are made using the terminology of morphologists. The first externally visible abdominal sternum is designated as the second, and so forth. Seevers (1955) says that "considerable reliance must be placed on the aedeagus and the terminal segments of the male abdomen for distinguish- ing closely allied species." I agree entirely with his point of view. Some members of the jelskii group are not easily recognizable without a careful examination of the apex of the aedeagus. For this reason, the genital arma- tures were dissected with fine needles, then mounted in Canadian balsam and drawn with the aid of a camera lucida. I prepared the illustrations of the aedeagus. All other drawings were made by Maria A. Vulcano, of the above-mentioned Departmento de Zoolo- gia, to whom I am very much indebted for her kind cooperation. Amblyopinus tiptoni, new species. Figures 31A, B ; 32. Near A. bolivari (figs. 31E, F; 33B) but readily separated as follows: eyes visible from above ; mentum with two very small marginal bristles on each side ; metasternal process with more than six bristles on the anterior surface ; eighth sternum of male with one or two bristles. Ventral margin of apex of aedeagus (fig. 31A) forming a well sclerotized lobe followed proximally by a wide sinus. Marginal row of spiniform microsetae ex- tending from behind the fourth to the first submarginal apical bristles of the aedeagus. DESCRIPTION, MALE: Head. — Clypeal margin straight, with one very short, blunt seta on each frontolateral angle. Labrum bilobed; each lobe with two very long thin sub- marginal setae on the internal side. Lateral margins divergent, convergent behind the eyes. Lateral lobes separated from the posterolateral or occipital lobes by a relatively wide sinus. Seven eye facets visible from above, just in front of the supraocular bristle. Occipital region with two very well-developed lateral setae, these longer than the post- antennal bristles. Mentum with two very small marginal bristles on each side. Thorax. — Prosternum with two large median bristles. Mesosternum with about 25 bristles arranged as in fig. 31B. Characteristically six small bristles of the 25 are arranged in two apical rows. In A. isabelae n. sp. there are three large apical bristles (fig. 31D). Metasternum as in A. isabelae n. sp. Abdomen and genitalia. — Caudal margin of terga I to VI with a long, pigmented lateral bristle. Margin of tergum VII almost rounded. Tergum VIII relatively slender, with a fringe of thin long setae along the caudal margin. Margin of sternum VI with three conspicuously long, pigmented bristles on each side. Sternum VII bilobed, with two large bristles on the left lobe and four on the right. Apex of sternum VIII entire, rounded, with many slender marginal setae (fig. 32A, VIII). Urostyli with many thin setae on the anterior half of the external margin, and with the large bristles arranged as in fig. 32A. Dorsal apodeme of aedeagus relatively wide, with numerous spiniform microsetae well-developed from near the apex, behind fourth submarginal to the first submarginal bristle. A wide ventral sinus forms a conspicuous isthmus between apex and the anterior part of the aedeagal apodeme. Apical sclerite of internal tube of aedeagus long, slender, dorsally convex, the caudal apex flat, constricted, the proximal end rounded (fig. 31A). BARKER A: Amblyopinus 283 FEMALE: Abdomen and genitalia. — Caudal margin of sternum VI with two large pig- mented bristles on the right side, three on the left. Same condition is found in sternum VII, the bristles arranged as in fig. 32B, VII. Apex of urostyli with four long bris- tles. Two very long bristles on the apex of styliform appendages of segment VIII ar- ranged as in fig. 32B. TYPE MATERIAL : Holotype male from Peromyscus nudipes nudipes (host no. 5967), Bambito (Chiriqui), 5800 feet elevation, collected 9 February 1960 by V. J. Tipton. In the collection of Chicago Natural History Museum. Fig. 31. Apex of aedeagus (A,C,E) and mesosternal plate (B,D,F) of male. A,B, Amblyopinus tiptoni n. sp. C,D, Amblyopinus isabelae n. sp. E,F, Amblyopinus bolivari Barrera, Machado, and Muniz. Allotype female, same host and repository as the holotype, from Lava Flow (Chiriqui), 5000 feet elevation, collected 5 February 1960 by V. J. Tipton. The following paratypes, collected by V. J. Tipton and C. M. Keenan, are all from Peromyscus n. nudipes, with the one exception indicated. A male from Finca Martinz (Chiriqui) , 6800 feet elevation, 13 February 1960 (host no. 6004). In the collection of the Escuela de Biologia of the Universidad Central de Venezuela, Caracas, Venezuela. 284 ECTOPARASITES OF PANAMA Several males and females from Lava Flow and Finca Martinz, 7 Jan- uary 1961 (host nos. 6813, 6815, 6820, 6827, 6836). In the collection of the Laboratorio de Entomologia Economica y Experimental, Escuela Nacional de Ciencias Biologicas, I.P.N., Mexico. Also deposited here is the unique material collected from Reithrodontomys creper (host no. 6309), Cerro Baru, 10500 feet elevation, 2 May 1960. A male and female (host nos. 6844, 6875) both from Lava Flow, 5000 feet elevation, 8 and 9 January 1961, respectively. In the collection of the Departmento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil. From Lava Flow a long series, 27-29 January, 7-9 February, 1960 (host nos. 5794, 5806, 5807, 5935, 5936, 5965) and several males and females, 27 January 1960 and 5 February 1960 (host nos. 5781 and 5906). In the collection of V. J. Tipton. A long series from: Finca Martinz, 30 January 1960 (host no. 5827), 1 February 1960 (host no. 5847), 14 February 1960 (host nos. 6012 and 6022) ; and Casa Lewis (Chiriqui), 5700 feet elevation, 31 January 1960 (host no. 5838). In the collection of Gorgas Memorial Laboratory. This species is named for Lt. Col. Vernon J. Tipton, who has contributed much to the study of ectoparasites of the mammals of Panama. It is important to note that the modified abdominal segments of many paratypes of A. tiptoni n. sp. have been separately mounted. These speci- mens were dissected for examination of gastro-intestinal contents. Dis- section was always carefully made to avoid damage of important sclerotized parts used for descriptive purposes. Amblyopinus emarginatus Seevers Amblyopinus emarginatus Seevers, 1955, Fieldiana, Zoology, 37: 239, figs. 37m, 38f. I have examined three specimens of this species from Rancho Mojica (Bocas del Toro), elevation 5000 feet, from Peromyscus flavidus (host no. 8042), 10 September 1961, and from Oryzomys albigularis (host no. 8098), 12 September 1961, collected by V. J. Tipton and C. M. Keenan. The type specimens are from Huila, Colombia, from Thomasomys laniger. The spe- cies probably also occurs in Venezuela. Amblyopinus isabelae, new species. Figures 31C, D ; 33A. Near A. bolivari (figs. 31E, F; 33B) but readily separated as follows: eyes visible from above; mentum without marginal bristles; metasternal process with more than six bristles on the anterior surface ; eighth sternum of male with two or three bristles. Ventral margin of apex of aedeagus (fig. 31C) with a sclerotized flat extension between fourth and second submarginal apical bristles ; marginal row of spiniform microsetae well developed only from behind the fourth submarginal bristle toward the apex. DESCRIPTION, MALE: Head. — Clypeal margin nearly straight, with one very short and blunt seta on each frontolateral angle. Labrum bilobed; each lobe with three setae on the external side and three on the internal side of the margin. Lateral margins very divergent, but convergent behind the eyes, forming conspicuous lateral lobes separated from the posterolateral or occipital lobes by a short sinus. Five or six eye facets are visible from above just in front of the supraocular bristle. Occipital region with two BARRERA: Amblyopinus 285 B Fig. 32. Amblyopinus tiptoni n. sp., modified abdominal segments. A, eighth sternum (VIII), male. B, seventh sternum (VII), female. 286 ECTOPARASITES OF PANAMA B Fig. 33. Modified abdominal segments, male. A, Amblyopinus isabelae n. sp. B, Am- blyopinus bolivari Barrera, Machado and Muniz. BARRERA: Amblyopinus 287 abortive lateral setae, very much shorter than the postantennal ones. Mentum without marginal bristles. Thorax. — Prosternum with two large median bristles. Mesosternum (fig. 31 D) with about 14 large bristles arranged in two irregular longitudinal rows. Metasternum clothed with several relatively small bristles arranged in eight irregular rows; apex of metasternal process emarginate, nearly pentagonal. Abdomen and genitalia. — Caudal margin of terga I to VI with a long pigmented lateral bristle. Margin of tergum VII evenly rounded. Tergum VIII trapezoidal, with several long, slender marginal bristles; near the posterolateral angles one of these bris- tles is longer and more pigmented than the others. Margin of sternum V with one long pigmented bristle on each side. Margin of sternum VI with two long pigmented bristles each side. Sternum VII bilobed, with two large submarginal bristles on the left lobe and three on the right. Sternum VIII rhomboidal, but with the apex bilobed (fig. 33A). Urostyli with very few setae on the anterior half of the external margin and with the large bristles arranged as in fig. 33 A. Dorsal apodeme of the aedeagus (fig. 31 C) relatively slender, with spiniform microsetae well developed only at apex, behind fourth submarginal bristle. One spiniform microseta under each submarginal bristle. A flat extension runs between the second and fourth submarginal bristles. Lateral lobes wide, rounded. Apical sclerite of internal tube of aedeagus dorsally convex, smooth, with both proximal and distal ends rounded. Female unknown. TYPE MATERIAL: Holotype a male from Omiltemi, State of Guerrero, Mexico ; collected 29 December 1954 by A. Barrera and D. Pelaez. In the collection of the Laboratorio de Entomologia Economica y Experimental, Escuela Nacional de Ciencias Biologicas, I.P.N., Mexico, D.F. REMARKS: This species is named for my wife, the entomologist Isabel Bassols de Barrera, who has often helped me in collecting rodents and ectoparasites. Other ectoparasites collected at Omiltemi were described by Barrera (1958) in a previous paper. There is considerable agreement that the Amblyopinini are obligate ectoparasites. However, neither the literature nor field notes contain con- clusive reports on their feeding habits. Although Zikan (1939) concluded that Amblyopinus henseli Kolbe is a blood-sucking parasite, he offered no positive evidence. By means of the benzidine test for occult blood, I obtained a strong positive reaction for blood in some specimens of the large series of Amblyo- pinus tiptoni n. sp. I have come to the conclusion that, as far as the species tested is concerned, Amblyopinus do parasitize their hosts primarily for food. Normally, however, they feed on skin exudates or on other products that do not contain haemoglobin, although they may occasionally draw peripheral blood when embedding their mandibles deeply into the skin as they usually do. A forthcoming paper will specifically report the results of the examina- tion of gastro-intestinal contents of this and other Amblyopinus. References BARRERA, A. 1958. Insectos parasites de mamiferos salvajes de Omiltemi, Gro., y descripcion de un nuevo sifonaptero: Pleochaetis soberoni, n. sp. An. Esc. Nac. Cien. Biol., 9, (1-4), pp. 89-96, figs. BARRERA, A., MACHADO-ALLISON, C., and MUNIZ, R. 1960. Un nuevo coleoptero parasite de roedores; Amblyopinus bolivari, n. sp., (Col., Staph.). Cienca (Mexico), 20, (5-6), pp. 127-130, figs. SEEVERS, C. H. 1955. A revision of the tribe Amblyopinini : Staphylinid beetles parasitic on mam- mals. Fieldiana, Zoology, 37: 211-264, figs. ZIKAN, J. F. 1939. Amblyopinus henseli Kolbe, um Coleoptero da familia Staphylinidae que para- sita mammiferos. Revista Ent., 10: 219-226, figs. 288 The Fleas (Siphonaptera) of Panama VERNON J. TIPTON l AND EUSTORGIO MENDEZ 2 Published records of fleas from Panama are fragmentary. Jennings (1910) surveyed the two terminal cities of Colon and Panama City for rats and fleas in an attempt to assess the plague potential resident within these two cities. Dunn (1923) listed the fleas known from the more common animals collected in and near the Canal Zone and admitted that his list was incomplete. He later (1934) listed additional species as a result of a general entomological investigation in Chiriqui Province and Fuller (1943) reported on still other material collected by Dunn. Collections made by Dunn in the vicinity of the Canal Zone and J. H. Batty in Chiriqui Province provided new material for taxonomic papers by Jordan and Roths- child (1922, 1923), Jordan (1926) and Kohls (1942). Fairchild (1943) brought together the existing records which included 12 common species and gave additional host data. Mendez and Altman (1960) described a new species of the genus Kohlsia. Tipton and Mendez (1961) described four new species and listed four species not previously known in Panama. This study is based on material in our possession, for the most part collected by us. It attempts to provide data on host-parasite relationships and supplies keys and illustrations to facilitate the identification of the 37 species and subspecies of fleas known to occur in Panama. Six new species are described and seven species are recorded for the first time from Panama. In addition, the male of Rhynchopsyllus megastigmata Traub and Gam- mons, 1950, and the female of Strepsylla dalmati Traub and Barrera, 1955, are described. We wish to express our appreciation to several people whose coopera- tion has made this study possible. We are especially grateful to Lt. Col. Robert M. Altman who initiated the field work on which much of this study is based. It was intended originally that he be a co-author of the publica- 1 Lieutenant Colonel, Medical Service Corps, United States Army. 2 Gorgas Memorial Laboratory, Panama, Panama. 289 290 ECTOPARASITES OF PANAMA tion resulting from our joint study. However, it has been impractical for us to continue the work as a joint project. Colonel Altman has been very cooperative and kindly allowed us to use material collected during his tour of duty in Panama. Colonel Robert Traub (Ret.), formerly with the United States Army Medical Research and Development Command, Office of the Surgeon General, and Mr. F. G. A. M. Smit, British Museum (Natural His- tory), have compared many of our specimens with type material and have given helpful suggestions. We are grateful to Dr. Phyllis T. Johnson, Gorgas Memorial Laboratory, for her suggestions and permission to use illustrations contained in her outstanding study, Fleas of South America. Major Wallace P. Murdoch and Lt. Col. Gordon Field and their staff of il- lustrators at the 406th Medical General Laboratory in Tokyo prepared many of the figures contained herein and we are grateful for their coopera- tion and assistance. Charles M. Keenan has been most helpful with prep- arations for field trips as well as with collecting of host animals. Dr. Charles 0. Handley has identified our host animals and has been very help- ful in obtaining specimens, especially during a joint venture into the Chiri- qui area during March, 1962. We wish also to acknowledge with gratitude the assistance rendered by field workers and laboratory technicians. Prodi- gious feats of strength and endurance in the field were equaled only by patience and painstaking care in the laboratory. KEY TO THE PANAMANIAN GENERA OF FLEAS8 1. Sword-like ridge of mesocoxa usually present, mesonotum with or without pseudo- setae ; metanotum much broader dorsally than ventrally 2 Sword-like ridge of mesocoxa absent ; metanotum lacking pseudosetae ; metanotum rectangular, not especially broader dorsally than ventrally; never with more than one row of bristles on abdominal terga 3 2. With a combination of the following characters: anterior tentorial arm present, always inserting in front of eye; mesopleural rod not bifurcate dorsally; lacking head, thoracic and abdominal combs ; ventral margin of pronotum never bilobed ; fifth tarsal segment always with four lateral plantar bristles 10 Not with above combination of characters 11 3. Inside of metacoxae with spiniform bristles; sensillium with 14 or more pits on a side ; female with anal stylet 5 Inside of metacoxae without spiniform bristles; sensillium with eight pits on a side ; female without anal stylet 4 4. Anterior apical corner of hind coxa projecting downward as a broad tooth; no tooth at base of hind femur Tunga Jarocki Hind coxa without apical tooth ; hind femur with a large basal tooth Rhynchopsyllus Haller 5. Thorax compressed; measured dorsally, meso- and metathorax together shorter than first abdominal tergum Echidnophaga Olliff Thorax normal; measured dorsally, meso- and metathorax together longer than first abdominal tergum 6 6. With both genal and pronotal combs Ctenocephalides Stiles and Collins With only pronotal comb or no combs 7 7. With pronotal comb Hoplopsyllus Baker Without pronotal comb 8 Partially adapted from Hopkins and Rothschild (1953) and Johnson (1957). TIPTON AND MENDEZ : FLEAS 291 8. Dorsal sulcus above antennae absent or poorly sclerotized; mesopleural rod pres- ent; vestigial genal comb absent Xenopsylla Glinkiewicz Dorsal sulcus above antennae well sclerotized; mesopleural rod absent; vestigial genal comb present 9 9. Frons smoothly rounded, without a tubercle Pulex Linnaeus Frons angulate, with small tubercle Juxtapulex Wagner 10. Prosternosome projecting downward between the coxae; mesocoxa rectangular, margins parallel Rhopalopsyllus Baker Prosternosome not projecting downward between coxae ; mesocoxa not rectangular, obviously broadest basally Polygenis Jordan 11. Head with helmet Plocopsylla Jordan Head without helmet 12 12. Pronotal comb present 13 Pronotal comb absent Wenzella Traub 13. Genal comb present; male with third aedeagal rod arising from aedeagal pouch. . . .14 Genal comb absent; male with third aedeagal rod arising from ninth sternum 18 14. Genal comb of two spines 15 Genal comb of four spines Adoratopsylla Ewing 15. Genal comb of two overlapping spines (on rodents) Strepsylla Traub Genal comb of two large spines which do not overlap (on bats) 16 16. First abdominal tergum with well developed comb 17 First abdominal tergum without comb Sternopsylla Jordan and Rothschild 17. Metanotum and terga I through IV with combs. . .Hormopsylla Jordan and Rothschild Only first abdominal tergum with comb Ptilopsylla Jordan and Rothschild 18. Pronotal comb with 24 or more spines 19 Pronotal comb with fewer than 24 spines 20 19. Fifth segment of all tarsi with third pair of plantar bristles ventrally displaced. . Dasypsyllus Baker Fifth segment of all tarsi with third pair of plantar bristles not ventrally dis- placed Ceratophyllus Curtis 20. Males 21 Females 23 21. Distal arm of ninth sternum with stout, cephalad directed bristle; without apical or subapical spiniform bristles Jellisonia Traub Distal arm of ninth sternum without stout cephalad directed bristle 22 22. Distal arm of ninth sternum with distinct notch, lacking spiniform bristles Pleochaetis Jordan Distal arm of ninth sternum without notch, with spiniform bristles. . . .Kohlsia Traub 23. Female anal stylet with dorsomarginal bristle Pleochaetis Jordan Female anal stylet without dorsomarginal bristle 24 24. Tail of spermatheca almost twice length of head; head of spermatheca subglob- ular; bursa copulatrix lying in a horizontal plane Kohlsia Traub Tail of spermatheca only slightly longer than head; head of spermatheca ovoid; bursa copulatrix lying in a vertical plane Jellisonia Traub Superfamily PULICOIDEA Family Pulicidae Subfamily Pulicinae Genus Ctenocephalides Stiles and Collins Ctenocephalides Stiles and Collins, 1930, U.S. Publ. Hlth. Repts., Wash., 45: 1308. Type-species: Pulex cam's Curtis, 1826. 292 ECTOPARASITES OF PANAMA Ctenocephalides felis felis (Bouche). Plate 47. Pulex felis Bouche, 1835, Nova Acta Leop. — Carol., 17: 505. Ctenocephalides felis felis, Hopkins and Rothschild, 1953, Cat. Rothschild Coll. Fleas, 1: 145, figs. 14, 152, 155, 157, 161, 162; pis. 25E, 27A, B, D. Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 228, pis. 100, 101. MATERIAL EXAMINED : 346 males and 742 females collected from several hosts including Cam's familiaris, Felis cattus, Didelphis marsupialis caucae, Nasua nasua narica, Eira barbara biologiae, Conepatus semistriatus trichurus, Liomys adspersus, Coendou mexicanus laenatus, Peromys- cus nudipes nudipes, Felis onca centralis and Oryzomys caliginosus from various localities throughout Panama. In addition, several hundred spec- imens in alcohol, from July 1956 to March 1962, collected by R. M. Altman, C. M. Keenan and V. J. Tipton. REMARKS : Perhaps the most ubiquitous flea in Panama, C. felis felis, has been taken in all of our collecting localities and from a wide variety of hosts. Ctenocephalides canis (Curtis) Pulex canis Curtis, 1826, Brit. Ent., 3: 114, figs. A-E, 8. Ctenocephalides canis, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pp. 227-228. MATERIAL EXAMINED : None. REMARKS: We have not collected this species in Panama. However, Dunn (1923) reports a collection taken from a dog shortly after it had been brought to Panama from an island off the coast of Honduras. Genus Echidnophaga Olliff Echidnophaga Olliff, 1886, Proc. Linn. Soc. N. S. Wales, (2), 1: 171. Type-species : Echidnophaga ambulans Olliff, 1886. Echidnophaga gallinacea (Westwood). Plates 48 (fig. 2), 49 (figs. 3, 4). Sarcopsylla gallinaceus Westwood, 1875, Ent. Month. Mag., 11: 246. Echidnophaga gallinacea, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 232, pis. 108, 109. MATERIAL EXAMINED : None. REMARKS : We have not collected this species although it is likely quite common throughout Panama. Dunn (1923) made several collections from such hosts as poultry, dogs, cats, rats, and man. Genus Pulex Linnaeus Pulex Linnaeus, 1758, Syst. Nat., 10th ed., 1: 614. Type-species: Pulex irritans Linnaeus, 1758. Pulex irritans Linnaeus Pulex irritans Linnaeus, 1758, Syst. Nat., 10th ed., p. 614. Hopkins and Rothschild, 1953, Cat. Rothschild Coll. Fleas, 1: 105, figs. 3-6, ISA, 20B, 75, 124, 125; pis. 1, 4A, 21B, 22A-G. Smit, 1958, Jour. Parasit., 44, (5), pp. 523-526, fig. 2. MATERIAL EXAMINED : 2 females from Conepatus semistriatus trichurus, Cerro Punta (Chiriqui), 12 March 1962, collected by C. O. Handley and V. J. Tipton. TIPTON AND MENDEZ : FLEAS 293 REMARKS : Pulex irritans has been collected in Panama on a number of occasions but Hopla and Smit (personal communications, 1960) have in- dicated that these records are in error. Smit has kindly examined the Panamanian specimens labeled P. irritans in the British Museum. Accord- ing- to him, only one lot of 4 males and 4 females from Ancon are P. simu- lans Baker. An additional lot of 3 females from Bug-aba is in alcohol and could be of either species. We have examined all specimens labeled as irritans in the United States National Museum and are of the opinion that they fit Smit's (loc. cit.) figures of simulans. There is one lot of 1 male and 4 females, which are regarded as irritans, but the origin of these specimens is doubtful, because they were removed from an airplane em- ployed on the Balboa-New Orleans route. We were unable to collect P. irritans in the Chiriqui area where Cham- pion (Hopkins and Rothschild, 1953) found it at elevations from 4000 to 6000 feet, nor has it been found during- several years of collecting in the Canal Zone. However, we have 2 females from skunk (Conepatus semi- striatus) collected in Cerro Punta (Chiriqui) which are probably irritans. Pulex simulans Baker. Plates 48 (fig. 4), 51 (figs. 6, 7). Pulex simulans Baker, 1895, Can. Ent., 27: 65, 67. Smit, 1958, Jour. Parasit., 44, (5), pp. 523-526, fig. 1. MATERIAL EXAMINED : 31 males and 53 females collected by Wilbur Lowe from man in Colon (Panama), March, September and November, 1958. Also 1 male and 4 females from Rattus norvegicus, Panama City, May 1957, collected by E. Mendez. REMARKS: Dunn (1923) reported a heavy infestation of P. irritans in a labor camp (Camp Maru) near the Chagres River on the Atlantic side of the Isthmus. Fuller (1942) lists one male collected from a dog in Pan- ama City and Fairchild (1943) adds data on two lots from man at Moja Polio, Chagres River area. We do not know whether these records refer to P. irritans or to the closely related species, P. simulans. We are not yet convinced that simulans is a distinct species but for the present we choose to retain it as valid, in view of the paucity of our material from hosts other than man. Assuming that the two species are morphologically distinguishable, we suggest that simulans occurs in the lowlands of Pan- ama and that irritans is restricted to elevations of over 5000 feet. Smit (loc. cit.) indicates that irritans has a predilection for man and large carnivores while simulans occurs on rodents which live in colonies. Our records do not bear this out since most of our specimens (31 males and 53 females) of simulans were collected from man and the remaining speci- mens (1 male and 4 females) were collected from Rattus norvegicus. Genus Juxtapulex Wagner Juxtapulex Wagner, 1933, Mitt. Berlin Zool. Mus., 18: 431. Type-species: Juxtapulex echidnophagoid.es Wagner, 1933. 294 ECTOPARASITES OF PANAMA Juxtapulex echidnophagoides Wagner. Plates 48 (fig. 3), 49 (figs. 5, 6). Juxtapulex echidnophagoides Wagner, 1933, Mitteil. Mus. Berlin, 18: 343, figs. 1A, 2-6. Hopkins and Rothschild, 1953, Cat. Rothschild Coll. Fleas, 1: 100-102, fig. 120. Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 255-256. MATERIAL EXAMINED : 273 males and 326 females from Dasypus novem- cinctus fenestratus; 78 males and 138 females from Didelphis marsupialis caucae; 19 males and 19 females from Felis onca concolor; 1 male and 1 female from Canis familiaris ; 1 male from Sciurus granatensis chiriquensis ; 1 female from man. All, except F. onca concolor from El Volcan, were collected at Cerro Punta (Chiriqui) during July 1959, January and Feb- ruary 1960, January, April and May 1961, and March 1962, by C. 0. Hand- ley, C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS : Of 859 fleas of this species collected from 23 hosts, 599 (about 70 percent) were collected from 7 (34 percent) of the hosts, Dasypus novem- cinctus fenestratus, at a rate of 85 fleas per host. Twelve specimens of Didelphis marsupialis caucae were collected at elevations above 5000 feet. From 11 of these, 216 fleas were removed at a rate of approximately 20 fleas per host. Even though J. echidnophagoides may live quite successfully in association with marsupials, it is likely that edentates are true hosts. Genus Hoplopsyllus Baker Hoplopsyllus Baker, 1905, Proc. U. S. Nat. Mus., 29: 130. Type-species: Pulex anomalus Baker, 1904. Hoplopsyllus glacialis exoticus Jordan and Rothschild. Plates 50, 51 (figs. 1-5). Hoplopsyllus exoticus Jordan and Rothschild, 1923a, Ectoparasites, 1: 311-312, fig. 314. Hopkins and Rothschild, 1953, Cat. Rothschild Coll. Fleas, 1 : 192-193, fig. 181. Hoplopsyllus glacialis exoticus, Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 256-259, pis. 1,2. MATERIAL EXAMINED : 24 males and 47 females from Sylvilagus brasilien- sis gabbi, 5 males and 3 females from Didelphis marsupialis caucae, 1 male from Nasua nasua narica; Cerro Punta (Chiriqui), January 1960, January 1961 and March 1962, collected by C. O. Handley, C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS: H. glacialis exoticus is found most commonly, although not exclusively, on rabbits. Apparently it does not occur on rabbits in the lowlands. Genus Xenopsylla Glinkiewicz Xenopsylla Glinkiewicz, 1907, Sitzber. Akad. Wiss. Wien, Abt. 1, 116: 385. Type-species : Xenopsylla pachyuromyidis Glinkiewicz, 1907. Xenopsylla cheopis Rothschild. Plates 52, 53. Pulex cheopis Rothschild, 1903, Ent. Month. Mag., (2), 14: 85, pi. 1 (figs. 3, 9), 2 (figs. 12,19). TIPTON AND MENDEZ : FLEAS 295 Xenopsylla cheopis Hopkins and Rothschild, 1953, Cat. Rothschild Coll. Fleas 1: 248-260, figs. 20A, 76, 199, 220, 246, 255, 259, 266, 286, 305-308, 310, 391; pis. 2, 22D-F, 39A, 40E. Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pp. 226-227, pis. 98, 99. MATERIAL EXAMINED: 3 males and 5 females Rattus rattus; Arraijan and Santa Fe; February and April 1959, collected by R. M. Altman and C. M. Keenan. REMARKS: The distal arm of the ninth sternum is considerably more setose in our specimens than in those of Hopkins and Rothschild (1953). The shape of the spermatheca also differs but within the range of intra- specific variation. Jennings (1910) collected 3022 fleas from 2394 rats (presumably of the genus Rattus) , an average of 1.26 fleas per rat. Most of the fleas (2966 or 97.9 percent) were X. cheopis. In a later survey in Colon and Panama City in 1920-21, he collected 505 fleas from 1225 rats and mice. However, 1020 of the hosts were mice, from which only 29 fleas were removed. There were 476 fleas on 205 rats or an average of 2.33 fleas per rat. The rats captured on the drier Pacific side of the Isthmus were more heavily in- fested than those from the Atlantic side. In recent years there have been very few specimens of X. cheopis col- lected in the Canal Zone. Only eight specimens of X. cheopis were collected from 119 rats in Arraijan just outside of the Canal Zone. The widespread use of insecticides for the control of mosquitoes may be partially responsible for the paucity of fleas on domestic rats. Subfamily Tunginae Genus Tunga Jarocki Tunga Jarocki, 1838, Zoologiia, 6: 50. Type-species: Pulex penetrans Linnaeus, 1758. Tunga penetrans (Linnaeus). Plates 48 (fig. 1), 49 (figs. 1, 2). Pulex penetrans Linnaeus, 1758, Syst. Nat., 10th ed., p. 614. Tunga penetrans, Hopkins and Rothschild, 1953, Cat. Rothschild Coll. Fleas, 1: 39-43, figs. 21, 22A, 23, 26A, 28, 37; pis. 6A, B, 7A-C, 8B. MATERIAL EXAMINED: 31 males and 9 females from Sus scrofa; Santa Fe (Veraguas) and Las Palmitas (Los Santos) ; January 1956, February 1957 and January 1962, collected by R. M. Altman, C. 0. Handley and C. M. Keenan. REMARKS : This species does not appear to be highly variable. Hopkins and Rothschild (loc. cit.) compare the claspers of males from Brazil and Nyasaland and as expected, our specimens agree more closely with those from Brazil. T. penetrans is widespread throughout Panama according to those who have visited villages of the interior. Residents of these vil- lages are familiar with the "nigua," and in some areas of western Panama, it is a common pest of man and domestic pigs. 296 ECTOPARASITES OF PANAMA Genus Rhynchopsyllus Haller Rhynchopsyllus Haller, 1880, Arch. Naturg., 46, Bd. 1, p. 72. Type-species: Rhynchopsyllus pulex Haller, 1880. Rhynchopsyllus megastigmata Traub and Gammons. Plate 54. Rhynchopsyllus megastigmata Traub and Gammons, 1950, Jour. Parasit., 36, (3), pp. 271-272, figs. 6, 7. MATERIAL EXAMINED : 4 males and 3 females from bat guano under tile on church at Pacora, 20 June 1961 and 12 January 1961 ; 1 female from Tadarida yucatanica, same locality, 20 June 1961 ; collected by C. M. Keenan and V. J. Tipton. REMARKS : We are not certain that all of our specimens should be called R. megastigmata. Traub (personal communication, 1962) has pointed out that if the sexes are conspecific, there is extreme sexual dimorphism in this species. However, it seems unlikely that males of one species would be found occupying the same ecological niche as females of a closely re- lated species. On the other hand, our series is small and it is possible that we have collected four male specimens of one species and three female specimens of another species in the same guano deposit. Perhaps at a later date, when male specimens of R. pulex have been collected, it will be possible to determine the extent of sexual dimorphism in this genus. Our female specimens differ from the description and illustrations given by Traub and Gammons (loc. cit.) in several details. In our specimens, the spermatheca is extremely large, the abdominal spiracles are not quite as large, the lateral process of the prosternum and the maxillary lobe are longer and somewhat more acuminate, and the postoral process is broader basally. There also appear to be some differences in the arrangement of the bristles on the eighth tergum. The type material, consisting of two females, and our four specimens make a total of six female specimens available for study. Such a short series does not allow a proper evaluation of intraspecific variation. Despite the possibility that our specimens may represent two distinct species, we choose to refer them to R. megastigmata. DESCRIPTION, MALE: Head (pi. 54, fig. 1). — Dorsal margin of head shallow arc; ante- rior margin well rounded ; ventral margin almost straight, terminating in small postoral process. One large preantennal bristle cephalad of eye; 10-11 minute spiniforms scat- tered over preantennal area. Eye well developed, subovate; with small ventral sinus. Genal process long, subacuminate. Maxillary lobe short, reaches to apex of first segment of maxillary palpus; broad basally, suddenly reduced to subacuminate apex. Stilettos broad, serrations not prominent; reach slightly beyond apex of maxillary palpus. Labial palpus three-segmented; reach almost to apex of forecoxa. Antenna oblong; bristles of second segment reach beyond middle of club. Postantennal area with four long bristles plus five or six spiniforms. Thorax (pi. 54, fig. 2). — Pronotum with single row of four bristles. Prosternum broad, without lateral process. Mesonotum (MSN.) with row of five bristles. Meso- sternum (MST.) undivided; with one long bristle. Metanotum (MTN.) with two or three bristles. Lateral metanotal area (L.M.) well developed. Metepisternum (MTS.) with long dorsal bristle immediately ventrad of prominent, strongly sclerotized pleural arch (PL.A.). Metasternum (MTT.) fairly well developed; saber-like. Metepimere (MTM.) large; with five bristles. TIPTON AND MENDEZ : FLEAS 297 Legs. — Metafemur with basal spur. Tibia and first three tarsal segments much broader apically than basally. Third tarsal segment with two bristles which reach be- yond apex of fifth segment. Modified Abdominal Segments (pi. 54, figs. 3-5). — Eighth tergum (8T.) reduced. Tergum 9 with broad, cephalad-directed apodeme (T. AP. 9) ; rod-like dorsal apodeme. Manubrium much reduced, thumb-like; directed caudad. Immovable process of clasper (P.) large; with several small bristles scattered over surface. Movable process of clasper divided (Fi, F2) ; apical half of Fi broad, somewhat truncate; basal half with deep lateral sinus to accommodate ventral spur on F2; caudal margin of F2 strongly arched; anterior margin with truncate recess to receive apical portion of Fi. Distal arm of ninth sternum (D.A.9) roughly foot-like; with three small bristles on ventral margin, about 18 lateral bristles. Aedeagus (pi. 54, fig. 6). — Aedeagal apodeme short, broad; with prominent middle plate (MI. P.) strongly hooked apically. Proximal spur absent. Crescent sclerite (C.S.) inconspicuous. Medial dorsal lobe (M.D.L.) rounded subapically; apex hook-like. Sclero- tized inner tube (S.I.T.) long, straight; with prominent lateral process. Lateral lobe of adeagus (L.L.) broadly nipple-like. Crochet (CR.) broad basally; with short apical hook. Pseudocrochet (PS.C.) on broad stalk projecting from wall of aedeagal pouch; apex slightly rugose. Superfamily RHOPALOPSYLLOIDEA Family Rhopalopsyllidae Subfamily Rhopalopsyllinae Genus Polygenis Jordan Polygenis Jordan, 1939, Novit. Zool. 41 : 444. Type-species: Pulex roberti Rothschild, 1905. Polygenis atopus (Jordan and Rothschild) . Plate 55. Rhopalopsyllus atopus Jordan and Rothschild, 1922, Ectoparasites, 1: 267, figs. 259, 260. Polygenis atopus, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pp. 158-159. MATERIAL EXAMINED : 2 males and 3 females from Peromyscus nudipes nudipes and 1 male from Oryzomys albigularis, Rancho Mojica (Bocas del Toro), September 1961, collected by V. J. Tipton. REMARKS : In a comparison with the original description and figures we note that in our specimens the distal arm of the ninth sternum of the male is more setose and the apex not quite so truncate. The sinus in the process of the clasper is almost absent and PI is very short in our specimens. As mentioned elsewhere, the flea fauna of Peromyscus nudipes nudipes captured on the Pacific slope was quite different from that of the Atlantic slope although the two localities were separated by only a few miles and at essentially the same elevation. P. atopus was not found on any of the more than 200 specimens of P. nudipes nudipes collected on the pacific slope but was found only on the Atlantic slope above 5000 feet elevation. The oc- currence of atopus at this locality lends credence to the statement by Fair- child (personal communication, 1961) that it is not uncommon to find South American species of insects in Atlantic drainage localities on the Isthmus and that species of the Pacific drainage are characteristically North American. 298 ECTOPARASITES OF PANAMA Polygenis dunni (Jordan and Rothschild) . Plates 56 (fig. 1) , 57 (figs. 1, 2) . Rhopalopsyllus dunni Jordan and Rothschild, 1922, Ectoparasites, 1, (4), p. 269, figs. 261, 262. Jordan and Rothschild, 1923b, Ectoparasites, 1, (5), pp. 336, 351. Rhopalopsyllus (Polygenis) dunni, Ewing and Fox, 1943, Fleas N. Am., p. 22. Polygenis dunni, Costa Lima and Hathaway, 1946, Pulgas, p. 144. Polygenis ambersoni Traub and Johnson, 1952, Bol. Of. Sanit. Panam., 32: 112, figs. 1-4, 6, 9, 10, 12-14, 16. Polygenis dunni, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pp. 160-161 (synony- mizes ambersoni). MATERIAL EXAMINED : 19 males and 19 females from Liomys adspersus, 2 males and 1 female from Proechimys semispinosus panamensis, 1 male and 1 female from Zygodontomys microtinus, 1 male from Sigmodon his- pidus chiriquensis, 2 females from Metachirus nudicaudatus dentaneus, 1 female from Oryzomys capito, 1 female from Sciurus granatensis chiriquen- sis; Canal Zone, Cerro Azul, Rio Seteganti, Chiriqui Lagoon; September- December 1956, July 1958, February 1960, January and March 1961, col- lected by R. M. Altman, C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS: Our specimens conform closely to the original description and show very little variation. However, we have a rather short series (23 males and 25 females). Polygenis klagesi (Rothschild) . Plates 56 (fig. 3) , 57 (figs. 5, 5a, 6, 6a, 6b) . Pulex klagesi Rothschild, 1904, Novit. Zool., 11: 620, pis. 9 (fig. 28).. 10 (figs. 35, 39). Rhopalopsyllus klagesi samuelis Jordan and Rothschild, 1923b, Ectoparasites, 1: 331, figs. 342, 345. Polygenis klagesi samuelis Johnson, 1957, Mem. Ent. Soc. Wash., no. 5 : 164. Rhopalopsyllus klagesi klagesi Jordan and Rothschild, loc. cit., p. 332, figs. 343, 344. Polygenis klagesi klagesi Johnson, loc. cit., p. 163. MATERIAL EXAMINED: 966 males and 1347 females from Proechimys semispinosus panamensis, 30 males and 51 females from Hoplomys gym- nurus, 13 males and 18 females from Tylomys panamensis, 19 males and 24 females from Nasua nasua narica, 3 males and 8 females from Zygo- dontomys microtinus, 3 males and 7 females from Didelphis marsupialis caucae, 2 males and 2 females from Oryzomys capito, 2 females from 0. bombycinus, 1 male and 3 females from Dasyprocta punctata isthmica, 2 males and 1 female from Marmosa robinsoni, 2 females from Liomys adsper- sus, 1 female from Heteromys australis, 1 female from Sigmodon hispidus chiriquensis and several hundred specimens in alcohol ; from all collecting localities at 2500 feet elevation and below. In addition 8 males and 25 females from Coendou mexicanus laenatus, Cerro Punta (Chiriqui), col- lected by R. M. Altman, C. 0. Handley, C. M. Keenan, V. J. Tipton, and Gorgas Memorial Laboratory. REMARKS: Jordan and Rothschild (1923) referred Panamanian spec- imens of P. klagesi (Rothschild, 1904) to a new subspecies, P. klagesi samuelis, based on the shape of the spermatheca and the apical width of the movable process of the clasper compared with the immovable process of the clasper. It is our opinion that the figures and description given by Jordan and Rothschild are not adequate for an accurate assignment of our TIPTON AND MENDEZ : FLEAS 299 specimens to either subspecies. Furthermore, there is an unusual amount of variation in this species as expressed in the length of the distal arm of the ninth sternum. In specimens from the Canal Zone, the length of D.A.9 is 94 p.; from Cerro Azul, 140 ^; from Rio Changena, 156 /*; and from Cerro Punta, 187 p. The spermatheca of our specimens resembles the figure by Jordan and Rothschild for P. klagesi klagesi, while the male claspers more closely resemble their figures of P. klagesi samuelis. The width and length of the movable process of the clasper and the shape of the spermatheca are highly variable. Specimens from Coendou mexicanus laenatus (pi. 57, figs. 5a, 6a, 6b) collected in Chiriqui represent a distinct population and may be even a distinct species. Smit (personal communication, 1962) pointed out that the abdominal spiracles are larger and the seta below the eye is posi- tioned differently than in P. klagesi samuelis. The shape of the sperma- theca, the setal pattern of the clasper and the length of the distal arm dif- fer from lowland specimens. It will be necessary to obtain specimens from the entire range of P. klagesi and a detailed study of the aedeagus will be required to determine the status of subspecific populations of this species. Almost 99 percent of the fleas collected from Proechimys semispinosus panamensis were Polygenis klagesi and 51 percent of the specimens of P. s. panamensis harbored this flea. Over 92 percent of all the specimens of klagesi collected were from this host and 96 percent from hystricomorph rodents. Records maintained over a period of four years indicate there is a sea- sonal fluctuation in the population of P. klagesi. It appears that the numbers increase at the onset of the wet season but drop off sharply as the rainfall increases. As will be noted in the table below, the dry season corresponds roughly with the first quarter of the calendar year. Rainfall is expressed as an average of both the Atlantic and Pacific sides of the Isthmus. Average Rainfall Fleas per Rat Quarter per Month in Inches (Positive Rats Only) 1 1.7 2.8 2 7.3 14.4 3 10.7 4.0 4 11.7 1.7 Polygenis roberti beebei (I. Fox). Plates 56 (fig. 2), 57 (figs. 3, 4). Rhopalopsyllus beebei I. Fox, 1947, Zoologica, 32: 117, fig. 2. Polygenis roberti beebei, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pp. 168-169. MATERIAL EXAMINED: 13 males and 17 females from Oryzomys capita, 14 males and 29 females from O. caliginosus chrysomelas, 5 males and 5 females from O. bombycinus, 3 males and 7 females from Didelphis marsu- pialis caucae, 2 males and 4 females from Nectomys alfari, 2 males and 5 females from Proechimys semispinosus panamensis, 1 male and 2 females from Philander opossum fuscogriseus, 1 male and 2 females from Zygo- dontomys microtinus, 1 male from Metachirus nudicaudatus dentaneus, 3 females from Marmosa robinsoni, 1 female from Peromyscus nudipes nudi- pes, 1 female from Heteromys desmarestianus ; all collecting localities be- 300 ECTOPARASITES OF PANAMA tween 2000 and 3000 feet elevation plus a few specimens from the Canal Zone; July-August 1956, May-June 1957, January-February 1958 and September 1961, collected by R. M. Altman, C. M. Keenan, V. J. Tipton, C. E. Yunker, and Gorgas Memorial Laboratory. REMARKS: Most of our specimens were collected at 2550 to 3000 feet elevation. Although they are associated with a wide range of hosts they appear to have a predilection for oryzomine rodents. KEY TO THE PANAMANIAN SPECIES OF POLYGENIS JORDAN 1. Immovable process of clasper (P.) divided by shallow sinus; spermatheca without sharp distinction between head and tail P. atopus (Jordan and Rothschild) Immovable process of clasper (P.) not divided by sinus; spermatheca with sharp distinction between head and tail 2 2. Apex of movable process of clasper (F.) reaches to or beyond antero-dorsal mar- gin of immovable process (P.) ; head of spermatheca roughly ovoid, tail narrow and not recurved back over head P. klagesi (Rothschild) Apex of movable process of clasper (F.) does not reach to antero-dorsal margin of immovable process (P.) ; head of spermatheca roughly triangular, tail broad and recurved back over head 3 3. Bristles of distal arm of ninth sternum short, approximately as long as width of distal arm of ninth sternum. Labial palpus does not reach trochanter I P. roberti beebei (I. Fox) Bristles of distal arm of ninth sternum long, approximately four times as long as width of distal arm ; labial palpus reaches to trochanter I P. dunni (Jordan and Rothschild) Genus Rhopalopsyllus Baker Rhopalopsyllus Baker, 1905, Proc. U. S. Nat. Mus., 29: 128, 129, 143. Type-species: Pulex lutzi Baker, 1904. Rhopalopsyllus australis tupinus Jordan and Rothschild. Plates 58 (fig. 1),59 (figs. 1,2). Pulex australis Rothschild, 1904, Novit. Zool., 11: 613, pis. 9 (fig. 29), 10 (figs. 34, 36), partim. Rhopalopsyllus australis tupinus Jordan and Rothschild, 1923b, Ectoparasites, 1: 328, figs. 339. Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 176. MATERIAL EXAMINED: 96 males and 130 females from Dasyprocta punc- tata, 24 males and 39 females from Nasua nasua narica, 2 males and 5 females from Didelphis marsupialis caucae, 3 males and 3 females from Chironectes minimus, 2 males and 2 females from Tamandua tetradactyla, 3 males and 11 females from Proechimys semispinosus panamensis, 8 males and 11 females from Agouti paca virgatus, 1 male and 1 female from Zygo- dontomys microtinus, 1 male from Sciurus granatensis chiriquensis, 1 fe- male from Philander opossum fuscogriseus, 1 female from Eira barbara biologiae, 1 female from Galictis allamandi, 1 female from Canis familiaris, 1 female from man, 10 males and 20 females from Tayassu tajacu; mostly from Canal Zone but also from Chiriqui, Bocas del Toro, Los Santos, and Darien Provinces; November 1956 to March 1962, collected by R. M. Alt- man, C. O. Handley, C. M. Keenan, V. J. Tipton, C. E. Yunker, and Gorgas Memorial Laboratory. TIPTON AND MENDEZ : FLEAS 301 REMARKS : Fuller (1942) lists R. australis australis from D. marsupialis etensis, Alhajuela; R. australis tupinus from Tayassu tajacu, Alhajuela; and R. australis tamoyus taken from D. marsupialis etensis, Ancon and Chilibrillo Caves. The morphological differences in these subspecies are based on the position of a notch of the movable process and a groove in the immovable clasper in relation to the acetabulum. With few exceptions our specimens fit the figures given by Jordan and Rothschild (1923) for R. australis tupinus. However, there is an unusual amount of variation in our series. For example, a male from Dasyprocta punctata isthmica is easily referrable to R. australis tupinus while another male from the same host specimen more closely approaches R. a. tamoyus. We prefer to refer all of our specimens to a single subspecies in view of the variation found in a series from a single host specimen. Further, it is very easy to confuse R. lugubris with subspecies of R. australis; this may have happened in the past. The subspecies of R. australis are most frequently associated with agou- tis, pacas and other hystricomorph rodents, and the carnivores which prey upon them. Most of our specimens were collected during the dry season but this may be due to the more favorable collecting conditions. Rhopalopsyllus cacicus saevus Jordan and Rothschild. Plates 58 (fig. 3), 59 (figs. 5, 6). Rhopalopsyllus cacicus saevus Jordan and Rothschild, 1923b, Ectoparasites, 1 : 325, fig. 332. Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pp. 176-177. MATERIAL EXAMINED : 77 males and 104 females from Dasypus novem- cinctus fenestratus, 6 males and 12 females from Didelphis marsupialis caucae, 2 males and 3 females from Proechimys semispinosus panamensis, 5 females from Philander opossum fuscogriseus, 1 male and 1 female from Nasua nasua narica, 1 male from Agouti paca virgatus, 1 female from Metachirus nudicaudatus dentaneus, 2 females from Dasyprocta punctata, 105 males and 159 females from animal burrows (presumably Dasypus novemcinctus fenestratus) ; mostly from the Canal Zone, but a few from other sea level localities in Panama; July 1956 to March 1961, collected by R. M. Altaian, C. M. Keenan and V. J. Tipton. REMARKS: Fuller (1942) lists this species from Dasypus novemcinctus and Didelphis marsupialis from several localities in and near the Canal Zone. The variation due to orientation of specimens on microscope slides is frequently great enough to constitute what appear to be morphological differences. However, all of our specimens fit fairly closely the description and figures for R. cacicus saevus. In many of the animal burrows listed above we found the host present. We have collected leaves and soil from the burrows in which we found Dasypus novemcinctus fenestratus; from this debris we have removed large numbers of R. cacicus saevus. We surmise that D. novemcinctus fene- stratus is the true host of this flea at low elevations while J. echidnopha- goides replaces R. cacicus saevus on the same host at high elevations. 302 ECTOPARASITES OF PANAMA Rhopalopsyllus lugubris cryptoctenes (Enderlein). Plate 59 (fig. 3a). Rhopalopsyllus lugubris Jordan and Rothschild, 1908, Parasitology, 1: 74; pis. 3 (fig. 12), 6, (fig. 9). Rothschildella cryptoctenes Enderlein, 1912, Zool. Anz., 40: 72, figs. 1-5, 7. Rhopalopsyllus lugubris cryptoctenes, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 177, pis. 92 (figs. 1-3, 5), 93 (figs. 1, 2, 5). MATERIAL EXAMINED: 51 males and 64 females from Agouti paca virga- tus and 17 males and 19 females from Dasyprocta punctata isthmica; El Volcan and Cerro Punta (Chiriqui) ; July 1958, June-July 1959, January 1960 and March 1962, collected by C. M. Keenan, V. J. Tipton, and Gorgas Memorial Laboratory. REMARKS : All of our specimens of R. lugubris from paca and agouti in Chiriqui are R. I cryptoctenes. Dunn (1934) reports this species from agoutis collected in the lowlands in Chiriqui Province but they were prob- ably R. I. lugubris. Johnson (loc. cit.) lists a single male "ex human" col- lected by H. C. Mathes in Volcan (Chiriqui) . There is considerable variation in the characters used to separate R. lugubris cryptoctenes from the nominate subspecies. This is especially true of the incrassation which separates the lateral groove on the caudal margin of the process of the clasper from the acetabulum. In general, how- ever, this incrassation is very broad in cryptoctenes, and it extends at least two-thirds of the distance from the acetabulum to the apex of the finger of the clasper. In the nominate subspecies the incrassation extends less than one-half this distance. The notch on the finger of the clasper opposite the lateral groove of the process of the clasper is subapical in cryptoctenes and is roughly at the midpoint in lugubris. There are no demonstrable differences in the females. Rhopalopsyllus lugubris lugubris Jordan and Rothschild. Plates 58 (fig. 2), 59 (figs. 3, 4). Rhopalopsyllus lugubris Jordan and Rothschild, 1908, Parasitology, 1: 74, pis. 3 (fig. 12), 6 (fig. 9). Rhopalopsyllus lugubris lugubris, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 177, pis. 92 (fig. 4), 93 (fig. 4). MATERIAL EXAMINED : 15 males and 16 females from Agouti paca virga- tus, 1 male and 3 females from Dasyprocta punctata isthmica, 7 males and 3 females from Didelphis marsupialis caucae, 1 female from Zygodontomys microtinus, 1 female from Proechimys semispinosus panamensis and 1 male and 2 females from man; Canal Zone, Darien, San Bias, Bocas del Toro Provinces and other sea level collecting localities; October 1957 through July 1958, April-October 1959, March, September 1961 and February 1962, collected by R. M. Altman, C. M. Keenan, V. J. Tipton, and Gorgas Memorial Laboratory. REMARKS: All specimens of R. lugubris collected at elevations below 4000 feet are probably R. I. lugubris. In addition, 1 male and 1 female from man, Rancho Mojica (Bocas del Toro), elevation 5600 feet, have been assigned to this subspecies. None of our specimens of R. lugubris conform TIPTON AND MENDEZ : FLEAS 303 exactly to figures given by Jordan and Rothschild (1923b) or Johnson (loc. cit.) which makes clear the need for a revision of the genus Rhopalop- syllus. Rhopalopsyllus species, near mesus Wagner MATERIAL EXAMINED: 1 female from Sciurus granatensis chiriquensis, Cerro Punta (Chiriqui), January 1960, collected by C. M. Keenan and V. J. Tipton. REMARKS: This single specimen has a rather unique spermatheca and probably represents an undescribed species. Its description, however, will await the collection of additional material. KEY TO THE PANAMANIAN SPECIES OF RHOPALOPSYLLUS BAKER 1. Spiracle of metepimere oblong, prolonged dorsally; head of spermatheca globular. . R. lugubris (Rothschild) Spiracle of metepimere rounded or subrounded ; head of spermatheca not globular .... 2 2. Labial palpus reaching to apex of coxa I or beyond; movable process of clasper longer than distal arm of ninth sternum ; spermatheca not strongly S-shaped .... R. cacicus saevus Jordan and Rothschild Labial palpus not reaching apex of coxa I; movable process of clasper about as long as distal arm of ninth sternum ; spermatheca strongly S-shaped R. australis tupinus Jordan and Rothschild Superfamily CERATOPHYLLOIDEA Family Ischnopsyllidae Genus Hormopsylla Jordan and Rothschild Hormopsylla Jordan and Rothschild, 1921, Ectoparasites, 1, (3), p. 158. Type-species: Ceratopsylla fosteri Rothschild, 1903. Hormopsylla kyriophila, new species. Plates 60, 61. DIAGNOSIS : H. kyriophila is more heavily sclerotized than the other spe- cies of the genus. In chaeotaxy of the frons and dorsal incrassations of the occiput it closely resembles H. fosteri (Rothschild) . It may be sepa- rated from the latter species in that the male eighth sternum is subacum- inate and bears about 16 regularly spaced bristles on the ventral margin to form a comb whereas in fosteri there are only eight to ten bristles and the apex is rounded ; the distal arm of ninth sternum is not globular as in fosteri; the apical portion of the crochet is only slightly sinuate but is S- shaped in fosteri and the immovable process of the clasper is almost twice as wide as the movable process of the clasper while it is only slightly wider in fosteri. In the female the apex of the tail of the spermatheca almost touches the head of the spermatheca resulting in virtually a closed circle while the angle between the head and the tail of the spermatheca in fosteri is 30 to 40 degrees. DESCRIPTION: Head, Male (pi. 60, fig. 1). — Anterior margin evenly rounded; pre- antennal area covered with 62-64 short, spiniform bristles plus four longer bristles; prominent mesal sclerotization running parallel with anterior margin of head; vertical incrassation extending from dorsal region. Two genal teeth; anterior tooth broad, about two-thirds length of narrower tooth; preoral tuber distinct. Genal process long, 304 ECTOPARASITES OF PANAMA extends well beyond apex of antenna; apex rounded. Eye small, lightly pigmented. Postantennal area with five rows of marginal bristles of two bristles each ; four vertical incrassations between rows of bristles, middle two most pronounced; diagonal row of bristles along antennal fossa; pale, mesal, triangular area with four short, three mod- erately long and three long bristles. Thorax (pi. 60, fig. 2). — Pronotum with approximately nine bristles (three short, three moderately long, three long) ; about 24 teeth in pronotal comb which curves cephalad on ventral end. Mesonotum (MSN.) with six long bristles, four or five shorter bristles; two broad thickened areas extending ventrad from dorsal margin. Mesepimere (MPM.) with three long, six shorter bristles. Mesepisternum (MPS.) divided by broad horizontal band; two long, four shorter bristles above band; lower portion devoid of setae. Met- anotum (MTN.) with four dorsal incrassations; seven long, two short bristles; apex with comb of ten teeth. Lateral metanotal area (L.M.) elongate, with single long bristle. Metepisternum (MTS.) with one long bristle; squamulum well developed. Met- epimere (MTM.) with five bristles. Legs. — Procoxa with about 36 bristles plus marginals; profemur with two short, one longer bristle on outer lateral surface. Meso-, metacoxae with heavily, sclerotized in- ternal rods. Meso-, metafemora with single lateral bristle. Hind tibia with eight dorso- caudal notches with bristles arranged 2-2-1-2-2-1-2-2; single bristles between last five notches. Tarsal segment lengths (in /*) 156, 140, 104, 68, 125; four pairs lateral plantar bristles. Abdomen. — Terga 1-4 with well developed combs with teeth numbering 18, 14, 14, 14; each tergum with globular sclerotization extending from dorsal margin ; two bristles per segment with bases near dorsal margin plus two additional bristles more ventrad with spiracle between them. Minute bristle at base of single, long antesensillial bristle. Sterna with darkened, heavily sclerotized areas near ventral margins; one bristle per sternum. Modified Abdominal Segments, Male (plate 61, figures 4, 5, 6, and 7). — Tergum 8 (ST.) with four stout bristles caudad of sensillium; caudal margin rounded. Sternum 8 (SS.) with broad, subacuminate process bearing 15—16 small bristles on ventral margin resembling comb. Distal arm of ninth sternum (D.A. 9) with pronounced lobe on caudoventral margin bearing one short, one long setae; apex shaped like head of bird with prominent cephalad-directed beak; small marginal seta proximad of throat; two longer setae on back of neck. Immovable process of clasper (P.) membranous; outline indistinct; apex reaches to approximately midpoint on movable process of clasper (F.). F. prominent ; with five or six latsral bristles distad of P. ; point of articulation with P. medial. Aedeagus (pi. 60, fig. 3). — Aedeagal apodeme (AE.A.) long, slender; with upturned, acuminate apex; proximal spur (P.S.) prominent; apex angular. Accessory lateral lobe (A.L.L.) with prominent dorsal hump. Crescent sclerite (C.S.) small, rod-like. Median dorsal lobe (M.D.L.) a membranous flap with distal half doubled. Lateral lobes (L.L.) with caudal margin subtruncate. Crochet (CR.) very broad basally; apex slightly sinuous, acuminate. Modified Abdominal Segments, Female (pi. 61, figs. 1-3). — Seventh sternum (IS.) with sloping, almost straight caudal margin; bearing six or seven lateroventral bristles. Eighth tergum (ST.) with two long, lateral bristles plus several marginals. Eighth sternum (SS.) indistinct, broadly rounded caudal margin. Dorsal anal lobe of proctiger with five or six bristles. Anal stylet about twice as long as wide; long apical bristle plus minute subapical bristle. Ventral anal lobe of proctiger with shallow sinus in caudal margin ; with two long submarginal, four or five marginal bristles. Spermatheca swollen slightly on head-end, otherwise little distinction between head and tail; curved so that apex of tail almost touches head. Bursa copulatrix club-shaped; with slightly sinuous handle. TYPE MATERIAL: Holotype male (host no. 7583) from Tadarida yucatan- ica, collected under roof tile on belfry of church at Pacora, about 25 miles northeast of Panama City (Panama), elevation sea level, 21 June 1961, TIPTON AND MENDEZ : FLEAS 305 collected by C. M. Keenan and V. J. Tipton. Allotype female (host no. 7284) , same locality and collectors, 20 June 1961. Paratypes : 2 males and 9 females, same host, locality, and collectors, 6 June 1961 (host no. 7231), 20 June 1961 (host nos. 7291, 7504, 7509), 21 June 1961 (host nos. 7563, 7566, 7589) ; 3 females, same locality and collectors, from Molossus coibensis (host nos. 7297, 7501), 20 June 1961 ; 11 males and 6 females, same locality and collectors, from bat guano (coll. no. 8824) 12 January 1962. Holotype male and allotype female in the collection of the United States National Museum. Paratypes (1 pair each) deposited in the British Museum (Natural History); Chicago Natural History Museum; Gorgas Memorial Laboratory in Panama City; Environmental Health Branch, Corozal, Canal Zone; private collections of Robert Traub and the senior author. Genus Ptilopsylla Jordan and Rothschild Ptilopsylla Jordan and Rothschild, 1921, Ectoparasites, 1, (3), p. 158. Type-species: Ptilopsylla leptina Jordan and Rothschild, 1921. Ptilopsylla dunni Kohls. Plates 62, 63. Figure 34. Ptilopsylla dunni Kohls, 1942, Jour. Parasit., 28, (5) , pp. 361-362, 1 fig. Hopkins and Rothschild, 1956, Cat. Rothschild Coll. Fleas, 2: 207-208, figs. 301, 335, 357, 358, pis. 19A, 23A, 25A. Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, pis. 44-46. MATERIAL EXAMINED: 83 males and 163 females from Tadarida yucatan- ica, 66 males and 129 females from Molossus coibensis and 2667 males and 5305 females from bat guano. Bats were collected in mist nets, or, like bat guano, were collected from under roof tile on the belfry of the church at Pa- cora about 25 miles northeast of Panama City (Panama) , elevation sea level, by C. M. Keenan, C. E. Yunker and V. J. Tipton in June and July of 1961 and January of 1962. REMARKS : P. dunni is represented by a series of several thousand spec- imens. They conform quite well to the figures given by Johnson (1957) except for the crochet. We have a few specimens with crochets like those in Johnson's drawings but the majority of them are oriented so that the crochet resembles an inverted high-heeled boot or a raised thumb and point- ing index finger with a membrane between (pi. 63, figs. 2, 3). In some of our male specimens, the length of P is greater than in specimens shown by Johnson, and the spicules on the base of the eighth sternum are shorter and more numerous. There is some variation in the shape of F, especially in the dorsocaudal margin. There is no apparent difference in specimens taken from hosts and reared specimens taken from bat guano. P. dunni is extremely rare in collections and we had been unable to ob- tain it during two years of intensive collecting. There were no fleas on hundreds of specimens of M. coibensis collected in various localities through- out Panama. This may indicate that T. yucatanica is the true host and M. coibensis was infested only because of close association with T. yucatanica. We can only speculate about the length of time the bat colony has in- habited the Pacora church where we obtained the hosts and guano. Bat guano was six inches deep in places when Bloedel (1955) observed the 306 ECTOPARASITES OF PANAMA colony in 1953. Residents of the town claim that the church is very old. This is substantiated by a 1785 date inscribed on a silver incense burner which was made in Mexico especially for the church. However, it is reported that the church was rebuilt in 1941 or 1942. The estimated 1000 specimens of Tadarida yucatanica observed by Bloe- del were said to constitute the major portion of the bat population. There were also small colonies of Myotis nigricans nigricans and Noctilio labialis minor. Bloedel observed no immatures or pregnant females. In June and July of 1961 when we made our collections, small numbers of Molossops p. planirostris and Noctilio I. labialis and several hundred Tadarida yucatanica were living under the tile on the roof of the belfry of 600 550 500 450 400 350 300 250 200 150 100 50 .11. ll.mll lit 111 Im- 15 20 25 Weeks beginning June 21 30 35 40 Fig. 34. Numbers of Ptilopsylla dunni which emerged from bat guano over a 40 week period (see text). the church. Bloedel had observed the latter two species hanging from the floor joists in the attic. We observed many immature specimens of T. yucatanica. We also captured many specimens of Molossus coibensis, a few Glossophaga soricina leachii, and one Phyllostomus hastatus panamensis in a mist net near the church. We presume they came from the attic of the church where we had observed bats earlier. Bat guano was abundant under the tile on the belfry of the church and we removed approximately 45 pounds during the June collecting trip and an additional 32 pounds on a subsequent trip in July. Guano was divided into three-pound lots and placed in paper bag containers. The tops of the bags were fastened with paper clips. As the adult fleas emerged they TIPTON AND MENDEZ : FLEAS 307 moved to the surface of the guano and jumped on the sides of the paper bags. The bags were examined daily or every other day and the fleas were removed with applicator sticks, the tips of which had been dipped in 70 percent alcohol. As the guano began to dry out, wet paper towels were added to each bag after the adult fleas had been removed. At first, it appeared that this was a pure colony of Ptilopsylla dunni Kohls and during the first month over 3000 specimens were removed from the gu- ano. Eventually three male specimens of Rhynchopsyllus megastigmata Traub and Gammons emerged. The latter species was somewhat smaller and moved more sluggishly, so perhaps other specimens were present but were not observed earlier in the collecting. By late April 1962, it appeared that the guano would produce no more specimens. Over the 9-month period, 3634 adult fleas had emerged continuously from the 45-pound batch of guano. This suggests that development is arrested by unfavorable conditions or that some mechanism insures survival of the species during times when the host species is absent from the roosting place. After a small amount of guano had been sterilized in an autoclave, 16 adult fleas of both sexes were introduced into this material. Although it was examined at frequent inter- vals during the subsequent two months, no adult fleas were recovered. Additional bat guano was collected during the dry season (January 1962) and adult fleas (P. dunni) emerged at about the same rate as from the wet season material. This was also apparently true for Hormopsylla kyriophila and R. megastigmata. Figure 34 shows the number of fleas (P. dunni only) removed from the bat guano collected in June. The erratic emergence rate may be due to humidity variations. Wet paper towels were not added to the paper bags until after the first month when the guano had begun to dry out. Genus Sternopsylla Jordan and Rothschild Sternopsylla Jordan and Rothschild, 1921, Ectoparasites, 1, (3), p. 158. Type-species: Ischnopsyllus texanus C. Fox, 1914. Sternopsylla distincta speciosa Johnson. Plates 64, 65. Sternopsylla distincta speciosa Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 100, pis. 48 (figs. 3, 4), 50 (figs. 3, 8). MATERIAL EXAMINED: 11 males and 12 females from Myotis nigricans nigricans, 1 male and 3 females from mixed collection of M. nigricans nigricans and M. chiloensis, 2 males and 1 female from Tadarida brasilien- sis, 1 male and 1 female from Tadarida yucatanica, 313 males and 250 fe- males from bat guano from cave where both species of Myotis and T. brasiliensis were roosting; all from Cerro Punta (Chiriqui) except speci- mens from Tadarida yucatanica from Rancho Mojica (Bocas del Toro) ; April, May and September 1961 and March 1962, collected by C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS : Johnson (1957) indicates there is some variation in the shape of the crochet of S. distincta distincta. Some variation in this character is found in S. distincta speciosa too. Compared with her figures, the bases 308 ECTOPARASITES OF PANAMA of the crochets of our specimens tend to be broader and to taper less ab- ruptly. Our material agrees closely with her figures of the eighth and ninth sterna and the clasper. However, in the female, she shows the bursa copulatrix straight and lying in a vertical plane (S, d. distincta) , whereas in our specimens only the apical portion lies in a vertical plane and there is an abrupt bend. Species of the genus Sternopsylla are most frequently found on molossid bats and various species of Myotis. The cave from which our host bats were collected was inhabited by Tadarida brasiliensis, Myotis nigricans nigricans and Myotis chiloensis. We have not collected specimens of S. d. speciosa below 5000 feet elevation. Family Ceratophyllidae Genus Ceratophyllus Curtis Ceratophyllus Curtis, 1832, British Ent., 9: pi. 417, text. Type-species : Ceratophyllus hirundinis Curtis, 1832. Ceratophyllus altus, new species. Plate 66. DIAGNOSIS : C. altus is near C. idius Jordan and Rothschild but may be distinguished from that species as follows : the membranous apical flap of the eighth sternum has a narrow, saber-like, horizontal lobe without a fringe, while in idius this lobe is broad and fringed at the base ; the vertical lobe of the apical flap of the eighth sternum is fringed in both species but the pseudosetae are more prominent in altus (pi. 66, fig. 7) ; the movable process of the clasper extends beyond the apex of the immovable process of the clasper for about one-half its length in altus but much less than one- fourth its length in idius. DESCRIPTION, MALE: Head (pi. 66, fig. 1). — Frontoclypeal margin rounded; with moderately well developed frontal tubercle ; presetal area pitted ; with two rows of three bristles each. Eye ovate, well pigmented. Genal process subacuminate. Maxillary lobe extends to distal end of penultimate segment of maxillary palpus. Apical setae of second antennal segment about one-half length of club. Postantennal area with row of fine bristles on margin of antennal fossa ; two long bristles ; posterior row of five or six bristles interspersed with fine setae. Thorax (pi. 66, fig. 2). — Pronotum with five or six long bristles interspersed with fine setae; pronotal comb of 28-29 spines. Mesonotum (MSN.) with three rows of bristles arranged 5-5-5. Mesepisternum (MPS.) with two small medial bristles plus one minute seta near caudal margin. Mesepimere (MPM.) with three long bristles. Met- anotum (MTN.) with two rows of bristles arranged 5-3; intercalary setae between bases of bristles of second row. Lateral metanotal area (L.M.) with one long bristle. Met- episternum (MTS.) with single long bristle. Metepimere (MTM.) with three long bristles. Legs. — Procoxa with 24-25 bristles on lateral surface. Meso-metacoxae with strongly sclerotized internal rods; several long bristles in distal angle. Profemur with two bristles on outer surface plus five or six bristles on inner surface. Meso-metafemora with single bristle on outer surface ; four or six bristles on inner surface. Metatibia with about 13-15 medial bristles on outer surface. Length of metatarsal segments 270.4, 156, 104, 62.4, 124.8 M- No tarsal bristles reach beyond apex of next segment. Fifth tarsal segment of all legs covered with microsetae. TIPTON AND MENDEZ : FLEAS 309 Abdomen. — Apical spinelets on terga 1-4 paired; single on terga 5, 6. Terga with two rows bristles of about five bristles per row with intercalary microsetae between bases of second row. Basal sternum with submarginal bristles; sterna 3-7 with row of three stout bristles. Single antesensillial bristle between two minute bristles. Modified Abdominal Segments (pi. 66, figs. 3-7). — Tergum 8 large, membranous flap, almost entirely covering genitalia; with four stout bristles on dorsal margin. Eighth sternum triangular basally; punctate; becoming long, moderately narrow apically, with three apical bristles; six marginal bristles; sword-like apical flap bearing caudally directed spiculose membranous process. Immovable process of clasper (P.) with broadly rounded apex bearing three short bristles; two long acetabular bristles midway on caudal margin between apex and base. Movable finger of clasper (F.) ham-like; broad medially, tapering somewhat to broadly rounded apex; extending for almost one-half its length beyond apex of P.; with one long apical bristle surrounded by three or four smaller bristles; three additional bristles on caudal margin. Proximal arm of ninth sternum curved caudad subapically. Distal arm of ninth sternum very narrow basally, abruptly swollen into proximal lobe bearing 17-18 marginal or submarginal bristles; narrower distal lobe resting on proximal lobe bears 12-15 marginal to submarginal bristles; apex rounded, with 15-18 small setae scattered on medial surface plus three or four larger marginal setae. Aedeagus (pi. 66, fig. 4). — Aedeagal apodeme long, narrow; with long coiled apical appendage; neck short. Lateral lobe (L.L.) narrow, inapparent; median dorsal lobe (M.D.L.) bifid, open apically. Crochet (CR.) swollen ventrally; apex subacuminate, beak-shaped. TYPE MATERIAL: Holotype male (host no. 7477) from Glaucidium jar- dinii, about five miles beyond Cerro Punta on the Boquete Trail (Chiriqui) , elevation 7800 feet, 3 May 1961, collected by C. E. Yunker and C. M. Keenan. Deposited in the collection of the United States National Museum. Genus Dasypsyllus Baker Dasypsyllus Baker, 1905, Proc. U. S. Nat. Mus., 29: 129. Type-species : Ceratophyllus perpinnatus Baker, 1904. Dasypsyllus gallinulae perpinnatus (Baker). Plate 67. Ceratopsyllus gallinulae Dale, 1878, Hist. Glanville's Woot., pp. 291, 292. Ceratophyllus gallinulae, Rothschild, 1903, Ent. Month. Mag., (2), 14: 145, 146. Dasypsyllus gallinulae, Wagner, 1927, Konowia, 7: 104, 106, figs. 2, 4B. Ceratophyllus perpinnatus Baker, 1904, Proc. U. S. Nat. Mus., 27: 386, 391, 445, figs. 1-6. Dasypsyllus perpinnatus, Baker, 1905, loc. cit., 29: 129, 146. Ceratophyllus gallinulae perpinnatus, Jordan, 1926, Novit. Zool., 33: 386. Dasypsyllus gallinulae perpinnatus, Wagner, 1930, Kat. Siphonap., p. 14. MATERIAL EXAMINED: 2 males and 6 females from Sciurus granatensis chiriquensis, 5 males and 7 females from nest of Zonotrichia, capensis costari- censis, 3 males and 7 females from nest of Vireo leucophrys chiriquensis; near Cerro Punta (Chiriqui) , January and May 1960, April 1961 and March 1962, collected by C. L. Hayward, C. M. Keenan, V. J. Tipton, and C. E. Yunker. REMARKS: We have examined many birds and bird nests for fleas but have found this species only at high elevations. It is probably much more common than our records would indicate since we have concentrated on mammal collecting in the Chiriqui mountains. 310 ECTOPARASITES OF PANAMA Dasypsyllus lasius venezuelensis (I. Fox and Anduze). Plates 68, 69. Ceratophyllus lasius Rothschild, 1909, Novit. Zool., 16: 63, fig. 10. Dasypsyllus lasius, Jordan, 1933, Novit. Zool., 39: 76. Avesopsylla venezuelensis I. Fox and Anduze, 1947, Bol. Ent. Venez., 6: 108, pi. 1, figs. 1-3. Dasypsyllus lasius venezuelensis, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 117, pis. 53-55. MATERIAL EXAMINED : 193 males and 242 females from nests of Notio- chelidon cyanoleuca, 1 male and 5 females from nest of unknown host which contained both bird and rodent fleas and ticks, 1 female from N. cyanoleuca; near Cerro Punta (Chiriqui), March 1962, collected by C. L. Hayward and V. J. Tipton. REMARKS: This species was very abundant in swallow nests, both in building's as well as earth banks. In one hole in an earth bank the fleas were very numerous in the sand below the scanty nesting material. Genus Jellisonia Traub Jellisonia Traub, 1944, Field Mus. Nat. Hist, Zool. Ser., 29: 211. Type-species : Jellisonia klotsi Traub, 1944. Jellisonia johnsonae Tipton and Mendez. Plates 70, 71. Jellisonia johnsonae Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 259-262, pis. 3, 4. MATERIAL EXAMINED : 14 males and 27 females from Scotinomys teguina, 4 males from S. xerampelinus, 6 males and 3 females from Peromyscus nudipes nudipes, 3 males and 10 females from Reithrodontomys sumichrasti vulcanius, 1 male from R. creper, 1 female from R. mexicanus garichensis, 1 female from Bassaricyon gabbii gabbii, and 5 males and 5 females from nest; near Cerro Punta (Chiriqui), January, February and May 1960, January and April 1961, March 1962, collected by C. O. Handley, C. M. Keenan, V. J. Tipton, and C. E. Yunker. REMARKS: Specimens collected subsequent to the type material do not vary greatly from the original figures and description. Some two years after the type material was obtained, about 500 small animals were collected in the same general area but at a little higher elevation. There was a de- cided paucity of /. johnsonae on this group of hosts. There is much to be learned about host preferences and seasonal and altitudinal distribution of fleas in this area. Jellisonia species One female from Reithrodontomys mexicanus, Rancho Mojica (Bocas del Toro), collected by V. J. Tipton, 10 September 1961, may be J. john- sonae but the dorsal lobe of the female seventh sternum is more pronounced. Another female from Oryzomys capito, Cerro Hoya (Los Santos), collected by C. 0. Handley, 12 February 1962, is quite distinct and undoubtedly repre- sents an undescribed species. The dorsal lobe of the female seventh sternum is pronounced and the ventral margin is concave. TIPTON AND MENDEZ: FLEAS 311 Genus Pleochaetis Jordan Pleochaetis Jordan, 1933, Novit. Zool., 39: 77. Type-species : Ceratophyllus mundus Jordan and Rothschild, 1922. Pleochaetis altmani Tipton and Mendez. Plates 72, 73. Pleochaetis altmani Tipton and Mendez, 1960, Ann. Ent. Soc. Amer., 54, (2), pp. 262-263, pis. 5, 6. MATERIAL EXAMINED: A total of 58 males and 58 females as follows: 28 males and 29 females from Reithrodontomys creper, 2 males from R. sumichrasti vulcanius, 4 males and 1 female from R. mexicanus garichensis, 16 males and 12 females from Scotinomys xerampelinus, 5 males and 12 females from Peromyscus nudipes nudipes, 2 males and 2 females from Oryzomys fulvescens vegetus, 1 male and 2 females from O. albigularis near Cerro Punta (Chiriqui), collected by C. M. Keenan, V. J. Tipton, and C. E. Yunker, between January 1960 and March 1962. REMARKS: In addition to the type material collected in 1960, an addi- tional lot of 6 males and 7 females was collected in May of 1961 and a long series of 49 males and 50 females was collected in March of 1962. We have a relatively long series of specimens of this species and it is remark- able for its lack of variation. In some male specimens the dorsal margin of the crochet is straighter than in the type material and the dorsal and apical margins are slightly serrate. The outline of the female seventh sternum is quite constant. Pleochaetis dolens dolens (Jordan and Rothschild). Plates 74, 75. Ceratophyllus dolens Jordan and Rothschild, 1914, Novit. Zool., 21: 257, figs. 1, 2. Pleochaetis dolens, Jordan, 1923, Novit. Zool., 29: 27. Pleochaetis dolens dolens, Traub, 1950, Fieldiana, Zool. Mem., 1: 34-36, pis. 18, 19. Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 263-264. MATERIAL EXAMINED : A total of 194 males and 285 females as follows : 104 males and 140 females from Sciurus granatensis chiriquensis, 47 males and 58 females from Peromyscus nudipes nudipes, 8 males and 6 females from Reithrodontomys sumichrasti vulcanius, 4 males and 8 females from R. mexicanus garichensis, 2 males and 4 females from R. creper, 4 males and 5 females from Scotinomys xerampelinus, 1 male and 3 females from S. teguina, 1 male and 6 females from Oryzomys fulvescens vegetus, 1 male and 3 females from O. alfaroi alfaroi, 3 females from O. albigularis, 3 fe- males from Nyctomys sumichrasti, 1 male from Mus musculus, 1 male from Heteromys desmarestianus, 4 males and 8 females from Didelphis mar- supialis caucae, 5 males and 9 females from Bassaricyon gabbii gabbii, 1 female from Procyon lotor, 1 female from Nasua nasua, 1 female from Mustela frenata, 11 males and 23 females from nests; all collected near Cerro Punta (Chiriqui) and Rancho Mojica (Bocas del Toro) by C. M. Keenan, V. J. Tipton, and C. E. Yunker, from January 1960 to March 1962. REMARKS : This is a highly variable species but the aedeagal structures, especially the crochet, are rather constant. Fifty-one per cent of all the spe- cimens of P. dolens dolens collected in Panama have been taken from Sciurus 312 ECTOPARASITES OF PANAMA granatensis chiriquensis and 22 per cent have been collected from Peromys- cus nudipes nudipes. The table below indicates the distribution of this flea on Sciurus granatensis according to elevation : Specimens of Number of Sciurus Total P. dolens dolens Elevation granatensis P. dolens dolens per host Below 4000 ft. 63 00 4000-5000 ft. 200 5000-6000 ft. 10 18 2 Over 6000 ft. 48 227 5 There may be some seasonal variation in the number of fleas per host since there were eight specimens of P. dolens dolens per host collected during the latter part of April and early part of May at elevations in excess of 6000 feet. Genus Kohlsia Traub Kohlsia Traub, 1950, Fieldiana, Zool. Mem., 1 : 45. Type-species : Kohlsia osgoodi Traub, 1950. Kohlsia azuerensis, new species. Plates 76, 77. DIAGNOSIS: K. azuerensis n. sp. resembles K. uniseta Traub (1950) and K. graphis graphis (Rothschild) . It is easily distinguished from uniseta in possessing two acetabular bristles. It may be separated from graphis graphis in that there is a single, long, apical bristle on the reduced male eighth sternum and the distal arm of the ninth sternum is broader and more setose. In addition, the caudal margin of the female seventh sternum has a deep sinus not present in the other two species. DESCRIPTION: Head, Male (pi. 76, fig. 1). — Small frontal tubercle on broadly rounded anterior margin of head ; micropunctate preantennal area with large ovate pores cephalad of 15-16 bristles. Eye subovate, not heavily pigmented. Ovate trabecula centralis. Apex of genal process subrounded. Maxillary lobe reaches slightly beyond distal end of second segment of five-segmented labial palpus. Maxillary palpus four-segmented. Postantennal area with about 11-13 bristles arranged in three rows. Thorax (pi. 76, fig. 2). — Pronotal comb of about 18 spines caudad of row of five or six bristles. Mesonotum (MSN.) with row of three thin bristles, row of five or six slightly longer bristles, then row of three or four long bristles; about six microsetae on flange; two or three subapical pseudosetae. Mesepisternum (MPS.) with patch of four small, thin setae in cephalodorsal angle plus two or three bristles in caudoventral area. Mes- epimere (MPM.) with five bristles; does not cover lateral metanotal area. Metanotum (MTN.) with six or seven bristles arranged in two irregular rows. Lateral metanotal area (L.M.) broader than long; with two long bristles. Metepisternum (MTS.) with single long bristle. Pleural arch well defined. Metepimere (MTM.) with six or seven long bristles. Legs. — Coxae and femora not unusual. Metatibia with nine dorsomarginal notches bearing bristles arranged 2-2-1-2-2-1-1-2-3. Last segment of metatarsus with six pairs plantar bristles with sixth pair medially displaced. Abdomen. — Terga 1-4 with apical spinelets; paired on terga 2-3; two rows of bristles on all terga ranging from three to five bristles per row. Bristles on sterna 2-7 arranged l-3(4)-3-3(4)-3-3. Dorsal antesensillial bristle about three times longer than ventral bristle. TIPTON AND MENDEZ : FLEAS 313 Modified Abdominal Segments, Male (pi. 77). — Tergum 8 indistinct but appears to cover most of genitalia. Sternum 8 with short, thin process bearing long apical bristle plus short, thin, subapical bristle. Distal arm of ninth sternum broad, with well sclerotized rod reaching almost to apex; ventral lobe bearing 11-12 marginal or submarginal bristles; patch of 11 medial bristles which becomes diagonal row of additional nine or ten bristles terminating in two subapical spiniform bristles; small apical bristle plus three more small bristles on dorsal margin ; apex not produced caudad into lobe. Immovable process of clasper (P.) with slightly concave caudal margin bearing two long acetabular bristles near base. Movable process of clasper (F.) reaches apex of P.; caudal margin bears three heavy marginal bristles plus two or three small marginal bristles; several medial microsetae. Aedeagus (pi. 77, fig. 1). — Aedeagal apodeme more than twice as long as aedeagus proper; prominent proximal spur (P.S.) arising near base of blade-like accessory lateral lobe (A.L.L.) Median dorsal lobe (M.D.L.) with smoothly rounded anterior margin; convoluted to form handle-like paradorsal lobe (P.D.L.) ; primary medial dorsal lobe (P.M.D.) with flared, hyaline apex. Lateral lobes (L.L.) well developed; ventral margin bulbous. Crochet (CR.) well developed but lightly sclerotized; triangular, with apex, dorsal and caudoventral margins tufted. Sclerotized inner tube (SJ.T.) rod-like, heavily sclerotized ; with prominent, lateral, "seed-cluster" sclerite. Subacuminate, curved caudal process (A. M.S.) arising mesad of hooked anterior process of armature of inner tube (A.I.T.). Crescent sclerite (C.S.) prominent. Penis rods strongly recurved but not coiled. Modified Abdominal Segments, Female (pi. 76, figs. 3-5). — Caudal margin of seventh sternum (7S.) with acuminate dorsal lobe followed by deep sinus; ventral lobe broadly rounded; with medial row of six bristles. Eighth tergum (87\) with nine or ten long, three or four short medial bristles plus three or four marginals. Eighth sternum (SS.) not clearly defined. Anal stylet more than twice as long as broad. Head of spermatheca subglobular; tail more than twice as long as head; bent at right angle to head. Bursa copulatrix lying in diagonal plane with cephalad end inflated; bent slightly to lie in horizontal plane. TYPE MATERIAL: Holotype male (host no. 9955) from Peromyscus species near P. pirrensis, about ten airline miles west of Las Palmitas (Los Santos), elevation 3000 feet, 20 February 1962, collected by C. O. Handley. Allotype female (host no. 9934) same data as above except 17 February 1962. Paratypes : 3 males and 2 females, same data as holotype ; 1 female, same data as allotype; 2 females (host no. 9855), same data as holotype but 11 February 1962; 1 male and 1 female (host no. 9915), same data as holotype but 15 February 1962; 1 male (host no. 10040), same data as holotype but 26 February 1962. Holotype and allotype deposited in the United States National Museum ; 1 pair of paratypes deposited with each of the following named institutions and individuals : British Museum (Natural History), Chicago Natural History Museum, Gorgas Memorial Laboratory, and private collections of Robert Traub and the senior author. Kohlsia graphis graphis (Rothschild). Plate 78. Ceratophyllus graphis Rothschild, 1909, Novit, Zool., 16: 62, pi. 10, figs. 3, 4. Kohlsia graphis graphis, Traub, 1950, Fieldiana, Zool. Mem., 1: 50-51, pis. 29 (fig. 6), 30 (figs. 5-7). Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 272. MATERIAL EXAMINED: 5 males and 11 females from Sciurus granatensis chiriquensis, near Cerro Punta (Chiriqui), collected by C. M. Keenan, V. J. Tipton, and C. E. Yunker from January 1960 to March 1962. REMARKS : Our specimens differ slightly from Traub's (1950) figures. In 314 ECTOPARASITES OF PANAMA our males the distal arm of the ninth sternum is more setose and the dorsal margin distad of the sharp angle is shorter. The proximal hump on the latero-ventral, curved lobe of the apodemal strut of the aedeagus is similar to that figured for K. g. erana Traub but the apex is like K. g. graphis (Roths- child) . The caudal margin of the seventh sternum of the female has a nipple- like bump suggesting a dorsal lobe and the ventral portion of the margin is concave while Traub shows this margin to be straight. The spermatheca also has some slight differences such as the concavity in the ventral margin of the head. Our failure to collect a longer series of this species is probably related to season and altitude. A total of 60 specimens of Sciurus granatensis chiriquensis was collected from localities that are above 5000 feet and fleas were removed from 47 of the specimens. All of the 60 specimens were collected during the dry season or the early part of the wet season and none were collected above 7000 feet elevation. The majority of our specimens of K. g. graphis were collected late in the dry season or early wet season. Nests from which squirrel ticks and Pleochaetis d. dolens were collected contained no specimens of K. g. graphis. Kohlsia keenani Tipton and Mendez. Plates 79, 80. Kohlsia keenani Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 265-269, pis. 7, 8. MATERIAL EXAMINED : A total of 5 males and 13 females as follows : 2 males and 3 females from Peromyscus nudipes nudipes, 2 males and 4 fe- males from Oryzomys fulvescens vegetus, 1 male and 1 female from O. albigularis, 2 females from O. alfaroi alfaroi, 2 females from Scotinomys xerampelinus, 1 female from S. teguina, all collected by C. M. Keenan, V. J. Tipton, and C. E. Yunker near Cerro Punta (Chiriqui), January 1960 to March 1962. REMARKS: The majority of the specimens (three males and eight fe- males) were collected during the first collecting trip in May, 1960. Although hundreds of small rodents were collected during subsequent trips in a much wider variety of ecological situations, only two additional males and five fe- males were obtained. All specimens conform closely to figures given. Kohlsia mojica, new species. Plates 81, 82. DIAGNOSIS : Kohlsia mojica n. sp. is very close to K. traubi Tipton and Mendez in size, shape and setation of the processes of the clasper; in the general structure of the aedeagus ; in' the outline of the seventh sternum and shape of the spermatheca of the female. K. mojica differs from traubi in that the distal arm of ninth sternum of the male is bilobed, somewhat as in keenani Tipton and Mendez, while there is a single ventral lobe in traubi; the caudal process of eighth sternum of the male is much broader and while the crochet is roughly conical as in traubi, the top of the cone is rectangular and the bottom portion of the cone is contiguous with the proximal corner of the rectangle. DESCRIPTION: Head, Male (pi. 81, fig. 1). — Anterior margin of head broadly rounded; TIPTON AND MENDEZ : FLEAS 315 with small but distinct frontal tubercle ; preantennal area micropunctate ; three irregular rows of bristles arranged 8 (9) -5-3 with intercalary setae; eye subovate; lightly pig- mented. Genal process subrounded. Maxillary lobe reaches almost to midpoint of distal segment of four-segmented maxillary palpus. Labial palpus five-segmented; reaches to about two-thirds length of forecoxa. Postantennal area with three rows bristles arranged 3 (4) -5 (6) -5 (6). Microsetae along margin of antennal fossa. Pedicel of antenna without long bristles. Trabecula centralis prominent, ovate. Thorax (pi. 81, fig. 2). — Pronotum with row of five (six) bristles; intercalary setae between bases of bristles; about 20-21 spines in pronotal comb. Mesonotum (MSN.) with small scattered setae on anterior portion followed by medial, irregular row of six or seven bristles, then row of four stout bristles with intercalary setae ; two pseudosetae on flange. Mesepisternum (MPS.) with two to four stout bristles plus one to three fine medial setae; anterodorsal area with three to four fine setae. Mesepimere (MPM.) with about eight stout bristles. Metanotum (MTN.) with three rows bristles arranged 2(3)- 6 (5) -4; first row incomplete, second row irregular, third row with intercalary setae. Lateral metanotal area (L.M.) broader than long; with two stout bristles. Metepis- ternum (MTS.) with one long stout bristle. Squamulum well developed. Pleural arch well defined. Metepimere (MTM.) with five to seven bristles. Legs. — Procoxa with about 35-38 bristles exclusive of marginals. Outer surface of profemur with six to eight thin setae; mesal surface with single seta. Protibia with seven dorso-caudal notches bearing bristles arranged (base to apex) 1-2-2-2-1-2-3; plus five to six medial bristles. Fifth tarsal segment on all legs longest; fourth segment shortest; six pairs plantar bristles, basal and distal pairs displaced medially. Internal rods of meso- and metacoxae strongly sclerotized. Meso-, metatibiae with eight dorso- caudal notches bearing bristles arranged (base to apex) 1-2-1-2-2-1-2-3; with 10-13 medial bristles. Abdomen. — Terga 1-4 with apical spinelets; sometimes paired on terga 1-3. Tergum 1 with three rows bristles arranged 2-6-5 with some variation; terga 2-7 with two rows of bristles usually arranged 6-7 but may vary from 3-7 in first row and 5-7 in second row. Basal sternum with one seta; sterna 3-7 each with one row of three bristles, oc- casionally four bristles per row. One long antesensillial bristle between two short bristles ; ventral bristle less than one-third length of middle bristle and three times longer than dorsal bristle. Modified Abdominal Segments, Male (pis. 81, figs. 3-5; 82, figs. 4, 5). — Tergum 8 indistinct but caudal margin lies in close proximity to caudal margin of immovable process of clasper ; with two long and three shorter bristles in dorsocaudal area. Sternum 8 with well developed process ; highly variable in shape, number and position of bristles ; usually broad, with two to three apical bristles plus three to five medial bristles. Distal arm of ninth sternum with well sclerotized mesal rod running from base almost to apex ; bilobed, with well rounded ventral lobe bearing three or four stout marginal bristles plus seven or eight small medial setae; apical lobe bearing three marginal spiniform bristles plus five or six medial setae; cephalodorsal margin broadly rounded, bearing single thin bristle. Immovable process (P.) of clasper reticulate; apex short, subrounded, bearing three small bristles; caudal margin almost straight, with two long acetabular bristles. Movable finger of clasper (F.) equal to P. in length; apex subacuminate ; caudal margin evenly rounded, bearing small subapical bristle, three stout marginal bristles plus three short, thin, submarginal bristles. Aedaegus (pi. 82, fig. 4). — Aedeagal apodeme about three times longer than aedeagus proper; proximal spur (P.S.) prominent, ribbon-like; accessory spur of aedeagus (A.S.P.) small, seta-like, arising from base of slightly sinuous, acuminate accessory lateral lobe (A.L.L.). Crescent sclerite (C.S.) distinct. Median dorsal lobe (M.D.L.) inflated apically to form blade-like primary median dorsal lobe (P.M.D.) ; primary paradorsal lobe (P.D.L.) buttonhook-like. Lateral lobe of aedeagus (L.L.) with truncate apex lying near primary median dorsal lobe; with angular caudal margin. Crochet (CR.) roughly cone shaped but with beak-like and nipple-like projections on distal margin. Penis rods strongly recurved but not coiled. 316 ECTOPARASITES OF PANAMA Modified Abdominal Segments, Female (pi. 82, figs. 1-3). — Caudal margin of seventh sternum (7S.) with conspicuous sinus; dorsal lobe short, subrounded; margin ventrad of sinus almost straight; with medial row of four or five bristles per side. Eighth tergum (8T.) with about six or seven small bristles cephalodorsad of spiracle; approx- imately six long plus five shorter medial bristles; four or five caudomarginal bristles. Eighth sternum (8S.) poorly defined. Ninth sternum (9S.) inapparent. Dorsal anal lobe of proctiger with nine or ten marginal bristles plus eight medial bristles. Anal stylet about twice as long as wide; long apical bristle less than twice length of ventral bristle; dorsomarginal and medial microsetae. Ventral anal lobe with eight or nine marginal and four medial bristles. Head of spermatheca subglobular; tail almost twice as long as head, bent at right angles to head. Bursa copulatrix sinuate, cephalad end dilated. REMARKS: This species is named for the type locality, Rancho Mojica (Bocas del Toro), comprised of two thatched huts and a few coffee trees but offering a welcome respite from two days on a mountainous jungle trail. TYPE MATERIAL: Holotype male (host no. 8129) and allotype female (host no. 8136) from Peromyscus nudipes nudipes, Rancho Mojica, about 30 miles west of Almirante near the Rio Changena (Bocas del Toro) , eleva- tion 5500 feet, 13 September 1961, collected by V. J. Tipton. Paratypes: 17 males and 15 females all from (15) Peromyscus nudipes nudipes with same data as above except collected from 8 thru 13 September 1961. Holo- type and allotype deposited in the United States National Museum ; one pair of paratypes in each of: the British Museum (Natural History), Chicago Natural History Museum, Gorgas Memorial Laboratory, private collections of Robert Traub, and the senior author. Kohlsia tiptoni Mendez and Altman. Plates 83, 84. Kohlsia tiptoni Mendez and Altman, 1960, Proc. Ent. Soc. Wash., 62, (1), pp. 45-50, figs. 1-11. MATERIAL EXAMINED: 1 female from Sciurus granatensis chiriquensis , 1 female from nest, near Cerro Punta (Chiriqui) ; May 1961 and March 1962, collected by C. L. Hayward, C. M. Keenan, V. J. Tipton, and C. E. Yunker REMARKS : Very little information can be added to that given by Mendez and Altman (1960). We have collected only 2 female specimens since the type material was collected. Kohlsia traubi Tipton and Mendez. Plates 85, 86. Kohlsia traubi Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), pp. 269-272, pis. 9, 10. MATERIAL EXAMINED: 180 males and 177 females from Peromyscus nudipes nudipes, 1 male and 1 female from Oryzomys alfaroi alfaroi, 1 female from 0. fulvescens vegetus, 3 females from Sciurus granatensis chiri- quensis, 1 male from Reithrodontomys creper, 1 male from Heteromys des- marestianus, 3 females from nests, 1 female from workbench used for prep- aration of animal skins ; all collected by C. M. Keenan, V. J. Tipton, and C. E. Yunker near Cerro Punta (Chiriqui), January 1960 to March 1962. REMARKS: There is some variation in this species, but the taxonomic TIPTON AND MENDEZ : FLEAS 317 characters of significance are fairly constant, i.e., the shape of the crochet, the median dorsal lobe, the distal arm of the ninth sternum and the claspers. The shape and setation of the caudal process of the eighth sternum varies considerably (pi. 85, figs. 5, a, b) as does the caudal margin of the female seventh sternum. Of a total of 365 specimens collected, almost 98 per cent (357) were from P. nudipes nudipes. No specimens were collected below 5000 feet and they appeared to be evenly distributed from 5000 feet to 8000 feet. KEY TO THE PANAMANIAN SPECIES OF KOHLSIA TRAUB 1. Movable finger of clasper with four stout bristles on caudal margin; legs with tibial combs K. tiptoni Mendez and Altman Movable finger of clasper with three stout bristles on caudal margin ; legs without tibial combs 2 2. Length of setae on proximal lobe of distal arm of ninth sternum greater than width of proximal lobe; female with combination of following characters: length of head of spermatheca not greater than one-half length of tail; seventh sternum with no deep sinus on caudal margin and not more than four bristles K. traubi Tipton and Mendez Length of setae on proximal lobe of distal arm of ninth sternum less than width of proximal lobe ; female without above combination of characters 3 3. Distal arm of ninth sternum bilobed; caudal process of eighth sternum with more than four bristles per side ; female seventh sternum with fewer than five bristles per side 4 Distal arm of ninth sternum not bilobed (apex not bent caudad to form apical lobe) ; caudal process of eighth sternum with fewer than four bristles per side ; female seventh sternum with more than five bristles per side 5 4. Median dorsal lobe of aedeagus with beak-like process ; distal arm of ninth sternum with only one or two bristles between proximal and apical lobes ; caudal margin of female seventh sternum with dorsal lobe subtruncate K, keenani Tipton and Mendez Median dorsal lobe of aedeagus smoothly rounded, without beak-like process ; distal arm of ninth sternum with at least five or six bristles between proximal and apical lobes; caudal margin of female seventh sternum with dorsal lobe sub- acuminate K. mojica n. sp. 5. Caudal process of male eighth sternum with single apical bristle; distal arm of ninth sternum with more than 36 bristles; crochet with tufted caudal process; caudal margin of female seventh sternum with pronounced sinus . .K. azuerensis n. sp. Caudal process of male eighth sternum with two or three apical or subapical bristles; distal arm of ninth sternum with fewer than 36 bristles; crochet with fang-like caudal process; caudal margin of female seventh sternum straight or only slightly concave K. graphis graphis (Rothschild) Comments on the Genus Kolilsia Traub There are now 18 described species and subspecies of the genus Kohlsia Traub of which four species are known only from Panama. K. graphis graphis (Rothschild) is known from both Panama and Nicaragua and K. tiptoni Mendez and Altman is known from Panama and Costa Rica. There are some peculiarities in the geographical distribution of the Panama spe- cies. In the mountains of western Panama within a radius of ten miles there are three distinct species of Kohlsia which occur on the same host species, Peromyscus nudipes nudipes. In the same area K. graphis graphis occurs on squirrels. Just over 150 airline miles to the southeast another 318 ECTOPARASITES OF PANAMA species, K. mojica n. sp., is associated with Peromyscus flavidus. K. tiptoni is represented in our collections by only a few specimens so we are unable to establish true host relationships for this species. In the mountains of eastern Panama there are other Peromyscus, Sciurus and Microsciurus species and undoubtedly additional undescribed species of Kohlsia. Such a rich fauna seems unusual for a relatively small geographical area, though Panama is remarkable in its great ecological variety. Climatic factors probably influence the distribution of fleas to a much greater extent than that of the hosts. Of the three species of Kohlsia on P. n. nudipes in western Panama one (mojica) was collected on the humid At- lantic slope and the other two on the drier Pacific slope. One of the latter two species, traubi, was very abundant and the other species, keenani, quite rare (total of 18 specimens) . It may be either a nest species or the true host is unknown. We suggest another possibility to account for the distribution of Kohlsia species in Panama. It seems unlikely that this flea genus and its host genera have evolved concurrently. It may be that some of the true hosts no longer occur in this area and certain of the present hosts have been "adopted." Family Hystrichopsyllidae Subfamily Ctenophthalminae Genus Adoratopsylla Ewing Adoratopsylla Ewing, 1925, Jour. Parasit., 12: 44. Type-species: Adoratopsylla bisetosa Ewing, 1925. Adoratopsylla intermedia copha (Jordan) . Plates 87, 88. Stenopsylla intermedia copha Jordan, 1926, Novit. Zool., 33: 391, fig. 15. Adoratopsylla intermedia copha, Ewing and Fox, 1943, Fleas N. Am., p. 82. Tritopsylla intermedia copha, Costa Lima and Hathaway, 1946, Pulgas, p. 228. Jor- dan, 1950, Bull. Wld. Hlth. Org., 2: 605. Adoratopsylla (Tritopsylla) intermedia copha, Johnson, 1957, Mem. Ent. Soc. Wash., no. 5, p. 33, pis. 15, 16 (of A. intermedia ssp.). MATERIAL EXAMINED: 138 males and 185 females from Didelphis mar- supialis caucae, 199 males and 190 females from Metachirus nudicaudatus dentaneus, 25 males and 16 females from Philander opossum fuscogriseus, 3 males and 3 females from Marmosa robinsoni, 1 female from Oryzomys caliginosus and 1 female from Proechimys semispinosus panamensis; El Valle, Cerro Azul, Cerro Campana, Cerro Hoya and Cerro Punta ; July 1956 to February 1962, collected by R. M. Altman, C. 0. Handley, C. M. Keenan, V. J. Tipton, and Gorgas Memorial Laboratory. REMARKS: The host animals on which A. intermedia ssp. occurs are large and are capable of harboring large populations of fleas. As many as 152 specimens have been collected from a single host animal. The host genera, Philander, Marmosa, Metachirus and especially Didelphis, are both widespread and numerous in Central America. They occur from sea level to elevations in excess of 5000 feet. Because the hosts are ubiquitous, their geographical range is large, and they are capable of harboring large num- bers of fleas, it follows that there must be a tremendous population of this TIPTON AND MENDEZ : FLEAS 319 species of flea in South and Central America. A wide range of variation is thus expected. In a series of 60 males and 92 females collected from a single host ani- mal, Didelphis marsupialis caucae from Cerro Punta (Chiriqui), elevation 5300 feet, there is some variation in the outline of the female seventh ster- num and in the shape of the movable clasper of the male. However, all of these specimens plus 225 additional males and 224 females collected west of the Canal Zone are readily identified as Adoratopsylla intermedia copha. An additional lot of 77 males and 77 females collected at Cerro Azul and Cerro Jefe (east of the Canal Zone) are much more variable and may rep- resent a distinct subspecific population. A detailed study is necessary to properly evaluate the intra-specific populations according to hosts and geographical distribution. Subfamily Neopsyllinae Genus Strepsylla Traub Strepsylla Traub, 1950, Fieldiana, Zool. Mem., 1 : 75. Type-species : Strepsylla mina Traub, 1950. Strepsylla dalmati Traub and Barrera. Plate 89. Strepsylla dalmati Traub and Barrera, 1955, Fieldiana Zool., 37: 541-544, pis. 10 (figs. 1-3), 13 (fig. 1). Tipton and Mendez, 1961, Ann. Ent. Soc. Amer., 54, (2), p. 272. MATERIAL EXAMINED: A total of 10 males and 20 females as follows: 5 males and 9 females from Peromyscus nudipes nudipes, 2 males and 3 females from Reithrodontomys creper, 1 male from R. sumichrasti vulcan- ius, 1 male and 3 females from Scotinomys xerampelinus, 1 male and 3 females from Sciurus granatensis chiriquensis and 1 female from Oryzomys fulvescens vegetus; near Cerro Punta and on the Boquete Trail (Chiriqui) ; January and May 1960, April and May 1961 and March 1962, collected by C. O. Handley, C. M. Keenan, V. J. Tipton and C. E. Yunker. REMARKS : For the most part our male specimens are in agreement with figures and description given by Traub and Barrera (1955). Differences in the shape of the distal arm of the ninth sternum may be more apparent than real, since the shape of the membranous portion is easily distorted in mounting. The entire caudal margin of the eighth sternum is convex in our specimens whereas there is a slight concavity shown in the original figure. Both the eighth and ninth sterna as well as the movable process of the clasper are slightly more setose in our specimens than in S, dalmati, s. str. We withstood the temptation to describe the female in an earlier pub- lication (1961) because of the possibility of having two distinct species represented in our relatively short series. Now, with a much longer series, we are quite confident that our 20 female specimens represent the unde- scribed female of S. dalmati. DESCRIPTION: Modified Abdominal Segments, Female (pi. 87, figs. 5-7). — Caudal- most portion of seventh sternum (IS.) strongly convex followed by shallow concavity; row of four or five long bristles preceded by seven to nine smaller bristles. Eighth 320 ECTOPARASITES OF PANAMA tergum (ST.) with row of five bristles over spiracle; second row of five bristles cephalo- ventrad of first; four or five thin bristles caudad of antesensillial bristles. Angular margin of eighth tergum ventral to ventral anal lobe, with five stout bristles; medial area with seven stout laterad-directed bristles plus eight or nine smaller caudad-directed bristles. Eighth sternum (85.) with subacuminate apex bearing three small dorso- marginal bristles, seven thin lateral bristles plus one long bristle on ventro-caudal angle. Anal stylet more than four times as long as wide; long apical bristle flanked by two minute subapical bristles. Ventral anal lobe angulate, strongly sclerotized; with six or seven strong marginal bristles plus two or three medial bristles. Spermatheca subovate; head almost twice as wide as tail; head deeply invaginated by dorsal margin of tail ; tail strongly recurved back over head. Bursa copulatrix lying in horizontal plane for most of its length ; slightly sinuate. Subfamily Rhadinopsyllinae Genus Wenzella Traub Wenzella Traub, 1953, Jour. Wash. Acad. Sci., 43: 77. Type-species: Wenzella obscura Traub, 1953. Wenzella yunkeri, new species. Plates 90, 91. DIAGNOSIS : Wenzella yunkeri n. sp. can be easily distinguished from the type-species, Wenzella obscura Traub. In the latter species, the movable process of the clasper is narrow and extends beyond the apex of the im- movable process of the clasper for almost one-half of its length, while in yunkeri it is broad and only slightly longer than the immovable process. The eighth sternum of yunkeri does not have the broadly rounded caudal margin as does obscura and it is more setose and the bristles are longer. The distal arm of the ninth sternum in obscura is straight and finger- like and the apical bristles are short. In yunkeri the apex is inflated and there are several stout apical and subapical bristles. The proximal arm of the ninth sternum of yunkeri has a subglobular apex but is truncate though somewhat inflated in obscura. The details of the aedeagus differ greatly in the two species although the basic design is quite similar. The crochet of obscura is a broad, curved, finger-like process which is simple and un- adorned. In yunkeri it is a rod-like sclerite with a dorsal membranous flap. DESCRIPTION: Head, Male (pi. 90, fig. 1). — Anterior margin of head shallow con- vexity; preantennal area with 20—21 scattered, fine setae; ocular row of two bristles; anterior arm of tentorium prominent, angular. Eye absent. Maxillary lobe (MX.) subacuminate, reaches apex of second segment of four-segmented maxillary palpus (M.P.). Genal process projecting ventrad to midpoint on second segment of maxillary palpus. Labial palpus reaches to apex of third segment of maxillary palpus. About 15 fine setae on postantennal area plus patch of about 20 fine setae on dorsal margin and submarginal area of antennal fossa.. Second segment of antenna with marginal row of seven or eight delicate setae. Club of antenna subovate; with setae on all but first segment. Tentorial bridge prominent; long, narrow, with laterad-turned apex. Thorax (pi. 90, fig. 2). — Prontotum with two irregular rows of bristles arranged 6-8; bristles of second row longer than first. Mesonotum (MSN.) with irregular row of 10-11 small, thin bristles followed by second row of six long bristles with intercalary setae of same size as bristles of first row; phragma prominent, triangular; flange with two pseudosetae. Mesepisternum (MPS.) with single lateral bristle in caudoventral angle; three or four minute setae on cephalodorsal margin. Mesepimere (MPM.) with two bristles in cephaloventral corner; third vinculum conspicuous, vermiform. Met- anotum (MTN.) with first row of nine setae followed by second row of six stouter, longer TIPTON AND MENDEZ : FLEAS 321 bristles with intercalary setae; phragma very conspicuous, beak-like. Metepisternum (MTS.) with single, long, caudomarginal bristle; internal furca prominent, slightly bent cephalad. Metepimere (MTM.) with two long bristles; fourth vinculum crescent- shaped. Legs. — Forecoxae with scattered bristles which become progressively heavier from base to apex. Mesocoxae with complete diagonal break; prominent notch in caudoventral angle; four stout bristles plus four or five setae on distal one-fifth. Metacoxae with six stout bristles plus several marginal and submarginal setae. Femora with row of 7-12 delicate setae on ventral margin. Tibiae with six (occasionally one metatibia has seven) dorsocaudal notches with stout bristles arranged 1-2-2-2-3; approximately 10 fine setae mediad of notches. One apical bristle of third tarsal segment reaches to apex of fourth segment. Fifth tarsal segment with five pairs plantar bristles; last two pairs delicate; tarsal claws weak. Abdomen. — Tergum 1 with two rows of bristles preceded by single bristle; first row of eight bristles; second row of about six stout bristles. Terga 2-7 with three rows of bristles; first row incomplete, with four or five thin bristles; second row with 11—12 bristles; third row of eight or nine stout bristles with intercalary setae; dorsal ends of second and third rows bent cephalad. Bases of antesensillial bristles ventrally displaced; bristles slender; in male middle bristle about four times length of dorsal bristle; almost twice length of ventral bristles; in female ventral bristle subequal in length to middle bristle; dorsal bristle about one-half length of middle bristle. Sterna 2-7 each with row of five or six stout bristles preceded by patch of several smaller bristles. Modified Abdominal Segments, Male (pis. 90, figs. 3-5; 91, figs. 4, 5). — Tergum 8 with broadly rounded apex bearing three or four small marginal bristles ventrad of sensillium. Eighth sternum (8S.) a broad caudal process with subtruncate apex bearing 19-20 long marginal-submarginal bristles plus 16-17 shorter medial bristles. Immovable process of clasper (P.) with broadly rounded caudal margin; knob-like apex bearing three or four small subapical bristles; medial row of four long bristles plus six smaller bristles ventrad of these. Movable process of clasper (F.) only slightly longer than immovable process; with broadly rounded caudal margin; 20-22 small marginal- submarginal setae plus two or three medial setae. Manubrium long, very narrow. Proximal arm of ninth sternum with subglobular apex; distal arm of ninth sternum (D.A. 9) inflated subapically on ventral margin to form bulbous apex; three or four stout submarginal bristles plus four or five smaller marginal-submarginal bristles; proximo- caudal margin of arm with eight or nine weak setae. Aedeagus (pi. 91, fig. 4). — Aedeagal apodeme almost four times as long as broad; long neck; no spur or apical appendage. Median dorsal lobe (M.D.L.) straight to slightly sinuate; terminating in flap-like apicomedian sclerite (AMS). Sclerotized inner tube (S.I.T.) not well defined; with beak-like armature (A.I.T.). Crochet (CR.) consisting of well sclerotized, rod-like process with slightly swollen apex and dorsal membranous flap. Lateral lobes (L.L.) extend to base of crochet; proximoventral margin broadly rounded; distoventral margin straight. Crescent sclerite (C.S.) well defined. Penis rods recurved but not coiled. Modified Abdominal Segments, Female (pi. 91, figs. 1-3). — Seventh sternum (IS.) with single caudal lobe; row of eight stout bristles preceded by patch of about eight smaller and two larger bristles. Eighth tergum (87\) with subacuminate caudal margin produced caudad almost to apex of ventral anal lobe ; many stout bristles scattered over caudoventral area as well as several smaller bristles near spiracle. Dorsal anal lobe with dense dorsal row of marginal-submarginal bristles plus five or six long bristles on caudoventral angle below anal stylet. Anal stylet about three times as long as wide; with long apical bristle plus too short subapical bristles. Ventral anal lobe well sclero- tized; with patches of long bristles on apex and caudoventral angle. Spermatheca sub- globular; tail only slightly longer than head. TYPE MATERIAL: Holotype male (host no. 10252) from Heteromys des- marestianus, Boquete Trail, about five miles beyond Cerro Punta (Chiri- qui), elevation 6900 feet, 9 March 1962, collected by C. O. Handley and V. 322 ECTOPARASITES OF PANAMA J. Tipton. Allotype female (host no. 10417), same data as the holotype, but 7300 feet elevation, 12 March 1962. Paratypes : A total of 22 males and 21 females same data as the holotype, but from (14) H. desmarestianus, between 6800 and 7700 feet elevation and between 7 and 14 March 1962. In addition, 1 male and 1 female (host. no. 7411) same data as holotype, but 7800 feet, collected by C. M. Keenan and C. E. Yunker on 2 May 1961. Holotype and allotype deposited in the U. S. National Museum ; one pair of paratypes deposited with each of the following named institutions and individuals: British Museum (Natural History), Chicago Natural History Museum, Gorgas Memorial Laboratory, Robert Traub and the senior author. REMARKS: Wenzella yunkeri is an extremely interesting species. Al- though its generic assignment is not in doubt it is certainly distinct from W. obscura. Both species were collected from Heteromys desmarestianus at high elevations. Heteromyid rodents are rather common in Central America and there are species in South America. However, this is the southernmost extension of the subfamily Rhadinopsyllinae. This species is named for Dr. Conrad E. Yunker to whom we owe a debt of gratitude for his cooperation and assistance in collecting some of the material reported in this paper. Family Stephanocircidae Subfamily Craneopsyllinae Genus Plocopsylla Jordan Plocopsylla Jordan, 1931, Novit. ZooL, 37: 138. Type-species: Craneopsylla achilles Rothschild, 1911. Plocopsylla scotinomi, new species. Plates 92, 93. DIAGNOSIS : P. scotinomi n. sp. is a member of a species complex which includes hector Jordan, heros Jordan, ulysses Hopkins, phobos Jordan, phyl- lisae Smit, and thor Johnson. In common with these species, scotinomi lacks a spine on the genal process, like hector it has a tibial comb and in details of the male genitalia and setation it closely resembles phobos, phyllisae and thor. P. scotinomi may be easily distinguished from phyllisae in that the lat- ter species has short stubby, genal spines. The shape and position of the two large spiniform bristles on the distal arm of the ninth sternum serve to separate the remaining three species. In thor these two bristles are sepa- rated by a distance not much greater than the width of the base of one of the bristles and the ventralmost- bristle is enlarged apically so as to resemble a wedge. In phobos the distance between the two bristles is equal to or greater than the length of the distalmost of the two bristles and the proximal bristle is rod-shaped and only slightly enlarged apically. In our species the distance between the two bristles is at least twice as great as the length of the distalmost bristle. The proximal bristle is rod-shaped as in phobos but is three to four times as long as the distal bristle, not twice as long. DESCRIPTION: Head, Male (pi. 92, fig. 1). — Anterior margin of helmet broadly rounded; pre-ctenidial area narrow, with horizontal striae. Helmet comb of 14 spines in TIPTON AND MENDEZ : FLEAS 323 both sexes. Genal comb of four long spines; apex of genal process truncate but caudo- dorsal angle acute, subacuminate. Two long bristles in preantennal area. Postantennal area with four irregular rows of bristles arranged 5-4-6-7, with some variation. Bristles of scape of antenna do not reach middle of club. Maxillary lobe broad, short, does not extend to midpoint of second segment of 6-segmented labial palpus. Maxillary palpus four-segmented. Thorax (pi. 92, fig. 2). — Pronotum with two rows of bristles of five bristles each; comb of 14 spines; with broad caudoventral lobe. Mesonotum (MSN.) with row of four bristles followed by row of five bristles. Metepisternum (MPS.) devoid of bristles. Metepimere (MPM.) with four or five bristles. Metanotum (MTN.) with two rows of bristles. Lateral metanotal area (L.M.) broader than long; with two bristles. Metepis- ternum (MTS.) with one long bristle. Metepimere (MTM.) with six long bristles (four broken off in holotype). Legs. — Procoxae with scattered bristles from base to apex. Meso-metacoxae devoid of bristles except for four or five stout bristles in subapical area. Distal half of all femora with four or five lateral bristles. Partial row of dorsocaudal bristles on fore and middle tibiae. Hind tibiae with seven dorsocaudal notches with bristles arranged 2-2-3-3-3-3-4; with six lateral bristles. None of bristles on tarsal segments reach to apex of next segment. Abdomen. — First three terga with apical spinelets. All terga with first row of bristles reduced; one exceptionally long bristle on each of first three terga. Male with one antesensillial bristle which barely reaches spiracle. Two antesensillial bristles slightly longer than head of spermatheca. Modified Abdominal Segments, Male (pis. 92, figs. 3-5 ; 93, figs. 1, 2) . — Eighth tergum (8T.) short, caudal margin truncate. Eighth sternum (8S.) with short, narrow process devoid of setae. Finger-like distal arm of ninth sternum (QS.) with two modified setae about midway on caudal margin; proximal bristle approximately same diameter for its length, almost one-half length of distal arm of ninth sternum ; distal bristle short, about one-quarter length of lower bristle; two bristles separated by distance equal to three- quarters length of longer bristle; two thin apical-subapical bristles. Immovable process of clasper (P.) very prominent, extends caudad of other elements of genitalia; with about 12 long marginal bristles; about four shorter marginal, six medial bristles. Internal sclerite five times longer than wide with sides roughly parallel. Movable process of clasper (F.) bent sharply dorsad so that distal half is at right angle to proximal half; bulbous subapically with apex acuminate; medial bristle proximad of swollen portion modified, spindle-shaped, bent, striated; three or four medial microsetae, one marginal seta on bulbous portion plus subapical seta. Aedeagus (pi. 93, fig. 1). — Distolateral lobe (D.L.L.) with broad apex; internal sclerotized band reaching to apex. Crochet (CR.) poorly developed; short, narrow band enlarged slightly in middle. Lateral lobe (L.L.) long, narrow. Sclerotized inner tube (S.I.T.) short, curved caudad. Lateral sclerite (L.S.I.) spade-like. Modified Abdominal Segments, Female (pi. 93, figs. 3-5). — Caudal margin of seventh sternum (IS.) undulating but with no pronounced lobes or sinuses; with two rows of bristles of four bristles each plus three bristles cephalodorsad of rows. Eighth tergum (8T.) with five bristles in first row, three bristles in second row plus two bristles not in rows. Eighth sternum (8S.) with ventral patch of 12-14 bristles plus two additional bristles dorsad of patch. Dorsal anal lobe of proctiger reduced; with three bristles plus short spiniform cephalad of anal stylet; two bristles dorsad of anal stylet. Ventral anal lobe of proctiger with two bristles. Anal stylet almost three times as long as broad ; long subapical bristles plus two shorter apical bristles. Head of spermatheca oblong, twice as long as tail. TYPE MATERIAL : Holotype male (host no. 7409) from Scotinomys xeram- pelinus on Boquete Trail, about five miles beyond Cerro Punta (Chiriqui), elevation 7800 feet, 2 May 1961, collected by C. M. Keenan and C. E. Yunker. Allotype female, same data as the holotype. Paratypes: 1 female (host 324 ECTOPARASITES OF PANAMA no. 6832), same data but elevation 6800 feet, 7 January 1961 ; 5 males and 10 females, same data, but from 14 host animals from 6900 to 7800 feet elevation, 7 to 15 March 1962, collected by C. 0. Handley and V. J. Tipton. Holotype and allotype deposited in the United States National Museum; 1 pair of paratypes each deposited in the collections of the following in- stitutions and individuals: British Museum (Natural History), Chicago Natural History Museum, Gorgas Memorial Laboratory, Robert Traub, and the senior author. REMARKS: This is the first record of a stephanocircid flea in Central America. However, it is not an unusual finding in view of the host rela- tionships. Species of the genus Scotinomys apparently fill the same eco- logical niche in Central America as that occupied by species of the closely related genus Akodon in South America. Host-Parasite Relationships Although our data are insufficient to be conclusive, some broad patterns are apparent. In Panama the flea fauna is much richer at high elevations (above 5000 feet) . Host specificity is rather loose, two or three host genera, or perhaps a family, harboring a single species of flea. The flea fauna of a host group at low elevations, when it exists, is distinct from that of the same host group at high elevations. There are some 95 recorded species of bats in Panama representing 40 genera and seven families. Only two bat families contain species which have been found to harbor fleas even though hundreds of bats of each family have been collected. Myotis nigricans, Tadarida yucatanica, T. brasiliensis and Molossus coibensis, the only species on which fleas have been found consistently, all live in closely-knit colonies. The molossids generally live under roof tile, under eaves or in low-ceilinged caves only a few inches from their droppings, making it a comparatively simple matter for adult fleas to gain access to a host. Generally, these bat species appear to rather consistently utilize the same roosts year after year. Whether or not a bat species harbors fleas may be determined in large measure by its roosting habits. From sea level to the highest elevations where they are found in Pan- ama, marsupials are infested with fleas, but below 2500 feet they apparently have no distinctive flea fauna. Both Adoratopsylla intermedia and Juxtapu- lex echidnophagoides occur in large numbers on Didelphis marsupialis eten- sis at elevations in excess of 5000 feet, but it is likely that the latter flea species is more commonly associated with the common armadillo. Of the seven species of edentates in Panama, only one, the common armadillo, is consistently parasitized with fleas. At low elevations Rhopa- lopsyllus cacicus saevus is numerous on this host as well as in burrows, while J. echidnophagoides was collected from all host specimens obtained above 5000 feet. There is a rich and interesting flea fauna on cricetid rodents above 5000 feet elevation but virtually none on cricetid hosts collected at low elevations. This is also true of the squirrels. However, fleas are commonly TIPTON AND MENDEZ : FLEAS 325 found on one species of pocket mouse and some hystrichomorph rodents at low elevations. The number of fleas found on carnivores and larger ani- mals, including domestic animals and man, has not been great although the cat flea is abundant on cats and dogs in most localities. HOST-PARASITE LIST BY ELEVATION For each host (bold face type) listed below, column 1 shows the total number of specimens found to be infested by fleas (numerator), followed by the total number examined for parasites (denominator). Columns 2, 3, and 4 show these totals divided according to elevations of collecting local- ities. Some hosts which were found to be negative for fleas are also included. Parasite statistics (light face) indicate the numbers of males (m) and females (f ) for each species of fleas taken from the respective host, accord- ing to elevation of collecting localities. Hosts and Parasites Host From Elevations : Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. Class AVES Order STRIGIFORMES Family Strigidae Glaucidium jardinii 1/2 . . . . 1/2 Ceratophyllus altus n. sp. . . . . 1m/ Order PASSERIFORMES Family Ptilogonatidae Ptilogonys caudatus 1/1 . . . . 1/1 Dasypsyllus gallinulae perpinnatus (Baker) .. .. /If Family Hirundinidae Notiochelidon cyanoleuca 1/1 . . . . 1/1 Dasypsyllus lasius venezuelensis (I. Fox and Anduze) . . . . /If Class MAMMALIA Order MARSUPIALIA Family Didelphidae Caluromys derbianus 0/11 0/10 0/1 Monodelphis adusta 0/1 0/1 326 ECTOPARASITES OF PANAMA Hosts and Parasites Host Totals From Elevations: below 2500- over 2500 ft. 5000 ft. 5000 ft. Marmosa mexicana 0/2 0/2 . . Marmosa robinsoni 5/108 3/86 2/19 0/3 Adoratopsylla intermedia copha (Jordan) /If 3m/2f Polygenis roberti beebei (I. Fox) /If /2f Polygenis klagesi (Rothschild) 2m/lf Philander opossum 15/109 6/91 9/18 . . Adoratopsylla intermedia copha (Jordan) 2m/ 33m/28f Rhopalopsyllus australis tupinus J. and R. /If Rhopalopsyllus cacicus saevus J. and R. /5f Polygenis roberti beebei (I. Fox) 1m/ /2f •• Metachirus nudicaudatus 27/41 0/7 27/34 Adoratopsylla intermedia copha (Jordan) 199m/190f Rhopalopsyllus cacicus saevus J. and R. /If Polygenis roberti beebei (I. Fox) 1m/ Polygenis dunni (J. and R.) /2f Didelphis marsupialis 43/207 23/183 9/12 11/12 Adoratopsylla intermedia copha (Jordan) 30m/28f 108m/157f Juxtapulex echidnophagoides Wagner . . 78m/138f Hoplopsyllus glacialis exoticus J. and R. . . 5m/3f Ctenocephalides felis felis (Bouche) 2m/2f 3m/2f llm/27f Pleochaetis dolens dolens (J. and R.) 4m/8f Rhopalopsyllus lugubris lugubris (J. and R.) 7m/2f /If Rhopalopsyllus australis tupinus J. and R. 2m/5f /If Rhopalopsyllus cacicus saevus J. and R. 5m/10f lm/2f Polygenis klagesi (Rothschild) 3m/7f Polygenis roberti beebei (I. Fox) /2f 3m/5f Chironectes minimus 1/2 1/2 . . . . Rhopalopsyllus australis tupinus J. and R. 3m/3f . . . . Order INSECTIVORA Family Soricidae Cryptotis nigrescens . 0/4 Order CHIROPTERA Family Vespertilionidae Myotis albescens 0/1 Myotis chiloensis 0/1 Myotis nigricans 7/143 Sternopsylla distincta speciosa Johnson 0/4 0/1 0/94 0/1 0/1 7/48 llm/12f TIPTON AND MENDEZ : FLEAS 327 Hosts and Parasites Host Totals From below 2500 ft. Elevations : 2500- over 5000 ft. 5000 ft. Mixed collection of Myotis chiloensis (78) and Myotis nigricans (30) 3/108 3/108 Sternopsylla distincta speciosa Johnson • - lm/3f Eptesicus chiriquinus 0/1 • • 0/1 Eptesicus f uscus 0/2 • • 0/2 Rhogeessa tumida 0/14 0/14 .. Lasiurus borealis 0/1 0/1 Lasiurus ega 0/1 0/1 .. Family Molossidae Molossops planirostris 0/15 0/15 .. Tadarida brasiliensis 2/8 0/1 2/7 Sternopsylla distincta speciosa Johnson 2m/lf Tadarida yucatanica 60/88 58/86 2/2 Sternopsylla distincta speciosa Johnson . . Im/lf Ptilopsylla dunni Kohls 83m/163f . . Hormopsylla kyriophila n. sp. 2m/9f . . Rhynchopsyllus megastigmata Traub and Gammons /If • • • • Bat guano (from roosting place of Tadarida yucatanica) 3/3 3/3 Ptilopsylla dunni Kohls 2667m/5305f . . Rhynchopsyllus megastigmata Traub and Gammons 4m/3f Hormopsylla kyriophila n. sp. llm/5f . . Molossus bondae 0/32 0/32 . . Molossus coibensis 25/222 25/222 . . . . Ptilopsylla dunni Kohls 66m/129f . . Hormopsylla kyriophila n. sp. /3f .. Molossus sinaloae 0/13 0/13 Promops centralis 0/11 0/11 . . Eumops auripendulus 0/4 0/4 .. Eumops bonariensis 0/1 0/1 Bat guano (from roosting place of Myotis nigricans, M. chiloensis, and Tadarida brasiliensis) Sternopsylla distincta speciosa Johnson 1/1 1/1 313m/250f 328 ECTOPARASITES OF PANAMA Hosts and Parasites Host From Elevations: Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. 3/7 /If Im/lf /If 1/2 2m/lf 0/1 Order PRIMATES Family Hominidae Homo sapiens 14/25 11/18 Ctenocephalides felis felis (Bouche) 9m/6f Pulex simulans Baker 30m/51f Rhopalopsyllus lugubris lugubris J. and R. /If Rhopalopsyllus lugubris cryptoctenes (Enderlein) Rhopalopsyllus australis tupinus J. and R. /4f Juxtapulex echidnophagoides Wagner Order EDENTATA Family Myrmecophagidae Tamandua tetradactyla 2/7 1/5 Rhopalopsyllus australis tupinus J. and R. /If Cyclopes didactylus 0/3 Bradypus infuscatus 0/9 Choloepus hoffmanni 0/5 Cabassous centralis 0/2 Dasypus novemcinctus 18/20 Juxtapulex echidnophagoides Wagner Rhopalopsyllus cacicus saevus J. and R. 77m/104f Order LAGOMORPHA Family Leporidae Sylvilagus brasiliensis 7/14 3/9 Ctenocephalides felis felis (Bouche) /If Hoplopsyllus glacialis exoticus J. and R. Rhopalopsyllus australis tupinus J. and R. lm/2f Order RODENTIA Family Sciuridae Sciurus granatensis 46/129 0/63 Pleochaetis dolens dolens (J. and R.) Kohlsia graphis graphis (Rothschild) Kohlsia tiptoni Mendez and Altman Kohlsia traubi Tipton and Mendez Dasypsyllus gallinulae perpinnatus (Baker) Strepsylla dalmati Traub and Barrera 7/7 273m/326f 0/1 4/4 24m/47f 3/10 43/56 104m/141f 5m/llf /If /3f 2m/6f lm/2f TIPTON AND MENDEZ : FLEAS 329 Hosts and Parasites Host From Elevations: Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. Sciurus granatensis (cont.) Polygenis dunni (J. and R.) Polygenis roberti beebei (I. Fox) Rhopalopsyllus australis tupinus J. and R. Rhopalopsyllus sp. Juxtapulex echidnophagoides Wagner Sciurus variegatoides 0/6 Microsciurus alfari 0/3 Family Geomyidae Macrogeomys cavator 0/5 Family Heteromyidae Liomys adspersus 11/62 Polygenis dunni (J. and R. ) Polygenis klagesi (Rothschild) Ctenocephalides felis felis (Bouche) Heteromys australis 1/21 Polygenis klagesi (Rothschild) Heteromys desmarestianus 18/80 Pleochaetis dolens dolens (J. and R.) Wenzella yunkeri n. sp. Polygenis roberti beebei (I. Fox) Kohlsia traubi Tipton and Mendez Family Cricetidae Oryzomys albigularis 7/25 Pleochaetis altmani Tipton and Mendez Pleochaetis dolens dolens (J. and R.) Kohlsia keenani Tipton and Mendez Polygenis atopus (J. and R.) Oryzomys alfaroi 7/26 Kohlsia keenani Tipton and Mendez Kohlsia traubi Tipton and Mendez Pleochaetis dolens dolens (J. and R.) Polygenis roberti beebei (I. Fox) Oryzomys bombycinus 3/7 Polygenis roberti beebei (I. Fox) Polygenis klagesi (Rothschild) 0/5 11/61 19m/19f /2f 1m/ 1/11 /If 0/4 /If /If 1m/ 0/1 0/3 0/1 0/9 1/42 /If 3/5 3m/4f 3/7 5m/5f /2f /If 1m/ 0/5 0/1 17/34 1m/ 25m/34f 1m/ 7/25 lm/2f /3f Im/lf 1m/ 4/21 /2f Im/lf lm/3f 330 ECTOPARASITES OF PANAMA Hosts and Parasites Host Totals From below 2500 ft. Elevations : 2500- over 5000 ft. 5000 ft. Oryzomys caliginosus 19/94 1/24 18/70 Polygenis roberti beebei (I. Fox) Im/lf 13m/28f Adoratopsylla intermedia copha (Jordan) /If Im/lf . . Ctenocephalides felis felis (Bouche) /If . . Rhopalopsyllus lugubris lugubris J. and R. 1m/ Oryzomys capito 21/100 4/57 17/43 . . Polygenis roberti beebei (I. Fox) /If 26m/22f Polygenis klagesi (Rothschild) Im/lf Im/lf . . Polygenis dunni (J. and R.) /If . . Jellisonia sp. /If Oryzomys f ulvescens 12/44 0/2 0/1 12/41 Pleochaetis dolens dolens (J. and R.) lm/6f Strepsylla dalmati Traub and Barrera /If Kohlsia keenani Tipton and Mendez 2m/4f Kohlsia traubi Tipton and Mendez /2f Pleochaetis altmani Tipton and Mendez 2m/2f Nectomys alfari 2/3 2/3 Polygenis roberti beebei (I. Fox) • • 2m/4f Tylomys panamensis 5/9 3/7 2/2 Polygenis klagesi (Rothschild) llm/16f 2m/2f Nyctomys sumichrasti 1/4 0/2 1/2 Pleochaetis dolens dolens (J. and R.) /3f Reithrodontomys creper 36/67 36/67 Jellisonia johnsonae Tipton and Mendez 1m/ Strepsylla dalmati Traub and Barrera 2m/3f Pleochaetis altmani Tipton and Mendez 28m/29f Pleochaetis dolens dolens (J. and R.) 2m/4f Juxtapulex echidnophagoides Wagner 1m/ Kohlsia traubi Tipton and Mendez 1m/ Reithrodontomys mexicanus 9/40 9/40 Pleochaetis altmani Tipton and Mendez 4m/lf Pleochaetis dolens dolens (J. and R.) 4m/8f Jellisonia johnsonae Tipton and Mendez . /2f Strepsylla dalmati Traub and Barrera Im/lf Reithrodontomys sumichrasti 18/67 18/67 Jellisonia johnsonae Tipton and Mendez 3m/10f Pleochaetis dolens dolens (J. and R.) 8m/6f Pleochaetis altmani Tipton and Mendez 2m/ Strepsylla dalmati Traub and Barrera 1m/ Peromyscus flavidus 4/7 4/6 0/1 Kohlsia azuerensis n. sp. 6m/7f TIPTON AND MENDEZ : FLEAS 331 Hosts and Parasites Host From Elevations: Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. Peromyscus nudipes 182/322 . . 1/1 181/ 321 Kohlsia traubi Tipton and Mendez . . . . 180m/177f Kohlsia keenani Tipton and Mendez . . . . 2m/3f Kohlsia mojica n. sp. . . . . 18m/16f Pleochaetis dolens dolens ( J. and R.) . . . . 47m/58f Pleochaetis altmani Tipton and Mendez . . . . 5m/12f Jellisonia johnsonae Tipton and Mendez . . . . 6m/3f Strepsylla dalmati Traub and Barrera . . . . 5m/9f Polygenis atopus (J. and R.) . . . . 2m/3f Polygenis roberti beebei (I. Fox) . . /If Ctenocephalides felis felis (Bouche) . . 1m/ Zygodontomys microtinus 9/74 5/61 4/13 Polygenis klagesi (Rothschild) lm/5f 2m/3f Polygenis roberti beebei (I. Fox) . . lm/2f Polygenis dunni (J. and R.) Im/lf Rhopalopsyllus australis tupinus J. and R. Im/lf Rhopalopsyllus lugubris lugubris J. and R. /If Scotinomys teguina 16/38 .. .. 16/38 Jellisonia johnsonae Tipton and Mendez . . . . 14m/27f Kohlsia keenani Tipton and Mendez . . . . /If Pleochaetis dolens dolens (J. and R.) . . . . lm/3f Plocopsylla scotinomi n. sp. . . . . /If Scotinomys xerampelinus 37/83 . . . . 37/83 Jellisonia johnsonae Tipton and Mendez . . . . 4m/ Kohlsia keenani Tipton and Mendez . . . . /2f Pleochaetis altmani Tipton and Mendez . . . . 16m/ 12f Pleochaetis dolens dolens (J. and R.) . . . . 4m/5f Plocopsylla scotinomi n. sp. . . . . 6m/llf Strepsylla dalmati Traub and Barrera . . . . lm/4f Sigmodon hispidus 2/153 2/152 0/1 Polygenis dunni (J. and R.) 1m/ Polygenis klagesi (Rothschild) /If Family Muridae Rattus norvegicus 1/12 1/12 Pulex simulans Baker lm/4f Rattus rattus 3/110 2/97 1/13 Xenopsylla cheopis Rothschild 2m/4f Im/lf Mus musculus Linnaeus 1/17 . . . . 1/17 Pleochaetis dolens dolens (J. and R.) . . . . 1m/ 332 ECTOPARASITES OF PANAMA Hosts and Parasites Host From Elevations: Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. 4/5 8m/25f /If 1m/ 3/3 51m/64f /3f Family Erethizontidae Coendou mexicanus 4/5 Polygenis klagesi (Rothschild) Ctenocephalides felis felis (Bouche) Juxtapulex echidnophagoid.es Wagner Coendou rothschildi 0/4 0/4 Family Hydrochaeridae Hydrochaeris hydrochaeris 0/2 . . 0/2 Family Dasyproctidae Agouti paca 10/13 3/4 4/6 Rhopalopsyllus lugubris lugubris J. and R. llm/14f 4m/2f Rhopalopsyllus lugubris cryptoctenes (Enderlein) Rhopalopsyllus cacicus saevus J. and R. 1m/ Rhopalopsyllus australis tupinus J. and R. . . 7m/8f Dasyprocta punctata 23/35 Rhopalopsyllus australis tupinus J. and R. Rhopalopsyllus cacicus saevus J. and R. Rhopalopsyllus lugubris lugubris J. and R. Polygenis klagesi (Rothschild) Family Echimyidae Proechimys semispinosus Polygenis klagesi (Rothschild) Polygenis dunni (J. and R.) Polygenis roberti beebei (I. Fox) Adoratopsylla intermedia copha (Jordan) . . /If Rhopalopsyllus lugubris lugubris J. and R. /If Rhopalopsyllus australis tupinus J. and R.( 3m/llf Rhopalopsyllus cacicus saevus J. and R. lm/3f Hoplomys gymnurus 5/9 2/3 3/6 Polygenis klagesi (Rothschild) 5m/llf 25m/40f Diplomys labilis 0/3 0/3 Order CARNIVORA Family Procyonidae 18/26 5/9 64m/91f 32m /39f /2f •• 2m/3f /If lm/3f , . 314/616 294/595 20/21 815m/1153f 151m/194f 2m/lf 2m/2f Procyon cancrivorus 0/2 0/2 TIPTON AND MENDEZ I FLEAS 333 Hosts and Parasites Host From Elevations: Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. Procyon lotor 1/1 1/1 Pleochaetis dolens dolens (J. and R.) /If Nasua nasua 14/27 10/23 1/1 3/3 Pleochaetis dolens dolens (J. and R.) /If Rhopalopsyllus lugubris cryptoctenes (Enderlein) 4m/4f Rhopalopsyllus australis tupinus J. and R. 24m/31f /2f /If Rhopalopsyllus cacicus saevus J. and R. Im/lf Polygenis klagesi (Rothschild) 19m/24f Dasypsyllus gallimdae perpinnatus (Baker) 1m/ Ctenocephalides felis felis (Bouche) 5m/9f /If Hoplopsyllus glacialis exoticus J. and R. /2m Potos flavus 0/5 0/2 0/3 Bassaricyon gabbii 2/3 0/1 2/2 Pleochaetis dolens dolens (J. and R.) 5m/9f Jellisonia johnsonae Tipton and Mendez /If Family Mustelidae Mustela frenata 1/1 1/1 Pleochaetis dolens dolens (J. and R.) /If Eira barbara 2/3 1/1 1/2 Rhopalopsyllus australis tupinus J. and R. . . /If Ctenocephalides felis felis (Bouche) 4m /5f Galictis allamandi 1/1 1/1 Rhopalopsyllus australis tupinus J. and R. /If Conepatus semistriatus 1/1 1/1 Ctenocephalides felis felis (Bouche) 2m/5f Pulex irritans Linnaeus /2f Lutra annectens 0/1 0/1 Family Canidae Canis familiaris 23/30 19/23 4/7 Ctenocephalides felis felis (Bouche) 161m/412f 2m/9f Juxtapulex echidnophag aides Wagner Im/lf Rhopalopsyllus australis tupinus J. and R. /If Family Felidae Felis cattus 6/6 5/5 1/1 . . Ctenocephalides felis felis (Bouche) 135m/197f 3m/9f . . 334 ECTOPARASITES OF PANAMA Hosts and Parasites Host From Elevations: Totals below 2500- over 2500 ft. 5000 ft. 5000 ft. Felis onca 1/3 0/1 1/2 Juxtapulex echidnophag aides Wagner 19m/19f Ctenocephalides felis felis (Bouche) /If Felis pardalis 0/1 0/1 • - Felis yagouaroundi 0/1 . . 0/1 . . Order PERISSODACTYLA Family Tapiridae Tapirella bairdii Order ARTIODACTYLA Family Tayassuidae Tayassu pecari Tayassu tajacu Rhopalopsyllus australis tupinus J. and R. Family Suidae Sus scrofa Tunga penetrans (Linnaeus) Family Cervidae Odocoileus virginianus NESTS Rodent nests Jellisonia johnsonae Tipton and Mendez Pleochaetis dolens dolens (J. and R.) Bird nests Dasypsyllus gallinulae perpinnatus (Baker) Dasypsyllus lasius venezuelensis (I. Fox and Anduze) Animal burrows (probably Armadillo) Rhopalopsyllus cacicus saevus J. and R. 0/1 0/5 1/1 4/4 0/4 2/8 3/4 1/1 2/2 17m/4f 0/2 0/1 2/2 14m/5f 0/2 0/5 2/8 5m/5f 8m/16f 3/4 5m/9f 194m/247f 16/20 16/20 105m/159f References BAKER, C. F. 1895. Preliminary studies in Siphonaptera. Can. Ent., 27: 19-22. 1904. A revision of American Siphonaptera, or fleas, together with a complete list and bibliography of the group. Proc. U.S. Nat. Mus., 27, no. 1361, pp. 365-469, pis. 10-26. 1905. The classification of the American Siphonaptera. Loc. cit., 29, no. 1417, pp. 121-170. BLOEDEL, P. 1955. Observations on the life histories of Panama bats. Jour. Mam., 36, (2), pp. 232-235. BOUCHE, P. F. 1835. Beitrage zur Insectenkunde. 1. Bemerkungen iiber die Larven der zweifluge- ligen Insecten. 2. Bemerkungen iiber die Gattung Pulex. Nova Acta Acad. Leo- pold., 17: 1, fig. COSTA LIMA, A. DA, AND HATHAWAY, C. R. 1946. Pulgas : Bibliografia, catalogo e animais por elas sugados. Monog. Inst. Os- waldo Cruz, no. 4, pp. 1-522. CURTIS, J. 1824-40. British Entomology. London. 16 vols. DALE, C. W. 1878. History of Glanville's Wootton Dorset, including its Zoology and Botany. London, viii + 392 pp., 2 photos. DUNN, L. H. 1923. Fleas of Panama, their hosts, and their importance. Amer. Jour. Trop. Med., 3, (4), pp. 335-344. 1934. Entomological investigations in the Chiriqui region of Panama. Psyche, 41, (3), pp. 166-183. ENDERLEIN, G. 1912. Rothschildella, eine neue Aphanipterengattung vom Aguti. Zool. Anz., 40: 72-76, figs. 1-8. EWING, H. E. 1925. Notes on the siphonapteran genus Doratopsylla J. & R., together with a de- scription of a new genus and species of fleas. Jour. Parasit., 12: 43-46. 335 336 ECTOPARASITES OF PANAMA EWING, H. E., AND Fox, I. 1943. The fleas of North America. U. S. Dept. Agr. Misc. Publ., no. 500, pp. 1-142, figs. 1-13. FAIRCHILD, G. B. 1943. An annotated list of the bloodsucking insects, ticks and mites known from Panama. Amer. Jour. Trop. Med., 23, (6), pp. 569-591. Fox, I. 1947. Notes on ectoparasites from Venezuela (Siphonaptera and Acarina). Zo- ologica, New York Zool. Soc., 32: 117-119, figs. 1, 2. Fox, I., AND ANDUZE, P. I. 1947. A new bird flea from Venezuela (Avesopsylla venezuelensis, new genus and species). Bol. Ent. Venezolana, 6: 107-110, figs. 1-3. FULLER, H. S. 1942. Notes on neotropical Siphonaptera. Revista Ent., Rio de Janeiro, 13, (1-2), pp. 39-44. GLINKIEWICZ, A. 1907. Ergebnisse der mit Subvention aus der Erbschaft Treitl unternommenen zoologischen Forschungsreise Dr. Franz Werner's nach dem agyptischen Sudan und Nord-Uganda. X. Parasiten von Pachuromys duprasi Lat. Sitzber. Akad. Wiss. Wien, 116, (3), pp. 381-386, pis. 1, 2. HALLER, G. 1880. Rhynchopsyllus, eine neue Puliciden-Gattung, in einigen Worten gekennzeich- net. Arch. Naturg., 46, Bd. 1, pp. 72-87, text figs. 1-13. HOPKINS, G. H. E., AND ROTHSCHILD, M. 1953. An illustrated catalogue of the Rothschild collection of fleas (Siphonaptera) in the British Museum (Natural History). 1, Tungidae and Pulicidae. British Museum (Natural History), pp. i-xv, 1-361, text figs. 1-465, pis. 1-45. JAROCKI, F. P. 1838. Zoologiia czyli zweirzepotismo Ogolne podlug Naynowszego Systematu. [Zo- ology, or general descriptions of animals in accordance with the latest system.] 6: 50-52. Warsaw. (Translation of portion on ectoparasites published by Roth- schild, 1921, in Ectoparasites, 1: 129.) JENNINGS, A. H. 1910. Rats and fleas in their relation to bubonic plague (with special reference to Panama and the Canal Zone). Proc. Canal Zone Med. Assn., 54th Meeting, pp. 119-133. JOHNSON, P. T. 1957. A classification of the Siphonaptera of South America with descriptions of new species. Mem. Ent. Soc. Wash., no. 5, pp. 1-299, pis. 1-114. JORDAN, K. 1926. New Siphonaptera. Novit. Zool., 33: 385-394, figs. 1-22. 1931. Further records and descriptions of fleas from Ecuador. Loc. cit., 37: 135-143, text figs. 1-12. TIPTON AND MENDEZ : FLEAS 337 1933. A survey of the classification of the American species of Ceratophyllus s. lat. Loc. cit., 39: 70-79. 1939. On Rkopalopsylhfs Baker 1905 (Siphonaptera). Loc. cit., 41: 443-448, figs. 323-332. 1950. Notes on a collection of fleas from Peru. Bull. World Hlth. Org., 2: 597-609, figs. 1-5. JORDAN, K., AND ROTHSCHILD, N. C. 1908. Revision of the non-combed eyed Siphonaptera. Parasitology, 1: 1-100, pis. 1-7. 1914. Katalog der Siphonapteren des Koniglichen Zoologischen Museums in Berlin. I. Nachtrag. Novit. Zool., 21: pp. 255-260, 2 text figs. 1920. On American b\rd-Ceratophylli. Ectoparasites,!, (2) , pp. 65-76, figs. 65-72. 1921. New genera and species of batfleas. Ibidem, 1, (3) , pp. 142-162, figs. 116-147. 1922. New Siphonaptera. Ibidem, 1, (4), pp. 266-283, figs. 258-275. 1923a. New American Siphonaptera. Ibidem, 1, (5), pp. 309-319, figs. 312-327. 1923b. On the genera Rhopalopsyllus and Parapsyllus. Ibidem, 1, (5), pp. 320-370, figs. 328-383. KOHLS, G. M. 1942. Siphonaptera; Ptilopsylla dunni, a new species of bat flea from Panama. Jour. Parasit., 28, (5), pp. 361-362, 1 fig. LINNAEUS, C. 1758. Systema Naturae, 10th ed. Holmiae. 1, ii + 824 pp. MENDEZ, E., AND ALTMAN, R. M. 1960. A new species of Kohlsia from Central America (Siphonaptera: Ceratophyl- lidae). Proc. Ent. Soc. Wash., 62, (1), pp. 45-50, figs. 1-11. OLLIFF, A. S. 1886. Description of a new aphanipterous insect from New South Wales. Proc. Linn. Soc. N. S. Wales, (2), 1, (1), pp. 171-172. ROTHSCHILD, C. N. 1903a. New species of Siphonaptera from Egypt and the Sudan (with two plates). Ent. Month. Mag., (2), 14: 83-87, 2 pis. 1903b. Types of Siphonaptera in the Daleian collection. Loc. cit., pp. 144-146. 1904. Further contributions to the knowledge of the Siphonaptera. Novit. Zool., 11: 602-653, pis. 7-14. 1909. On some American, Australian, and Palearctic Siphonaptera. Loc. cit., 16: 61-68, pis. 10, 11. SMIT, F. G. A. M. 1958. A preliminary note on the occurrence of Pulex irritans L. and Pulex simulans Baker in North America. Jour. Parasit., 44, (5), pp. 523-526, figs. 1, 2. STILES, C. W., AND COLLINS, R. J. 1930. Ctenocephalides, new genus of fleas, type Pulex cam's. U. S. Pub. Hlth. Rpts., Wash., 45:1308-1310. TIPTON, V. J., AND MENDEZ, E. 1961. New species of fleas (Siphonaptera) from Panama. Ann. Ent. Soc. Amer., 54, (2), pp. 255-273, pis. 1-10. 338 ECTOPARASITES OF PANAMA TRAUB, R. 1950. Siphonaptera from Central America and Mexico. A morphological study of the aedeagus with descriptions of new genera and species. Fieldiana : Zool. Mem. Chicago Nat. Hist. Mus., 1, (1), pp. 1-127, pis. 1-54. 1953. Wenzella obscura, a new genus and new species of flea from Guatemala (Siphonaptera). Jour. Wash. Acad. Sci., 43, (3), pp. 77-85, figs. 1-11. TRAUB, R., AND BARRERA, A. 1955. Three new fleas of the genus Strepsylla Traub (Siphonaptera: Hystrichop- syllidae). Fieldiana, Zool., 37, pp. 541-550, pis. 10-13. TRAUB, R., AND GAMMONS, J. G. 1950. Two new fleas of the family Tungidae. Jour. Parasit., 36, (3), pp. 270-273, figs. 1-7. TRAUB, R., AND JOHNSON, P. 1952. Fleas collected during a plague survey in Venezuela. Bol. Oficina Sanit. Panamer., 32, (2), pp. 111-133, figs. 1-35. (Spanish summary, pp. 133-135). WAGNER, J. 1927. Ueber die Einteilung der Gattung Cera to phyllus Curtis. Konowia, 6, (2), pp. 101-113. 1930. Katalog der palaearktischen Aphanipteren. Vienna. F. Wagner. 55 pp. 1933. Aphanipteren-Material aus der Sammlung des Zoologischen Museums der Berliner Universitat. Mitt. Zool. Mus. Berlin Univ., 18, (3), pp. 338-362, figs. 1-21. WESTWOOD, J. 0. 1875. Description of a new pulicideous insect from Ceylon. Ent. Month. Mag., 11: 246. TIPTON AND MENDEZ : FLEAS PLATE 47. Ctenocephalides felis felis (Bouche). 1, male, head, prothorax and procoxa. 2, female, spermatheca and seventh abdominal segment. 3, male genitalia. PLATE 48. ECTOPARASITES OF PANAMA Head, prothorax and procoxa, male. 1, Tunga penetrans (Linnaeus). 2, Echidnophaga gallinacea (Westwood). 3, Juxtapulex echidnophag aides Wagner. 4, Pulex simulans Baker. TIPTON AND MENDEZ : FLEAS PLATE 49. Tunga penetrans (Linneaus). 1, male genitalia. 2, female, spermatheca and seventh abdominal segment. Echidnophaga gallinacea (Westwood). 3, male genitalia. 4, female, spermatheca and seventh abdominal segment. Juxtapulex echidnophagoid.es Wagner. 5, male genitalia. 6, female, spermatheca and seventh abdominal segment. PLATE 50. ECTOPARASITES OF PANAMA Hoplopsyllus glacialis exoticus Jordan and Rothschild, male. 1, head, prothorax and pro- coxa. 2, meso- and metathorax and first abdominal tergum. 3, eighth sternum. 4, process and movable finger of clasper. 5, modified abdominal segment. TIPTON AND MENDEZ : FLEAS PLATE 51. Hoplopsyllus glacialis exoticus Jordan and Rothschild. Female: 1, modified abdominal segments; 2, spermatheca; 3, anal stylet and ventral and lobe; Male: 4, distal arm of ninth sternum; 5, apex of aedeagus. Pulex simulans Baker, male. 6, genitalia; 7, process and movable finger of clasper. PLATE 52. ECTOPARASITES OF PANAMA Xenopsylla cheopis Rothschild, male. 1, head, prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, process and movable finger of clasper. 4, apex of aedeagus. After Johnson, 1957. TIPTON AND MENDEZ : FLEAS PLATE 53. Xenopsylla cheopis Rothschild. Female: 1, modified abdominal segments. Male: 2, distal arm of ninth sternum ; 3, modified abdominal segments. PLATE 54. ECTOPARASITES OF PANAMA MTN MSN. Rhynchopsyllus megastigmata Traub and Gammons. Male: 1, head, prothorax and pro- coxa; 2, meso- and metathorax and first abdominal tergum; 3, modified abdominal seg- ments; 4, process and movable finger of clasper; 5, distal arm of ninth sternum; 6, apex of aedeagus. Female: 7, spermatheca and modified abdominal segments. TIPTON AND MENDEZ : FLEAS PLATE 55. DL.L. CR. S.I.T. MENDEZ Polygenis atopus (Jordan and Rothschild). Female: 1, modified abdominal segments; 2, spermatheca; 3, anal stylet and ventral anal lobe. Male: 4, apex of aedeagus; 5, process and movable finger of clasper ; 6, distal arm of ninth sternum. PLATE 56. ECTOPARASITES OF PANAMA Head, prothorax and procoxa, male. 1, Polygenis dunni (Jordan and Rothschild). 2, P. roberti beebei (I. Fox). 3, P. klagesi samuelis (Jordan and Rothschild). TIPTON AND MENDEZ : FLEAS PLATE 57. 6b Polygenis dunni (Jordan and Rothschild). 1, male genitalia. 2, female, spermatheca and seventh abdominal segment. P. roberti beebei (I. Fox). 3, male genitalia. 4, female, spermatheca and seventh abdominal segment. P. klagesi samuelis (Jordan and Roth- schild). 5, male genitalia; 5a, male, distal arm of ninth sternum of specimen from Coendou mexicanus laenatus showing variation. 6, female, spermatheca and seventh abdominal segment; 6a, 6b, female, spermathecae of specimens from Coendou mexicanus laenatus showing variation. PLATE 58. ECTOPARASITES OF PANAMA Head, prothorax and procoxa, male. 1, Rhopalopsyllus australis tupinus Jordan and Rothschild. 2, R. lugubris lugubris Jordan and Rothschild. 3, R. cacicus saevus Jordan and Rothschild. TIPTON AND MENDEZ : FLEAS PLATE 59. Rhopalopsyllus australis tupinus Jordan and Rothschild. 1, male genitalia. 2, female, spermatheca and seventh abdominal segment. R. lugubris lugubris Jordan and Roth- schild. 3, male genitalia. 4, female, spermatheca and seventh abdominal segment. R. lugubris cryptoctenes (Enderlein). 3a, male, process and movable finger of clasper. R. cacicus saevus Jordan and Rothschild. 5, male genitalia. 6, female, spermatheca and seventh abdominal segment. PLATE 60. ECTOPARASITES OF PANAMA M.D.L. Hormopsylla kyriophila, new species, male. 1, head prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, apex of aedeagus. TIPTON AND MENDEZ : FLEAS PLATE 61. MENDEZ Hormopsylla kyriophila, new species. Female: 1, modified abdominal segments; 2, anal stylet and ventral anal lobe; 3, spermatheca. Male: 4, modified abdominal segments; 5, distal arm of ninth sternum; 6, process and movable finger of clasper; 7, eighth sternum. PLATE 62. ECTOPARASITES OF PANAMA Ptilopsylla dunni Kohls. Male: 1, head, prothorax and procoxa; 2, meso- and metathorax and first abdominal tergum; 3, process and movable finger of clasper. Female: 4, sper- matheca and bursa copulatrix. After Johnson (1957). TIPTON AND MENDEZ : FLEAS PLATE 63. \ Ptilopsylla dunni Kohls. Male: 1, eighth sternum; 2, apex of aedeagus; 3, crochet. Fe- male : 4, modified abdominal segments ; 5, anal stylet. After Johnson (1957) , except fig. 3. PLATE 64. ECTOPARASITES OF PANAMA MSN MTN MENDEZ Sternopsylla distincta speciosa Johnson, male. 1, head, prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, eighth sternum. 4, process and movable finger of clasper. 5, modified abdominal segments. TIPTON AND MENDEZ : FLEAS PLATE 65. M.D.L. A. IT. LPT. C.S. Sternopsylla distincta speciosa Johnson. Female: 1, modified abdominal segments; 2, spermatheca; 3, anal stylet and ventral anal lobe. Male: 4, apex of aedeagus; 5, distal arm of ninth sternum. PLATE 66. ECTOPARASITES OF PANAMA Ceratophyllus altus, new species, male. 1, head, prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, modified abdominal segments. 4, apex of aedeagus. 5, process and movable finger of clasper. 6, distal arm of ninth sternum. 7, eighth sternum. TIPTON AND MENDEZ : FLEAS PLATE 67. 1 Dasypsyllus gallinulae perpinnatus (Baker). Male: 1, head, prothorax and procoxa; 2, genitalia. Female: 3, spermatheca and seventh abdominal segment. PLATE 68. ECTOPARASITES OF PANAMA Dasypsyllus lasius venezuelensis (I. Fox and Anduze). 1, male, head, prothorax and procoxa. 2, female, spermatheca; 3, male, meso- and metathorax and first abdominal tergum. 4, female, modified abdominal segments. 5, female, anal stylet, dorsal and ventral anal lobes. 6, male, process and movable finger of clasper. After Johnson (1957) . TIPTON AND MENDEZ : FLEAS PLATE 69. Dasypsyllus lasius venezuelensis (I. Fox and Anduze), male. 1, modified abdominal seg- ments. 2, apex of aedeagus. 3, eighth sternum and distal arm of ninth sternum. After Johnson, 1957. PLATE 70. ECTOPARASITES OF PANAMA Jellisonia johnsonae Tipton and Mendez, male. 1, head, prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, modified abdominal segment. 4, process and movable finger of clasper. 5, eighth sternum. TIPTON AND MENDEZ : FLEAS PLATE 71. Jellisonia johnsonae Tipton and Mendez. Female: 1, modified abdominal segments; 2, spermatheca; 3, anal stylet and ventral anal lobe. Male: 4, distal arm of ninth sternum. 5, apex of aedeagus. PLATE 72. ECTOPARASITES OF PANAMA Pleochaetis altmani Tipton and Mendez, male. 1, head, prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, modified abdominal segments. 4, process and movable finger of clasper. 5, eighth sternum. TIPTON AND MENDEZ : FLEAS PLATE 73. L.S.I. M.D.L. S.I.T. Pleochaetis altmani Tipton and Mendez. Female : 1, modified abdominal segments ; 2, anal stylet and ventral anal lobe; 3, spermatheca. Male: 4, apex of aedeagus; 5, distal arm of ninth sternum. PLATE 74. ECTOPARASITES OF PANAMA 1 MENDEZ Pleochaetis dolens dolens (Jordan and Rothschild), male. 1, head, prothorax and pro- coxa. 2, process and movable finger of clasper. 3, eighth sternum. 4, modified abdominal segments. TIPTON AND MENDEZ : FLEAS PLATE 75. Pleochaetis dolens dolens (Jordan and Rothschild). 1, female, modified abdominal seg- ments. 2, female, anal stylet and ventral anal lobe. 3, male, distal arm of ninth sternum. 4, male, apex of aedeagus. 5, female, spermatheca. PLATE 76. ECTOPARASITES OF PANAMA MENDEZ Kohlsia azuerensis, new species. Male: 1, head, prothorax and procoxa; 2, meso- metathorax and first abdominal tergum. Female: 3, anal stylet and ventral anal lobe; 4, modified abdominal segments : 5, spermatheca. TIPTON AND MENDEZ : FLEAS PLATE 77. MENDEZ Kohlsia azuerensis, new species, male. 1, apex of aedeagus. 2, distal arm of ninth sternum. 3, process and movable finger of clasper. 4, eighth sternum. 5, modified ab- dominal segments. PLATE 78. ECTOPARASITES OF PANAMA Kohlsia graphis graphis (Rothschild). Female: 1, modified abdominal segments; 2, spermatheca; 3, anal stylet and ventral anal lobe. Male: 4, eighth sternum; 5, process and movable finger of clasper; 6, distal arm of ninth sternum; 7, apex of aedeagus. TIPTON AND MENDEZ : FLEAS PLATE 79. Kohlsia keenani Tipton and Mendez, male. 1, head, prothorax and procoxa. 2, meso and metathorax and first abdominal tergum. 3, modified abdominal segments. 4, process and movable finger of clasper. 5, eighth sternum. PLATE 80. ECTOPARASITES OF PANAMA Kohlsia keenani Tipton and Mendez. Female: 1, modified abdominal segments; 2, anal stylet and ventral anal lobe; 3, spermatheca. Male: 4, distal arm of ninth sternum; 5, apex of aedeagus. TIPTON AND MENDEZ : FLEAS PLATE 81. Kohlsia mojica, new species, male. 1, head, prothorax and procoxa. 2, meso- and meta- thorax and first abdominal tergum. 3, process and movable finger of clasper. 4, modified abdominal segments. 5, eighth sternum; a, b, variations. PLATE 82. ECTOPARASITES OF PANAMA MENDEZ Kohlsia mojica, new species. Female: 1, modified abdominal segments; 2, spermatheca; 3, anal stylet and ventral anal lobe. Male: 4, apex of aedeagus; 5, distal arm of ninth sternum. TIPTON AND MENDEZ : FLEAS PLATE 83. Kohlsia tiptoni, Mendez and Altaian, male (figs. 1, 2, 3, 6, holotype male; 4, paratype male; after Mendez and Altaian, 1960). 1, head, prothorax and procoxa. 2, modified abdominal segments. 3, 4, movable and immovable process of clasper of holotype and paratype, respectively. 5, antesensillial bristles. 6, distal arm of ninth sternum. PLATE 84. ECTOPARASITES OF PANAMA L.L L.S. Kohlsia tiptoni Mendez and Altman (1-4, allotype female; 5, holotype male). Female: 1, modified abdominal segments; 2, anal stylet and ventral anal lobe; 3, spermatheca; 4, femur and tibia of hind leg. Male: 5, endchamber of aedeagus. After Mendez and Altman, 1960. TIPTON AND MENDEZ : FLEAS PLATE 85. MENDEZ Kohlsia traubi Tipton and Mendez, male. 1, head, prothorax, and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, process and movable finger of clasper. 4, modified abdominal segments. 5, eighth sternum ; a, b, variations of eighth sternum. PLATE 86. ECTOPARASITES OF PANAMA Kohlsia traubi Tipton and Mendez. Female: 1, modified abdominal segments; 2, sper- matheca ; 3, anal stylet and ventral anal lobe. Male : 4, apex of aedeagus ; 5, distal arm of ninth sternum. TIPTON AND MENDEZ : FLEAS PLATE 87. Adoratopsylla intermedia copha (Jordan). Male: 1, head, prothorax and procoxa; 2, genitalia; 3, apex of aedeagus, specimen from Cerro Punta (Chiriqui) ; 4, apex of aedea- gus, specimen from El Valle (Cocle) ; 5, apex of aedeagus, specimen from Cerro Jefe (Panama). Female: 6, spermatheca and seventh abdominal segment. PLATE 88. ECTOPARASITES OF PANAMA Adoratopsylla intermedia copha (Jordan). Female: Variations of seventh sternum. 1-5, specimens from Cerro Punta (Chiriqui). 6-9, specimens from El Valle (Code). 10-13, specimens from Cerro Azul (Panama). TIPTON AND MENDEZ I FLEAS PLATE 89. Strepsylla dalmati Traub and Barrera. Male: 1, head, prothorax and procoxa; 2, process and movable finger of clasper; 3, apex of aedeagus; 4, distal arm of ninth sternum. Fe- male: 5, modified abdominal segments; 6, anal stylet; 7, spermatheca. PLATE 90. ECTOPARASITES OF PANAMA MENDCZ Wenzella yunkeri, new species, male. 1, head, prothorax and procoxa. 2, meso- and meta- thorax and first abdominal tergum. 3, eighth sternum. 4, modified abdominal segments. 5, process and movable finger of clasper. TIPTON AND MENDEZ : FLEAS PLATE 91. MENDEZ Wenzella yunkeri, new species. Female: 1, modified abdominal segments; 2, anal stylet and ventral anal lobe; 3, spermatheca. Male: 4, apex of aedeagus; 5, distal arm of ninth sternum. PLATE 92. ECTOPARASITES OF PANAMA MENDEZ Plocopsylla scotinomi, new species, male. 1, head, prothorax and procoxa. 2, meso- and metathorax and first abdominal tergum. 3, eighth sternum. 4, process and movable finger of clasper. 5, modified abdominal segments. TIPTON AND MENDEZ : FLEAS PLATE 93. DL.L. 1 MENDEZ Plocopsylla scotinomi, new species. Male: 1, apex of aedeagus; 2, ninth sternum. Fe- male: 3, modified abdominal segments; 4, anal stylet and ventral anal lobe; 5, sperma- theca. A Checklist of the Hippoboscidae of Panama (Diptera) GRAHAM B. FAmcniLD1 The Hippoboscidae (louse flies) of the New World have been exhaus- tively treated by Dr. Joseph C. Bequaert in a series of papers culminat- ing in his Monograph of the family (1953, 1954, 1955, 1957), which ap- peared in parts in the Entomologica Americana. Twenty-one species representing nine genera are recorded from Panama in this work. Our collecting of these insects has been incidental to other work, and only eight species have been secured. One of these appears to be a new record for Panama. The Hippoboscidae are obligate, blood-sucking ectoparasites of birds and mammals. The wings of most species are fully developed. In some genera and species, the wings are deciduous, reduced, or absent. The larvae undergo their complete growth and development within the body of the female. Except in Melophagus, fully grown larvae are deposited away from the host (e.g., in the nest) or dropped on the ground. They then form a hard, seed-like puparium. Some species of Hippoboscidae are the inverte- brate hosts of a number of blood parasites of vertebrates. The best known of these parasites are species of the genus Haemoproteus, which cause ma- laria-like diseases in birds. All but two species of the Panamanian Hippoboscidae are parasites of birds. Most of the species are only moderately host specific and generally parasitize hosts of several related species, genera or families. A few are limited to a single host species. The species known from Panama, together with their recorded hosts and their range within the country, are listed below. References are limited to Bequaert's monograph, where a full dis- cussion, descriptions, illustrations, and keys may be found. 1 Gorgas Memorial Laboratory, Panama, Panama. 387 388 ECTOPARASITES OF PANAMA Subfamily Ornithoicinae Ornithoica vicina (Walker) Bequaert (1954: 90, 574) lists it from Crypturellus, Ramphastos, and Myiodynastes, all from Chiriqui Province. It parasitizes a wide variety of hosts and occurs from Canada to Chile. Ornithoica confluenta (Say) Bequaert (1954: 107) did not record the species from Panama. We have taken it twice, both times from Casmerodius albus (Common Egret), Almirante (Bocas del Toro), 30 November 1961 and 3 February 1961, P. Galindo, collector. Subfamily Ornithomyinae Ornithoctona erythrocephala (Leach) Bequaert (1954: 187, 192) records specimens from seven unrelated birds from Chiriqui Province and San Jose Island in Panama Bay. We have taken it once from Eurypyga helias (Sunbittern), Rio Changena (Bocas del Toro), 21 September 1961, and once from Daptrius americanus (Red-throated Caracara), same locality, 7 September 1961. Ornithoctona fusciventris (Wiedemann) Bequaert (1954: 210, 213) records specimens from 10 species of hosts, nine of them Passeriformes, all from Chiriqui Province. He believes that the true breeding hosts are Passerine birds. In a supplement, Bequaert (1957 : 576) adds a number of additional host records, also all from Chiriqui Province. We have taken the species four times, twice from Rio Changena (Bocas del Toro) , from Dysithamnus puncticeps (Spot-crowned Ant Vireo) and Thryothorus atrogularis (Black-throated Wren) ; and twice from Cerro Hoya (Los Santos), from "yellow water thrush," 13 February 1962, C. O. Handley, collector, and from "wren," 14 February 1962, C. 0. Handley, col- lector. Ornithoctona nitens (Bigot) Bequaert (1954: 227, 228; 1957: 577) records specimens from several species of Trogonidae and a house wren, all from Chiriqui Province. Bigot's types were from Panama. Stilbometopa ramphastonis Ferris Bequaert (1955: 254) records the species from Ramphastos swainsonii (Chestnut-mandibled Toucan) and from a dove. The type came from Chiri- qui Province, and Bequaert saw other specimens from Gamboa (Canal Zone) and Tapia (Panama). We have taken a single female, which is be- lieved to be this species, from Monasa morphoeus grandior (White-fronted Nunbird), Rio Changena (Bocas del Toro) 20 September 1961. FAIRCHILD : HIPPOBOSCID FLIES 389 Lynchia americana (Leach) Bequaert (1955: 272, 291) records the species from Bubo virginianus (Great Horned Owl) from Chiriqui Province, and without host from Juan Diaz (Panama). Bequaert believes the true hosts to be owls, hawks, and gallinaceous birds. Lynchia angustifrons (van der Wulp) Bequaert (1955: 305, 307) records a number of specimens from hawks and toucan, from localities in and near the Canal Zone and from Chiriqui Province. We have taken a single male from Ramphastos swainsonii (Chestnut-mandibled Toucan), Rio Changena (Bocas del Toro), 8 Septem- ber 1961. Bequaert believes hawks probably are the main breeding hosts, possibly also owls and toucans. Lynchia wolcotti (Swenk) Bequaert (1955: 311, 312) records from Chiriqui and Bocas del Toro Provinces, 3 lots, all from hawks, which appear to be the principal, if not the only, breeding hosts. We have 1 female, from Otus guatemalae ( Vermic- ulated Screech Owl), Rio Changena (Bocas del Toro), 21 September 1961. Lynchia nigra (Perty) Bequaert (1955: 314, 319) lists one record, from Buteogallus anthra- cinus (Common Black Hawk) from Pacora (Panama). Lynchia albipennis (Say) Bequaert (1955: 330, 336) reports the species from Cochlearius co- chlearius (Boat-billed Heron) from Puerto de Chorrera (Panama) ; also from Ancon and Christobal (Canal Zone), no host. Wading birds of the Order Ciconiif ormes are the only true hosts. Microlynchia crypturelli Bequaert Bequaert (1955: 377) described the type specimens from Crypturellus soui (Little Tinamou) from La Vaca (Chiriqui). The only other known records of this rare species are from Southern Brazil, also mainly from Crypturellus. Pseudolynchia canariensis (Macquart) Bequaert (1955 : 390, 397) saw material from Ancon (Canal Zone) , and during the last war, the United States Army Signal Corps pigeons at Quarry Heights (Canal Zone) were heavily infested by this world-wide species. The only known host is the domestic pigeon, at least in the western hemi- sphere. Pseudolynchia brunnea (Latreille) Bequaert (1955: 411, 414) records specimens from Chordeiles acuti- pennis (Lesser Night Hawk) from Barro Colorado Island (Canal Zone). 390 ECTOPARASITES OF PANAMA We have 1 male and 2 females from 'capacho,' Caprimulgus rufus (Rufous Nightjar), Rio Changena (Bocas del Toro), 16 September 1961. Bequaert notes that this species is known almost exclusively from Caprimulgiformes. Olfersia spinifera (Leach) Bequaert (1957: 429, 430) notes specimens from Panama and Tabo- guilla Island in Panama Bay, no host. It is a specific parasite of frigate birds (Fregata), and only occasionally parasitizes other marine birds. Olfersia aenescens C. G. Thomson Bequaert (1957: 437), gives one record from Panama from Sterna fiiscata (Sooty Tern). The chief hosts are a variety of marine birds. Olfersia bisulcata Macquart Bequaert (1957: 457, 459) records a series of captures of this species, all from vultures of several species, of which it is a specific parasite. He notes it as a common species. There are specimens in the Gorgas Memorial Laboratory collection from turkey buzzard collected by L. H. Dunn. Re- corded localities suggest the species is well distributed in Panama. Olfersia fossulata Macquart Bequaert (1957: 464, 465) gives a single record for the Canal Zone, without host. He states it to be a common parasite of marine birds. Olfersia sordida Bigot Bequaert (1957 : 469, 470) lists 5 records for coastal Panama and Barro Colorado Island, the hosts being pelicans and, in one case, a cormorant. Pelicans appear to be the chief host. Olfersia coriacea van der Wulp Bequaert (1957: 474, 476) gives two records for this species, Juan Diaz, no host, and Upper Chagres River, from Agriocharis ocellata (Ocel- lated Turkey). This last seems to be an error, as this bird is not reported south of Guatemala. The species seems to occur only on Galliformes. Subfamily Melophaginae Lipoptena (Lipoptenella) mazamae Rondani Bequaert (1957: 499, 501) refers to earlier records and adds Panama City, no host. Dunn (1934) took specimens from Mazama at Camp Pital (Chiriqui) , and we have taken it several times from Mazama, Rio Chucuna- que (Darien), 18 February 1958, P. Galindo, collector; from Odocoileits, Cerro Hoya (Los Santos), 9 February 1962, C. 0. Handley, collector; from Mazama americana (Red Brocket), upper Rio Mono, Rio Tuira Basin (Darien), 25 and 27 June 1962, A. Adames, collector; and from Odocoileus virginianus (White-tailed Deer), Los Santos Province, 20 February 1962, V. J. Tipton, collector. It is a specific parasite of deer. FAIRCHILD : HIPPOBOSCID FLIES 391 Melophagus ovinus (Linnaeus) Bequaert (1957: 507, 516) reports one record for Panama, but the species does not thrive in the moist tropics. It is a specific parasite of domestic sheep and is world-wide in distribution. References BEQUAERT, J. 1942. A monograph of the Melophaginae, or ked-flies, of sheep, goats, deer and ante- lopes. Ent. Amer., (n.s.), 22: 1-220. 1953-1957. The Hippoboscidae or Louse-flies (Diptera) of mammals and birds. Part I. Structure, physiology and natural history. 1953, Ent. Amer., (n.s.), 33: 1-209, 211—442, figs. 1-21. Part II. Taxonomy, evolution and revision of American gen- era and species. 1954, ibidem, 34: 1-232, figs. 22-44. 1955, ibidem, 35: 233-416, figs. 45-82. 1957, ibidem, 36: 417-611, figs. 83-104. 392 Nycteribiid Batflies from Panama (Diptera: Nycteribiidae) LlNDOLPHO R. GUIMARAES l Two species of nycteribiids were hitherto known from Panama, Basilia myotis and B. dunni, both described by Curran (1935) from specimens taken from Myotis nigricans. In the present paper we add five more species of Basilia, of which two are new. Genus Basilia Miranda Ribeiro Basilia M. Ribeiro, 1903, Arch. Mus. Nac. Rio de Janeiro, 12: 177. Pseudelytromyia M. Ribeiro, 1907, ibidem, 14: 233. Guimaraesia Schuurmans-Stekhoven Jr., 1951, Acta Zool. Lilloana, 12:109. Type-species : Basilia ferruginea M. Ribeiro, 1903. Only two genera of nycteribiids are known from the New World, Basilia M. Ribeiro, 1903, and Hershkovitzia Guimaraes and D'Andretta, 1956. They are easily distinguished by several characteristics. However, the separation between Basilia and some Old World genera is rather subtle and is based primarily on the presence of two-faceted eyes in Basilia. The fol- lowing characters are common to all New World Basilia: Anterior region of head (vertex) sclerotized; eyes two-faceted; palps with sub-parallel margins. Noto-pleural sutures parallel, with lateral plates. Sternal plate with one suture on each side of the lateral margins. Tarsal segment I much longer than the remaining segments taken together. Abdomen of female with two or three sclerotized plates (visible tergites) on the dorsal side. KEY TO PANAMANIAN SPECIES FEMALES 1. Tibiae of all legs with four or five transverse rows of setae on the ventral margin. Tergal plate I transformed into two elongate lobes with short and long setae 2 Tibiae with three rows of setae. Tergal plate II transformed into two elongate lobes with short and long setae or with posterior margin arcuate 3 1 Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brasil. 393 394 ECTOPARASITES OF PANAMA 2. Tibiae with four transverse rows of setae on ventral margin. Tergal plate II with sparse setae next to the median longitudinal suture. Posterior elevation of mesonotum without a median digitiform process ferruginea M. Ribeiro Tibiae with five rows of setae. Tergal plate II with numerous setae next to the median suture. A small digitiform process on middle of posterior elevation of mesonotum handleyi n. sp. 3. Tergal plate II transformed into two elongate lobes 4 Tergal plate II with arcuate posterior margin 5 4. Lobes of tergal plate II very thin. Lateral connexivum of abdomen (dorsal) with short pustulate setae and with a converging row of larger setae wenzeli Guimaraes and D'Andretta Lobes of tergal plate II not very thin. Lateral connexivum of abdomen with medium and long pustulate setae tiptoni n. sp. 5. Tergal plate I short, with few discal setae. Each tergite of tergal plate I more than twice as long as broad. Third sternite obsolete anceps Guimaraes and D'Andretta Not so 6 6. Long setae of posterior margin of the tergal plate I reaching or crossing the pos- terior border of the tergal plate II. Discal setae of tergal plate II on lateral halves of tergite myotis Curran Long setae of posterior margin of the tergal plate I not reaching posterior margin of the tergal plate II. Discal setae of tergal plate II in irregular rows con- verging obliquely toward the mid-line dunni Curran Basilia ferruginea M. Ribeiro Basilia ferruginea M. Ribeiro, 1903, Arch. Mus1. Nac., 12: 179, pi. I, figs. 1, 2. M. Ribeiro, 1907, ibidem, 14: 231, pi. XXIV, fig. 1. Ferris, 1924, Ent. News, 35: 195. Stiles and Nolan, 1931, Bull. Natl. Inst. Health, no. 155, p. 648. Schuurmans- Stekhoven Jr., 1931, Zeitschr. Parasitenk., 3: 216. Curran, 1935, Amer. Mus. Nov., no. 765, p. 2. Scott, 1936, Linn. Soc. Jour., Zool., 39: 497, 503, fig. 11. Del Ponte, 1944, Ann. Med. Reg., 1: 118, 124. Guimaraes, 1946, Arq. Zool. Est. S. Paulo, 5: 14, 19, 20, figs. 12-20; Guimaraes and D'Andretta, 1956, ibidem, 10: 28, figs. 5, 6, 74. Previous records : Brazil, from Lasiurus borealis bonariensis; Paraguay, from unknown host ; Cuba, from Lasiurus pfeifferi. PANAMANIAN MATERIAL EXAMINED: One male and 2 females from Lasiurus borealis frantzii, (host no. 11699), Armila (San Bias), 17 March 1963, collected by C. O. Handley. REMARKS : Basilia ferruginea is characterized by the length of the pus- tulate setae of the lateral abdominal connexivum and by the presence of four transverse rows of setae on the ventral margin of all tibiae. The male has the same chaetotaxy on the tibiae and a large number (ca. 40) of spini- form setae on the posterior margin of the fourth abdominal sternite. Basilia handleyi, new species. Figure 35A. This species closely resembles B. ferruginea, but differs in having : much larger size; more abundant pilosity; a digitiform process on the posterior elevation of the mesonotum ; 14 or 15 notopleural setae instead of eight or 10; five, instead of four, rows of setae on the tibiae; the discal setae of second tergal plate more numerous and concentrated along the mid-line; anal segment more conical and pilose ; pustulate setae on lateral connexivum of abdomen more abundant, longer and thicker; 65 to 68 spines of the GUIMARAES : NYCTERIBIID BATFLIES 395 ctenidium of the distal margin of the basal sternite; two, instead of six, setae on the genital plate. DESCRIPTION, FEMALE : Head. — Vertex with three or four pairs of setae near the an- terior margin, between the eyes; anterior margin of each gena with eight setae. Post- gena with five or six scattered short setae. Palpi with two terminal setae (one very long) and three or four pairs near the lateral margin on the ventral side; one of the eyes typically with two ocelli, the other with only one ocellus. The paratype has two ocelli on the two eyes. Thorax. — Much wider than long (1.42 x 0.96 mm.). Mesonotum raised posteriorly, Fig. 35. Basilia handleyi, new species. A, dorsal view of abdomen, female. ceps, male. B, last two tergites, and C, terminal sternite. Basilia an- with a conspicuous median digitiform process; 14 to 16 notopleural setae; thoracic ctenidium with 22 or 23 spines. Legs long, little laterally compressed. Tibiae scalpel- shaped, with five rows of setae ventro-distally. The proximal row, although less well- developed than the others, is rather conspicuous. Abdomen. — Tergal plate I wider than long, with posterior margin slightly sinuate, paralleled by thick setae of medium length and showing, on each side of the mid-line, several blunt discal setae on the posterior half. Tergal plate II longitudinally divided; each half ending posteriorly as a lobe with three or four long, thick setae and some spine-like setae; lateral margins with medium and long setae; discal setae arranged in several irregular rows along the mid-line and clustering on the antero-lateral corners. Anal segment truncate, conical with numerous long, thick setae on each side of the 396 ECTOPARASITES OF PANAMA mid-line. Lateral connexivum with long and medium pustulate setae. Ctenidium of hind margin of basal sternite with 65 to 68 spines. Behind the basal sternite there are five others, the two anterior ones limited posteriorly by a row of setae longer than the discals. Fourth sternite represented by two plates more sclerotized than the connexivum, separated medially and showing two rows of setae, the posterior row denser and stronger. Fifth sternite also made of two plates separated in the middle and with setae as in fourth. Distal sternite trapezoid, more than 2.5 times as broad as long; discal setae in four or five irregular rows on posterior two-thirds ; lateral and posterior margins paral- leled by setae that are a little longer than the discal setae. Only two genital setae, connected to the anal sclerite by a strip slightly more sclerotized than the connexivum. Length, 2.72 mm. MALE: Unknown. TYPE MATERIAL: Holotype female from Lasiurus castaneus Handley (host no. 11176), Armila (San Bias), collected 23 February 1963, by C. 0. Handley. In the collection of Chicago Natural History Museum. Paratype female, same data as the holotype but collected 26 March 1963 (host no. 11911), deposited in the Departamento de Zoologia, Secretaria da Agri- cultura, Sao Paulo, Brazil. REMARKS : This is the only New World species of Basilia that has tergal plate II transformed posteriorly into two lobes and the fifth sternite longi- tudinally divided. All other species with lobes on tergal plate II have a whole fifth sternite. Basilia wenzeli Guimaraes and D'Andretta Basilia wenzeli Guimaraes and D'Andretta, 1956, Arq. Zool. Est. S. Paulo, 10: 42, figs. 25-33, 57, 80. Previous records: Venezuela, from Eptesicus fuscus and Lonchorhina aurita; Columbia, from Histiotus sp. PANAMANIAN MATERIAL EXAMINED: From Eptesicus brasiliensis pro- pinquus: Sibube (Bocas del Toro), 1 male on 16 January 1963 (host no. 10659), 3 males and 4 females on 17 January 1963 (host no. 10706), and 1 male on 25 January 1963 (host no. 10935) all collected by C. O. Handley; Los Santos Province, 1 male and 1 female (host no. 10029), 26 February 1962, V. J. Tipton; Armila (San Bias), 1 female (host no. 11761), 20 March 1963, C. 0. Handley. From Artibeus j. jamaicensis, Cerro Hoya (Los Santos), 1 female, 18 February 1962, V. J. Tipton. REMARKS : The female of this species differs from all others by the great length of the terminal segment and by the forward position of the anal segment. The posterior lobes of tergal plate II are rather thin, a character which indicates a close relationship of this species to B. corynorhini Ferris, 1916. Basilia tiptoni, new species. Figure 36. Very close to B. silvae (Brethes, 1913) , from which it differs in having: 12, instead of eight or ten, notopleural setae ; a small digitiform process on the posterior elevation of the mesonotum ; longer and more numerous pustu- late setae on the lateral connexivum ; lateral margins of anal segment sub- parallel (convergent in silvae). The main difference, however, is in the shape of tergal plate I, which has two median projections on the posterior GUIMARAES : NYCTERIBIID BATFLIES 397 border, each projection bearing- two setae. The male of B. tiptoni n. sp. has only 11 spiniform setae on the posterior margin of the fourth abdominal sternite ; there are 20 in silvae. DESCRIPTION, FEMALE: Head. — Anterior margin of vertex slightly concave, paralleled by two pairs of setae. Anterior margin of each gena with six or seven setae ; postgena with five or six scattered setae on each side. Palpi with a long apical seta and eight uneven setae near the sides of the ventral aspect. Eyes typically two-faceted. Fig. 36. Basilia tiptoni, new species, and C, claspers, male. A, dorsal view of abdomen, female. B, abdomen, Thorax. — Wider than long (1.08 x 0.95 mm.). Mesonotum raised posteriorly, with a median digitiform process; 12 notopleural setae. Thoracic ctenidium with 18 or 19 spines. Legs long and little laterally compressed; tibiae scalpel-shaped with three rows of setae on the distal half of the ventral aspect. Abdomen. — Posterior margin of tergal plate I with two slight lobes, each with two setae (deciduous in the type) ; three or four setae on each side of the posterior margin; some scattered spine-like setae on the disc. Tergal plate II longitudinally divided; each half ending posteriorly in a lobe with four long, thick setae; discal setae concentrated along the median suture and on the outer halves of the tergites. Anal segment with subparallel sides, bearing 11 or 12 thick setae on each side. Lateral connexivum with medium and long pustulate setae, some of which are as long as the discal setae of tergal plate II. Ctenidium of basal sternite with 44 or 45 spines. Posterior to the basal 398 ECTOPARASITES OF PANAMA sternite are five other sternites, well set off by the setae on their posterior margins. The second sternite is completely covered with setae; the third has, in addition to the setae of the hind margin, only a few lateral setae; the fourth sternite is divided into two plates medially and bears one row of setae; the fifth is entire and bears two irregular rows of setae. The terminal segment is wider than long, with rounded margins paral- leled by 13 or 14 uneven setae; the discal setae are few (four or five), restricted to the posterior half. Genital plate with three setae. Length, 2.59 mm. MALE : Head, thorax and legs as in the female ; however, in the single male specimen, the vertex bears three pairs of setae instead of two. Thorax slightly broader than long (1.03 x 0.93 mm.) and has no digitiform process on the mesonotum. It is possible to recognize seven abdominal tergites; the first is short, has two or three rows of small discal setae and a row of slightly larger setae on the hind margin; the second is longer, with 12 or 13 discal setae arranged in two irregular rows, plus, on each side, a group of setae as long as or shorter than the discals, and still, on the posterior margin, a row of interspersed medium and short setae; third dorsal segment with a posterior row of setae and four or five discals ; fourth to sixth segments with setae only along the posterior margin ; terminal tergite with one or two irregular rows of setae on the posterior margin. Ventrally, five sternites are recognizable; the ctenidium of the basal sternite has 44 or 45 spines; the second sternite has three irregular rows of discal setae and one row of larger ones on the posterior margin ; the third sternite has setae only along the posterior margin, while the fourth sternite, besides these, has an irregular row of discals near the posterior margin plus a group of 11 spiniforms grouped on the mid-line. Length, 2.39 mm. TYPE MATERIAL : Holotype female from Lonchorhina or Tonatia (host no. 8288) 22 miles south of Changuinola (Bocas del Toro), collected 6 Septem- ber 1961 by V. J. Tipton and C. M. Keenan. In the collection of Chicago Natural History Museum. Allotype male, same data and repository as holotype. ADDITIONAL MATERIAL EXAMINED : The following seem also to belong to this species : 1 male, same data as the holotype but from a bat "like Tonatia" (host no. 8294) ; 1 male from Mimon crenulatum keenani (host no. 10724) , Sibube (Bocas del Toro) , 18 January 1963, C. O. Handley, Jr. Basilia myotis Curran Basilia bellardii Schuurmans-Stekhoven Jr., 1931 (nee Rondani, 1878), Zeitschr. Parasitenk., 3, (2), p. 207, figs. 1-6. Hase, 1931, ibidem, p. 220, figs. 1-17. Scott, 1936, Jour. Linn. Soc., Zool., 39: 497 (partim). Bequaert, 1942, Bol. Ent. Venez., 1, (4), p. 83. Guimaraes, 1946, Arq. Zool. Est. S. Paulo, 5: 15, 62 (partim). Karaman, 1948, Rad. Acad. Yougosl., 273: 42, fig. 4. Basilia myotis Curran, 1935, Amer. Mus. Novit., no. 765, p. 3, figs. 3-5. Scott, 1936, Linn. Soc. Jour., Zool., 39: 497. Bequaert, 1942, Bol. Ent. Venez., 1, (4), p. 84. Del Ponte, 1944, Ann. Inst. Med. Reg., L (1), pp. 118, 124. Guimaraes, 1946, Arq. Zool. Est. S. Paulo, 5: 16, 19. Guimaraes and D'Andretta, 1956, ibidem, 10: 76, figs. 85, 106-111, 124, 129, 145. Guimaraesia bellardii Schuurmans-Stekhoven Jr., 1951, Acta Zool. Lilloana, 12: 112, fig. 4. Previous records : Guatemala, from Myotis nigricans (probably subsp. nigricans) , Molossus sp. (bondae?) ; Colombia, from Myotis nigricans (probably subsp. nigricans), Uroderma bilobatum; Peru, from unknown bat; Venezuela, from Myotis sp., Myotis n. nigricans, Dasypterus sp., Molossus crassicaudatus, unknown bat; British Guiana, from Myotis n. GUIMARAES : NYCTERIBIID BATFLIES 399 nigricans; Panama, type locality and host — Tapia, from Myotis nigricans (probably subsp. nigricans) — and Camoganti (Darien), from Myotis n. nigricans. PANAMANIAN MATERIAL EXAMINED: From Myotis n. nigricans, in the Canal Zone, collected by V. J. Tipton ; Gamboa, 2 females (host no. 6411) , 23 September 1960; Fort Davis, 1 male and 2 females (host no. 3934) on 28 July 1959, 1 male and 3 females (host no. 4767) on 14 October 1959, 3 lots of 1 male, 1 female, and 2 females (host nos. 5171, 5173-4) on 7 January 1960; Barro Colorado Island, 14 males and 15 females (host no. 5595) on 12 June 1960. REMARKS : The citations of B. bellardii as synonyms of this form are due to Schuurman-Stekhoven Jr., who described (1931) as Basilia bellardii what Curran described (1935) as B. myotis. Basilia myotis replaces B. speiseri in northern South America, as the most common nycteribiid of Myotis n. nigricans. It ranges from British Guiana south to Peru and north to Guatemala. The two species are morphologically very similar. However, B. myotis has a longer tergal plate I, with discal setae only on anterior two-thirds of the tergite and longer and more nu- merous setae on the posterior margin, the latter setae extending well be- yond crossing the hind margin of tergite II. Guimaraes and D'Andretta (1956, p. 76) have suggested that B. myotis may be a synonym of B. ferrisi Schuurman-Stekhoven Jr., 1931. Basilia dunni Curran. Figure 37. Basilia dunni Curran, 1935, Amer. Mus. Nov., no. 765, p. 3, figs. 1-2. Scott, 1936, Linn. Soc. Jour., Zool., 39: 497. Del Ponte, 1944, Ann. Inst. Med. Reg., 1, (1), pp. 118, 124. Guimaraes, 1946, Arq. Zool. Est. S. Paulo, 5: 16, 20. Guimaraes and D'Andretta, 1956, ibidem, 10: 95, figs. 152, 153. Previous Panamanian records: Santa Rosa (Colon), from Myotis n. nigricans (type locality and host) . MATERIAL EXAMINED: 1 male and 5 females from Myotis n. nigricans (host no. 5688) Juan Mina (Canal Zone), 28 July 1960, V. J. Tipton; 2 males and 2 females from Myotis albescens (host no. 6375), Rio Tuira (Darien), 2 March 1958, V. J. Tipton, and 2 females (host no. 6378), same data but 5 March 1958. REMARKS : This species was described from a single female ; it was re- described and figured by Guimaraes and D'Andretta (1956). There are, in the region, two species whose females may be confused with that of B. dunni, namely, myotis Curran and costaricensis Guimaraes and D'Andretta, 1956. B. dunni differs from myotis in having tergal plate I relatively shorter and with a larger number of discal setae, in the distribution of the discal setae of tergal plate II, in the shape of the anal segment, and in the chaetotaxy of the fourth sternite. Basilia costaricensis, although somewhat similar to dunni, is easily characterized by the chaetotaxy of the fourth and fifth abdominal sternites and by the shape of the distal sternite. It also has a small digitif orm process on the posterior elevation of the mesonotum ; this process is absent in dunni. 400 ECTOPARASITES OF PANAMA However, the species most closely related to dunni is carteri Scott, 1936, from Paraguay, southern Brazil, northern Argentina, and southern Bolivia. The differences are not pronounced but are constant. They are primarily in the shape and chaetotaxy of tergal plate I, which in dunni is more slender posteriorly and has a less rounded posterior margin ; the discal setae of this tergite are more numerous and extend close to the mid-line, not leaving the median longitudinal glabrous strip characteristic of carteri. Until now only the female of B. dunni was known ; we here describe the male. DESCRIPTION, MALE: Head. — Vertex with three pairs of setae forming two diverging lines between the eyes (the allotype has five setae on one row) ; anterior border of each gena with seven or eight setae ; same number of scattered setae on postgena. Palpi with 12 or 13 uneven setae on ventral aspect and one long setae on apex. Eyes typically two- faceted. Thorax. — Broader than long (0.95 x 0.78 mm.) with anterior margin distinctly arcu- ate and posterior slightly sinuate. Thoracic ctenidium with 18 or 19 spines. Twelve setae on notopleural sutures. Legs long, little laterally compressed; all tibiae with three transverse rows of setae on ventral margin. Abdomen. — Seven segments recognizable dorsally. The first tergite is short, with a row of medium setae on hind margin ; on the sides a group of short setae, increasing in length posteriorly. Tergites II to VI with a dense row of setae on the hind margin and another row, parallel to the former but not reaching the sides, consisting of fewer and shorter setae. Besides the posterior row, the second tergite has several discal setae. The terminal tergite is wider than long, truncate conical, the posterior two-thirds with nu- merous setae in uneven rows, the lateral margins with numerous setae, three or four long setae on the hind margin. Basal sternite shorter than in the female, with 59 to 62 spines. Second sternite with a row of setae on the posterior margin, and three uneven rows of small discals. Third sternite with one row of setae on the posterior margin plus one or two irregular rows of small setae. Fourth sternite with two rows of discal setae, longer than those on the third plus one row of setae on the posterior margin and a group of 27 or 28 spiniforms, arranged into two rows; distal sternite with a dense group of thick setae arranged into several irregular lateral rows. Length, 2.26 mm. Basilia anceps Guimaraes and D'Andretta. Figure 35B, C. Basilia anceps Guimaraes & D'Andretta, 1956, Arq. Zool. Est. Sao Paulo, 10: 113- 116, figs. 188-191. Previous records: Colombia, from Myotis n. nigricans; Peru, from Myotis n. nigricans. PANAMANIAN MATERIAL EXAMINED : From Myotis nigricans, Los Santos Province, collected by V. J. Tipton : 2 males and 1 female on 21 February 1962 (host no. 9962), and 1 male and 1 female on 24 February, 1962 (host no. 10003). From Myotis simus, Armila (San Bias), 2 females, 30 March 1963, collected by C. O. Handley and F. M. Greenwell (host no. 12049). REMARKS : This species and juquiensis Guimaraes, 1946, form a sharply differentiated group, being very similar to each other and differing from the rest of the genus in having only five sternites. They can be easily dis- tinguished by the length of the pustulate setae of the lateral connexivum. In juquiensis these are long and of even length; in anceps they are pre- dominantly long in front of spiracle IV, being very short from there back- wards, except for three or four on the posterior region. GUIMARAES : NYCTERIBIID BATFLIES 401 Tergal plate II also differs in the two species. In juquiensis it has al- most straight lateral and posterior margins, which are rounded and con- tinuous in anceps. The two San Bias females (from Myotis simus) have no setae on the anterior region of the tergite. To date, only the female has been known ; we here describe the male. DESCRIPTION, MALE: Head. — Vertex with three pairs of setae, one between the eyes and two near the front margin ; margin of each gena with seven or eight setae ; postgena with six or seven scattered setae on each side; palpi with eight uneven setae ventrally and one apical seta. Eyes two-faceted (one specimen with single facets on both eyes). Thorax. — Little broader than long (0.70 x 0.62 mm.). Thoracic ctenidium with 17 or 18 spines, notopleural sutures with six or seven setae. Legs relatively short and somewhat laterally compressed ; tibiae slightly dilated distally and with three transverse rows of setae ventrally. Fig. 37. Basilia dunni, male. A, terminal sternite. B, last two tergites. Abdomen. — Basal tergite wider than long, with a few short setae on the posterior margin, plus some discal setae. Tergites II to VI with setae only on the hind margin. Distal tergite with 19 or 20 discal setae on the hind margin, some setae on the sides and four or five uneven setae on each side of the posterior margin. Basal sternite with 50 to 52 spines on the ctenidium of the posterior margin; sternites II to IV with some discal setae besides the row of setae on the posterior border; in addition, the fourth sternite has eight spiniforms on the median region of the hind border. Distal sternite with few setae on the sides. Length, 180 mm. Distribution and Host-Parasite Relationships The present collection is comprised of 84 specimens from 27 lots. This raises the total of reported Panamanian specimens (Curran, 1935; Gui- maraes and D'Andretta, 1956) to 112 specimens, belonging to seven species of nycteribiids from 34 lots. 402 ECTOPARASITES OF PANAMA Panamanian nycteribiids seem to belong to the South American assem- blage of Basilia species. Four of the seven have been found in South Amer- ica more than once: myotis, 23 records; anceps, two; wenzeli, four and ferruginea, seven. The remaining three species, although thus far known only from Panama, have clear relationships with South American species. Basilia dunni is very close to carteri, the differences being rather subtle; handleyi n. sp. is related to ferruginea; tiptoni n. sp. is closer to silvae, so far known only from Chile, although in some ways it resembles boardmanni Rozeboom, 1934, from the United States east of the Mississippi, and rondanii Guimaraes and D'Andretta, 1956, known from Guatemala to Texas. On the subject of host-parasite relationships, the present data fully bear out our previous conclusions as to the close association of New World Basilia with vespertilionid bats. Among 32 host records, 26 refer to vesper- tilionids and only six to phyllostomids. Four of the species which occur in Panama (myotis, dunni, ferruginea and handleyi) have been found exclusively on vespertilionids ; two (anceps and wenzeli) on both vespertilionids and phyllostomids, and one (tiptoni) only on a phyllostomid. Basilia myotis, although recorded from Dasypterus (once) , Uroderma (once) and Molossus (twice) is predominantly a parasite of Myotis nigri- cans (24 records). On the southern part of the range of the host, myotis is replaced by speiseri. Basilia dunni has been found twice on Myotis nigricans and twice on M. albescens. For the closely related carteri there are five records from nigricans and three from albescens. Basilia anceps also seems to be associated with Myotis nigricans, al- though not as closely as B. myotis. Of seven records, four are from M. nigricans, one from M. simus and two from Artibeus lituratus palmarum. Basilia wenzeli is beyond doubt linked to Eptesicus. There are three records from Eptesicus fuscus, five from E. brasiliensis propinquus, one each from Histiotus, Lonchorhina and Artibeus. Three species of Basilia have been found on vespertilionid bats of the genus Lasiurus. Basilia speiseri was found once on Lasiurus borealis bonariensis, possibly as a straggler. However, B. ferruginea has been re- corded five times from this host subspecies, once from Lasiurus borealis frantzii and once from Lasiurus pfeifferi (from Cuba) . Basilia handleyi n. sp. is closely related to ferruginea but distinct. Its occurence on Lasiurus castaneus (whose distribution overlaps that of L. borealis frantzii in Panama) seems to confirm the status of castaneus, re- cently described by Handley (1960), as a good species. Basilia tiptoni n. sp. is the only Panamanian species of Nycteribiidae thus far recorded exclusively from Phyllostomidae. References BEQUAERT, J. 1942. The Diptera Pupipara of Venezuela. Bol. Ent. Venez., 1, (4), pp. 79-88. BRETHES, J. M. F. 1913. Une nouvelle espece de Diptere pupipare du Chile. Bol. Mus. Nac. Chile, 5, (2), pp. 297-299, 1 fig. CURRAN, C. H. 1935. New species of Nycteribiidae and Streblidae (Diptera). Amer. Mus. Nov., no. 765, pp. 1-11, figs. 1-12. DEL PONTE, E. 1944. Basilia romanai, nobis (Diptera, Nycteribiidae) de la Argentina. Ann. Inst. Med. Reg., Tucuman, 1, (1), pp. 117-128, 3 figs. FERRIS, G. F. 1924. The New World Nycteribiidae (Diptera Pupipara). Ent. News, 35, (6), pp. 191-199, text fig. 1, pi. III. GUIMARAES, L. R. 1946. Revisao das especies sul americanas do genero Basilia (Diptera-Nycteribiidae) . Arq. Zool. Est. S. Paulo, 5: 1-87, figs. 1-98, 1 map. GUIMARAES, L. R., AND D'ANDRETTA, M. A. V. 1956. Sinopse dos Nycteribiidae (Diptera) do Novo Mundo. Arq. Zool. Est. S. Paulo, 9: 1-184, 232 figs., 3 tables, 5 maps. HANDLEY, JR., C. 0. 1960. Descriptions of new bats from Panama. Proc. U. S. Nat. Mus., 112: 459- 479, 4 tables. HASE, A. 1931. tiber die Lebensgewohnheiten einer Fledermausfliegen in Venezuela: Basilia bellardii (Fam. Nycteribiidae-Diptera pupipara). Zeitschr. f. Parasitenk., 3, (2), pp. 220-257, figs. 1-17. KARAMAN, Z. 1948. Beitrag zur Kenntnis der Nycteribien IV. "Rad" Acad. Yougosl., 273: 37-46, 6 figs. MIRANDA RIBEIRO, A. 1903. Basilia ferruginea genero novo e especie nova da Familia das Nycteribias. Arch. Mus. Nac., Rio de Janeiro, 12: 175-179, pi. I. 1907. Alguns Dipteros Interessantes. Ibidem, 14: 231-239, pis. XXIII-XXV. 403 404 ECTOPARASITES OF PANAMA SCHUURMANS STEKHOVEN JR., J. H. 1931. Eine seltene, ungeniigend beschriebene Basilia art (Diptera Pupipara) aus Venezuela. Zeitschr. Parasitenk., 3, (2), pp. 205-219, figs. 1-6. 1951. Algunas especies del genero "Basilia" Ribeiro y creacion del nuevo genero "Guimaraesia." Acta Zool. Lilloana, 12: 101-115, figs. 1-4. SCOTT, H. 1936. Descriptions and records of Nycteribiidae (Diptera Pupipara), with a dis- cussion of the genus Basilia. Linn. Soc. Jour., Zool., 39, (267), pp. 479-505, figs. 1-11. STILES, C. W., AND NOLAN, M. 0. 1931. Key catalogue of parasites reported for Chiroptera (bats) with their possible public health importance. Bull. Nat. Inst. Health, no. 155, pp. 603-742. The Streblid Batflies of Panama (Diptera Calypterae: Streblidae) RUPERT L. WENZEL,1 VERNON J. TiPTON,2 AND ALIC JA KIEWLICZ 3 The Streblidae, like the Nycteribiidae, are blood-sucking pupiparous flies and are obligate parasites of bats. All are external parasites except the females of the Old World genus Ascodipteron, which embed themselves in the skin of the host and become endoparasitic. Like their hosts, the Streblidae are primarily tropical in distribution. Relatively few species oc- cur in the subtropical and warm temperate zones. Their distribution and host-parasite relationships suggest an ancient origin and dispersal. The Old and New World faunae have no streblid taxa in common, even on bats of the families Molossidae, Emballonuridae, and Vespertilionidae which are represented in both hemispheres. Of the streblid subfamilies, the Nycteri- boscinae (= Brachytarsininae, with three genera, and about 45 species) and the Ascodipterinae (one genus and 17 species), are restricted to the Old World; the Nycterophiliinae (one genus, four species), Trichobiinae (18 genera, about 69 species) , and the Streblinae (four genera, 21 species) occur only in the New World. The greatest diversification of genera and species has occurred on families of bats that are endemic to the New World, espe- cially the Phyllostomidae. The radiation of the New World Streblidae is correlated not only with taxonomic and biological diversity of the hosts, but also with adaptations of the flies to living and feeding on particular body regions of their hosts. Some bats like Phyllostomus hastatiis normally harbor three or four species of Streblidae at the same time, and most hosts harbor at least two. The co- IT!I Division of Insects, Chicago Natural History Museum. For purposes of subsequent citation, authorship of new taxa described herein should be attributed to the senior author alone, unless otherwise indicated. This study was assisted by Grant DA-MD-49-193-63- G73, United States Army Research and Development Command. 2 Lieutenant Colonel, Medical Service Corps, United States Army. 405 406 ECTOPARASITES OF PANAMA existence of these parasites seems to be possible because of their feeding specializations. The adaptive radiation of the Streblidae is reflected in a wide range of size, form, and structure. The minute mite- or louse-like species of Mastop- tera may be as small as 0.74 mm. long, while gravid females of Joblingia schmidti Dybas and Wenzel may attain a body length of 5.55 mm. The lat- ter is the largest species of streblid known. Reduced wings have evolved in- dependently a number of times in such apparently unrelated genera as Megistopoda, Aspidoptera, Mastoptera, Joblingia, and Metelasmus. Wings are absent in Paradyschiria. The legs of the louse-like species are usually short and powerful for clinging to the host. In Megistopoda and related genera, the venter forms a shield whose anterior margin is upwardly re- flexed like the tip of a ski. In these flies, the hind legs are greatly elongated and apparently assist the fly in moving through the fur of the host by push- ing. The Streblinae, which can appress themselves against the wing mem- branes of the host are compressed dorso-ventrally and have very short f ore- and midlegs. The species of Nycterophilia, which resemble fleas in form and movement, are very strongly, laterally compressed. A postgenal cte- nidium has apparently evolved independently in the Streblinae and in Eldunnia. Many of the relationships between genera are obscured by convergences in adaptive modifications like those mentioned. Comprehensive morphologi- cal and biological studies are needed to clarify both the intra- and inter- familial relationships. We agree with Hennig (1941) that the Streblidae and Nycteribiidae are closely related families of calypterate Diptera, and that neither are closely related to the Hippoboscidae. Cooper's (1941, p. 126) notes on the chromosomes and karyotypes of these families supports this view. Little more can be said at this time about their relationships with other Diptera. The principal earlier papers dealing with the New World Streblidae are those of Speiser (1900) , da Costa Lima (1921) , and Kessel (1925) . These are now of historical and nomenclatural interest only. From 1936 to 1949, Jobling (see references) published a series of eight important papers deal- ing with the taxonomy of New World species. He also published several papers (1929, 1949b, and 1951) that dealt rather extensively with the mor- phology of both Old and New World species. In these superbly illustrated studies, he described only three new species and one new genus, but he re- described 21 species, re-defined eight genera and revised three. During the same period, important contributions were also made by Guimaraes, Pessoa and Guimaraes, and Hoffmann (see references). We consider about 42 species of the previously described New World Streblidae to be valid. Seventeen of these have been reported from Panama by Kessel (1924, 1925), Curran (1934, 1935), Jobling (1936, 1938, 1939, 1949a), Bequaert (1940), and Cooper (1941). Except for the material re- ported upon by Cooper, most of the specimens treated by them were obtained by Major Lawrence H. Dunn, Medical Corps, United States Army (see In- troduction by Fairchild). In the present paper, we treat 84 species, 50 of them new. Sixty-six (44 new) are recorded from Panama. In the collec- WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 407 tions of Chicago Natural History Museum, there are more than 30 additional undescribed neotropical species. An explanation of the reasons ,for this relatively great increase in number of species is warranted. First, the collections upon which earlier authors based their studies were small and included mostly common species from abundant and widely dis- tributed hosts. The present study is based on about 12,000 specimens col- lected from more than 50 of the 100 species of host bats collected in Panama. We have also studied about 10,000 additional neotropical Streblidae in the collections of Chicago Natural History Museum. Obviously, one may expect to discover a number of completely new species in collections of this mag- nitude. This is an important factor in explaining the number of new species recognized by us, but it is only one. Second, the approach to streblid systematics was very conservative in the past. The paucity of specimens and the inadequacy of the data largely prevented one from using host data in parasite discrimination. There was also a reluctance to exploit certain taxonomic characters such as those of the male genital apparatus. The resulting taxonomic "picture" was that of rela- tively few species, quite variable morphologically and occurring on a variety of host species. Third, in past collecting, little care was taken to safeguard against con- tamination or errors of association. It was not uncommon to place several host species in the same field bag, or to collect specimens from a skinning table and to arbitrarily assign the number or name of the host which the col- lector thought most probable. Many of the specimens reported in the litera- ture were removed from preserved specimens in museum collections, or from the bottoms of jars which, at one time or another, contained several species of host bats. Finally, the hosts were frequently misidentified. While this was often due to hasty and uneducated guesses in the field, it was also due to the primitive state of bat taxonomy in the nineteenth and early twentieth cen- turies. Even today, a sophisticated specialist may be unable to adequately characterize and/or identify many neotropical bat species and so-called races. In this study, we were most fortunate in not being handicapped, as were earlier authors, by the inadequacy of collections and host data. The col- lections made in the Panama survey, consisting of several thousand lots, were taken from individual bats representing more than 50 host species. Errors of association, though they occurred, were kept to a minimum (see "Techniques and Methods," below.). The host bats were carefully and uniformly identified by one person, Dr. C. O. Handley, Jr., as he pursued his revisional studies on the mammals of Panama. Any changes in host identification have been made in the streblid data, too, insofar as possible. The accurate host identifications and associations were essential for parasite discrimination (for a discussion of parasite discrimination, see Mayr, Linsley and Usinger, 1953, p. 113). Consequently, we were able to examine and carefully compare large series of parasites from the same host as well as from different host species. This enabled us to re-evaluate taxonomic characters used in the past and to 408 ECTOPARASITES OF PANAMA exploit new ones such as the male gonapophyses. It was soon evident that series of closely related species had been confused under such names as Trichobius dugesii and Euctenodes mirabilis, and further, that many of them exhibit a remarkably high degree of host specificity. In our citations, we have omitted many of the earlier references. As noted above, the host records published in many earlier papers are unrelia- ble. Further, transfers of Streblidae to other hosts in the same roosts are fairly common, though apparently mostly transitory. In addition, a species may be replaced by another species on the same host in another geographic area (and possibly habitat) . This is common but not well understood. For these reasons and because of the extensive taxonomic and nomenclatural changes made in our study, it is not possible to verify or correct most of the published records without examining the specimens upon which they were based. Acknowledgments About 10,000 of the Panamanian Streblidae studied were collected by personnel of the Environmental Health Branch, Office of the Chief Surgeon, United States Army, Caribbean, including Charles M. Keenan, Lt. Col. (then Major) Vernon J. Tipton, and Lt. Col. (then Major) Robert M. Altman. They were assisted in the field by Vicente Alvarez, Wilbur Lowe, Edmund Mattaden, Pantaleon Sanchez, and Sulpice Modestin, and in the laboratory by Wilbur Lowe. Many other specimens were collected by : Dr. Pedro Galindo and Eustorgio Mendez of the Gorgas Memorial Laboratory; Dr. Charles 0. Handley, Jr. and Francis M. Greenwell of the Smithsonian Institution, Washington, D. C. ; Dr. Nathan Gale, Canal Zone Veterinarian ; and Dr. Conrad E. Yunker of the Middle America Research Unit. A few specimens collected earlier by Col. F. S. Blanton, while he was stationed in the Zone, were also examined. To all these persons, as well as to numer- ous unnamed individuals who assisted in the field, we express our apprecia- tion. The status of much of this material could not have been worked out had it not been for extensive ancillary material collected on various expeditions by former and present staff members, associates, and field collectors of Chi- cago Natural History Museum. Among these are : H. A. Beatty (Surinam) ; Philip Hershkovitz (Surinam and Colombia) ; Celestino Kalinowski (Peru) ; Rodger D. Mitchell and Luis de la Torre (Guatemala) ; the late Colin C. Sanborn (Trinidad and Peru) ; the late Franklin J. Schmidt (Guatemala) ; and the late Karl P. Schmidt (Guatemala and Peru). The enthusiastic co- operation of these people helped us acquire basic collections that made study of the Panamanian material possible. A small collection made in Guatemala by F. J. and K. P. Schmidt in 1936 first interested the senior author in study- ing the Streblidae. Later, in 1948, R. D. Mitchell and L. de la Torre accompa- nied the senior author to Guatemala. Here they collected the largest assem- blage of New World Streblidae that had been obtained up to that time. In addition, a number of other specimens were borrowed from other in- stitutions. For their cooperation in making this material available, we wish to thank: Dr. Thomas H. G. Aitken of the Trinidad Virus Laboratory WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 409 (Rockefeller Foundation) at Port-of -Spain ; Prof. Dr. Max Beier, Natur- historische Museum, Vienna, Austria ; Drs. Joseph Bequaert and Philip J. Darlington of the Museum of Comparative Zoology, at Harvard University ; Dr. Elli Franz and Dr. Heinz Felten, Senckenberg Naturforschende Institut, Frankfurt a/Main, Germany; Dr. Paul Kurd, California Insect Survey, University of California at Berkeley ; Dr. J. Knox Jones, Museum of Natural History, University of Kansas ; Dr. C. Machado-Allison and Dr. J. Racenis of the Department of Microbiology, Universidad Central de Venezuela, Caracas ; Dr. C. J. Marinkelle, Universidad de los Andes, Bogota, Colombia ; Dr. Harold Oldroyd, British Museum (Natural History), London; Dr. L. L. Pechuman, Cornell University, Ithaca, N. Y. ; Dr. Alan Stone, United States National Museum, Washington, D. C. ; Prof. Dr. Herbert Weidner, Zoolo- gisches Staatsinstitut und Zoologisches Museum, Hamburg, Germany ; Drs. Paul Arnaud and P. Wygodzinsky, American Museum of Natural History, New York. We were also assisted by a number of persons who supplied us with in- formation regarding collections in their care. For this we wish to thank Dr. Walter Forster, Zoologische Sammlung des Bayerischen Staates, Mu- nich, Germany; Dr. H. D. Folkhard, Naturhistorische Museum, Bern, Switzerland; and Dr. H. Schumann, Institut fur Spezielle Zoologie und Zoologisches Museum, Humboldt Universitat, Berlin. A special expression of gratitude is owed : Dr. B. Jobling, now retired, of the Wellcome Institute of Tropical Medicine (London) for supplying us with information regarding material on which he had published and for giving us permission to use a number of his illustrations, of which he fur- nished us copies; Dr. T. C. Maa for supplying unpublished information re- sulting from his study of streblid types and for his opinions on certain tax- onomic and nomenclatural problems; Prof. Oskar Theodor, Hebrew University Medical School, Jerusalem, for advice and assistance; Dr. Lin- dolpho Guimaraes for lending us his published illustrations of Paradys- chiria "dubia" ; Miss Marion Pahl, Staff Artist, Chicago Natural History Museum, for preparing some of the illustrations ; and Lt. Col. Hugh Keegan, MSC, Chief of the Department of Entomology, 406th Medical General Lab- oratory, Tokyo, Japan, for supervising the preparation of a number of beautifully executed drawings by members of his staff. We are also indebted to Dr. Luis de la Torre, College of Pharmacy of the University of Illinois, Dr. Karl Koopman, American Museum of Natural History, and Mr. Philip Hershkovitz, Chicago Natural History Museum, for supplying information regarding relationships of host bats and certain host data ; and Mr. Henry S. Dybas, who collaborated with the senior author in his early work on the Streblidae and who from time to time has given en- couragement. Above all, we are indebted to Dr. C. 0. Handley, Jr. not only for the invaluable material collected by him, but especially for his untiring coopera- tion in supplying and re-checking host identifications and other data, and for providing valuable information on the habits of host bats. Without his cooperation, this paper could not have been written. Unless otherwise specified, all Panamanian specimens were collected by 410 ECTOPARASITES OF PANAMA Charles M. Keenan and Lt. Col. V. J. Tipton, assisted by personnel of the Environmental Health Branch. Where initials of an institution are given in brackets, it indicates that the specimens concerned are from the collections of that institution. Where the initials GML appear following data, but not in brackets, the specimens concerned were collected by personnel of the Gorgas Memorial Laboratory. The initials MA preceding a host number in- dicate field numbers of Dr. Carlos Machado-Allison of the Universidad Central de Venezuela. Abbreviations used for collections are as follows : AMNH American Museum of Natural History, New York. BISHOP Bernice P. Bishop Museum, Honolulu, Hawaii. BMNH British Museum (Natural History), London. BVP B. V. Peterson, Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario. CAS California Academy of Sciences, San Francisco. CIS California Insect Survey, Department of Entomology and Parasitology, University of California, Berkeley. CNHM Chicago Natural History Museum, Chicago, Illinois. cu Department of Entomology, Cornell University, Ithaca, New York. DZSP Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil. EHB Environmental Health Branch, Office of the Chief Surgeon, United States Army Forces Southern Command, Fort Amador, Canal Zone. FCUCV Departamento de Microbiologia, Faculdad de Ciencias, Universidad Central de Venezuela, Caracas. GML Gorgas Memorial Laboratory, Panama, Panama. INM Institute Nacional de Microbiologia, Buenos Aires. KU Snow Entomological Museum, University of Kansas, Lawrence. LACM Los Angeles County Museum, Los Angeles, California MCZ Museum of Comparative Zoology at Harvard University, Cambridge, Mas- sachusetts. MNHN Museum Nationale d'Histoire Naturelle, Paris. OT Oskar Theodor, Department of Parasitology, Hebrew University, Jerusalem. SNI Senckenberg Naturforschende Institut, Frankfurt a/ Main. TVL Trinidad Regional Virus Laboratory (Rockefeller Foundation), Port-of- Spain. USNM United States National Museum, Washington, D. C. ZMHU Institut fur Spezielle Zoologie und Zoologische Museum, Humboldt Uni- versitat, Berlin. ZMH Zoologisches Staatsinstitut und Zoologische Museum, Hamburg. Techniques and Methods Collecting Hosts. — As one can determine from Dr. Handley's account (see "Checklist of Mammals"), bats roost in a remarkably wide variety of situations. Accordingly, collecting techniques were varied and often novel. Bats that could not be obtained in any other way were brought down with dust shot. This was often necessary where they were roosting high in a hol- low tree or cave. When possible, shooting should be avoided, because of un- due disturbance of the parasites. A variety of nets and net-collecting techniques were employed. Where bats roosted in culverts, it was common procedure to place a loose mesh net (fish seine) over one end and to send a man through the culvert from the other end. He literally pushed the bats ahead of him with a leafy branch. As the bats "hit" the net at the other end, they were quickly removed by WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 411 men wearing gloves. One person stood alert with a hand insect net, to cap- ture bats that escaped through side openings. If the culvert was impassible, firecrackers were thrown in the open end. Bats in tree holes were obtained by essentially the same technique, the net being strung over the opening, and a leafy branch or bough of appropriate size introduced through one side and pushed up into the cavity to dislodge the bats. Hand insect nets (with coarse mesh bags) are often useful in caves, especially near the openings. They may also be used to collect bats, such as Saccopteryx, that roost on tree trunks or under branches. The roosting places of many bats are inaccessible or unknown. The use of "mist" nets makes it possible to obtain many of these species and their parasites. Immediate removal of the bat and its parasites lessens the loss or incidence of transfer. Dr. Handley found it desirable to remove some of the very active streblids from the bats while they were still enmeshed in the net. Collecting Parasites. — Most of the host bats collected in the Panama sur- vey were, upon capture, immediately placed in individual paper or plastic bags, together with a field number, and etherized. The flies were removed as quickly as possible and placed in vials with the corresponding field num- bers. This is important, because they do not preserve as well, if transferred an hour or two after death. Paper bags were discarded after one use. Plastic bags were thoroughly washed and reversed, before being re-used. In collecting the host, it is essential that it be bagged as quickly and with as little unnecessary handling as possible. Some Streblidae, like Speiseria ambigua and Trichobius uniformis not only leave the host at the slightest disturbance but will often land on another nearby bat, frequently of another species. Such "disturbance transfers" are not uncommon, among Streblidae of bats caught in nets. These flies often return to the original host, if time and opportunity permits. Others like Strebla, though strong fliers, tend to stay with their host, even when disturbed. While the need for the most rigorous precautions in collecting ectopara- sites is well known, in the past they have rarely been observed in the case of batflies. Unfortunately, the same has often been true for many other ectoparasites. Our experience leads us to believe that a substantial number of the host records published for them are valueless. Preparation and Study. — When possible, identifications should be made using both slide preparations and specimens preserved in 70-75 percent ethyl alcohol. In preparing Streblidae for slide mounts, the following procedures were followed. With minuten nadeln, small punctures were made in the abdomen and other membranous parts of the body wall. The flies were then placed in a 10 percent solution of cold or warm KOH for from 5 to 20 hours, washed in distilled water, passed through a series of graded alcohols, includ- ing absolute alcohol, cleared in oil of cloves, and mounted in a good grade Canada balsam. The male terminalia were removed and mounted separately. A slit was made in either the dorsal or ventral wall of the abdomen while the specimen was in oil of cloves. The terminalia were removed through the slit by means 412 ECTOPARASITES OF PANAMA oc.l. oc.l. C D Fig. 38. Structures of streblid head. A-C, Trichobius sphaeronotus Jobling. A, dorsal, B, ventral, and C, lateral views of head. D, Trichobius sp. (tlongipes Rudow), dorsal view, semi-diagrammatic, setae omitted. A-C from Zeve and Howell (1962) with modified terminology. D adapted from Jobling (1929). See text for explanation. of very fine jewelers' forceps. In this way, they could usually be removed in toto with much less difficulty than by dissecting them out through the genital "pouch." In order to insure reasonably uniform interpretation of the characters of the gonapophyses, the terminalia were mounted on their right side. Illustrations and descriptions were based on a lateral view. However, in some Streblidae, the gonapophyses are frequently twisted to the left and sometimes only feebly, if at all, curved ventrally. This is true in certain species of Trichobius, especially of the dugesii and longipes WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 413 groups. When this is so, rotation of the terminalia, as by pressure of the cover slip, may orient the gonapophyses so that they appear to be curved dorso-ventrally when they are actually curved to the left. One must take Fig. 39. Structures of streblid head. A, B, Metelasmus pseudopterus Coquillet, dorsal and ventral views of head, respectively (post-ctenidial area not shown in B) . C, Eldunnia breviceps Curran, ventral view of posterior margin. D, Anastrebla sp., dorsal view, palpi and frontoclypeal area omitted. A and B from Jobling (1936) ; C after Jobling 1929. See text for explanation. this into account in mounting and interpreting male terminalia. We believe that we largely avoided such rotation in most of the preparations from which we made illustrations. It is desirable to remove the wings and mount them separately. This should be done with fine jewelers' forceps or dissecting needles before the fly is placed in KOH. Wings should not be subjected to KOH, but only passed through the graded alcohols and oil of cloves before mounting1. We strongly recommend the use of a high grade Canada balsam in pref- erence to such media as Hoyer's or polyvinyl alcohols. Pieces of glass cover- 414 ECTOPARASITES OF PANAMA slips, slides, or other supports such as glass or plastic rings of appropriate thickness, must be used to prevent the coverslip from crushing the speci- mens. We also use small squares, of various thicknesses, of good quality heavy paper such as manila cover paper. They are soaked in xylol before being introduced into the medium. For wing and genital preparation, we have found it very convenient to use % inch diameter coverslips. tr. t.3 hit: A ex. 3 Fig. 40. Trichobius sphaeronotus Jobling. A, dorsal and B, ventral views (semi-dia- grammatic). Adapted from Zeve and Howell (1962). See text for explanation. Measurements. — All measurements given are in millimeters. Wing length (WL) has been measured from the fractureline (fig. 42, f.l.) at the base of the radius to the apex of the wing; width (ww) was measured at the broadest portion. The base of the radius was selected as a measuring point, because it was rarely damaged or removed in dissecting off the wings for slide preparations, while axillary sclerites were frequently damaged or lost. Body length (BL) has been measured from anterior margin of fronto- clypeus to apex of abdomen, thoracic length (TL) from anterior margin of prescutum to posterior tip of metanotum. FL = length of femur. Illustrations. — The photographic illustrations were prepared as follows : paper negatives were made by projecting slide preparations onto enlarg- ing paper; positive prints were made on contact print paper, as with film negatives, emulsion to emulsion, the light exposure being made through the paper negative. Although this technique has many limitations, it saves much time in preparing illustrations of material for which it is suited. Other illustrations made by us were prepared from slide preparations with the aid of a microprojector. WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 415 Figures 76-78 (A, B), 97-99, and 122-124 were executed by artists of the 406th Medical General Laboratory in Tokyo. Illustrations borrowed from Mr. Jobling are credited to him in the legends. Most of the remaining draw- ings were prepared by the senior and junior authors, in collaboration. metn. cx.l stpl. Fig. 41. Thorax, lateral view (diagrammatic), Trichobius sp. Adapted from Zeve and Howell (1963) . See text for explanation. MORPHOLOGY AND SYSTEMATICS Morphology The comparative morphology of the Streblidae has been treated in sev- eral papers by Jobling (1929, 1936, 1938, 1958). Zeve and Howell (1962, 1963) treated the morphology of three North American species of Trichobius of the major group. These authors differ with Jobling in the interpretation of certain structures. It is obvious that further detailed studies covering a broad taxonomic representation of the family must be undertaken if one is to resolve these differences and to arrive at a better understanding of other structures not investigated in detail by either. We were not in a position to undertake such studies. However, we feel it is desirable to discuss some of these differences in interpretation and to explain the terminology used by us. Unless otherwise noted, the following discussion concerns only New World Streblidae. Head. — The head of New World Streblidae has been described by Jobling (1929,1936) and by Zeve and Howell (1962). The terminology employed for head structures in this paper is shown in figs. 38, 39. In the caecus, pallidus, and major groups of Trichobius, the subdivisions of the vertex are not clearly differentiated (fig. 38 A, C) into subregions. In these, there is a median, 416 ECTOPARASITES OF PANAMA membranous or less distinctly sclerotized mediovertex (mv.) ; the anterior subregions or laterovertices (lv.) and the posterior subregions (occipital lobes, ocJ.) are usually separately rounded or separated by a vague trans- verse groove; laterally they may be partially separated by a membranous strip (fig. 38C). In most other Trichobiinae and in the Nycterophiliinae, the laterovertices and occipital lobes are well differentiated (fig. 38D) ; espe- cially in those species in which the head is flattened dorsally.4 In these, the occipital lobes may be more or less flattened plates. Zeve and Howell (loc. cit.) appear to interpret each side of the entire dorsum posterior to the Sc+R H 2+3 f.l. (M3+4+Cu,) 1A (2A) \ (Cu0+2A) M4+Cu, (Cu,) Fig. 42. Wing, Trichobius sparsus Kessel. Adapted from Jobling (1938). See text for explanation. antennae as the laterovertices. Until a detailed comparative study can be made of the head, we prefer to use the terms indicated above. The postero- median area, called the postvertex (pv.) may be membranous (fig. 38D) or form a sclerite (fig. 39A, D). In the Streblinae, the postgenae extend posteriorly on the dorsal surface, as conspicuous plates (fig. 39 A, pg.) in Strebla and Metelasmus, or as a sclerotized strip in Anastrebla (fig. 39D). In the Streblinae the occipital plates have a festooned posterior margin, as does the postvertex, and, a conspicuous postgenal ctenidium is present on the underside (fig. 39B). This extends around to the dorsal surface (fig. 39A, D). A short ventral ctenidium is also present in Eldunnia (fig. 39C). Thorax. — The principal structures referred to in our descriptions are illustrated and labeled in figs. 40, 41. A few remarks are in order. The * Pessoa and Guimaraes (1940, figs. 2, 3) figured the head of Nycterophilia. Their illustrations do not show the laterovertices as distinct from the narrow occipital strips, as they appear in stained material. On the basis of their interpretation, they assigned the genus to the Nycteroboscinae (Brachytarsininae). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 417 mesonotum (fig. 40A) consists of mesoprescutum (prsc.), mesoscutum (sc.) and mesoscutellum (scl.) . In our descriptions, we have not qualified these with the prefix but have referred to them simply as prescutum, scutum, and scutellum. Zeve and Howell (1963) have referred to the upper portion of the mesepisternum and mesepimeron as the mesoanepisternum and meso- anepimeron, respectively. Since there is no suture separating the meso- anepisternum from the mesokatepisternum, etc., we prefer to simply use the term mesepisternum (fig. 41, eps. 2) and mesepimeron (epm. 2) for taxo- nomic purposes. Jobling (various papers) has referred to the mesepister- num as the mesopleuron. Since this includes the mesepimeron as well, we use the more restrictive term. Zeve and Howell (loc. cit.) have referred to the two major divisions of the venter of the thorax (fig. 40B) as meso- and metasternum. We have employed the terms sternopleura (stpl.) and pleurotrochantines (pltr.) pri- marily because of past usage by Jobling. We are not able to judge which terms more truly reflect the composition of these sclerites. The median posterior margin of the pleurotrochantines may be produced as a lobe, called by us the median pleurotrochantinal lobe (m.p.L). In some species, for ex- ample in the longipes group of Trichobius, this may ascend dorsally and unite with the metepimeron (fig. 62 A). Wings. — Jobling (1936 and 1938) treats the wing venation of the Streb- lidae. Zeve and Howell (1963) agree with him in their interpretation of the anterior veins of the wing, but differ as to the posterior ones. In fig. 42, we have shown the terminology of Jobling (1938). Where it differs, we have shown the terminology of Zeve and Howell in parentheses. In our de- scriptions, we have tried to avoid confusion by using names or symbols for the costa, RI, and Rs and referring to the other veins by number. In those species of Nycterophilia that have fully developed wings (fig. 52), only a few anterior veins are well developed and sclerotized; remnants of some vannal (?) veins are also sclerotized. Nine others seem to be represented by rows of setae. If these rows of setae actually represent veins, then a more complete complement is present in the Nycterophiliinae than in any other Streblidae known to date. The anterior venation somewhat resembles that of male Ascodipteron, as figured by Jobling (1940) , though there are obvious differences. In brachypterous and stenopterous wings (sensu Hackman, 1964) venation may be essentially complete and identifiable (Aspidoptera) or very much reduced (Noctiliostrebla, Neotrichobius) . In some unde- scribed species of Nycterophilia the wings are micropterous. Abdomen. — There are seven pairs of spiracles in all of the known New World Streblidae. Terga I and II are completely fused in the Trichobiinae and Streblinae. They are separated ventrally but fused dorsally in the Nycterophiliinae. Sternum I is much smaller than II in the Trichobiinae and Streblinae, but longer than II in the Nycterophiliinae. In the following discussion, we refer to the abdominal membrane as the connexivum. Male: The structures of the male abdomen (fig. 44B) are labeled accord- ing to the interpretations of Jobling (1951) . The fifth sternum (st.V) may 418 ECTOPARASITES OF PANAMA t.I+II st. VII sup. a. p. sub. a. p. sub. a. p. v. a. st. I B st. II st.VII Fig. 43. Female abdominal structures, Trichobius sphaeronotus Jobling. A, dorsal, and B, lateral views. Adapted from Zeve and Howell (1963). See text for explanation. WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 419 be entirely absent, it may be a single transverse sclerite (in most Streb- lidae), or it may be divided into two sclerites as in Anatrichobius and some species of the Trichobius major group. Sternum VI (st. VI) may be pres- ent, or absent, in related species (Trichobius) or well developed in all of the species of a genus (Strebla). Jobling (loc. cit.) and Zeve and Howell (loc. cit.) differ greatly in their interpretation of the composition of the male hypopygium (hyp.). Zeve and Howell, contending that the spiracles of segment VII could not be en- closed by a sternum,5 maintain that the dorsal synsternum VII+VIII of Jobling is a syntergum (VII+VIII). To account for the position of these spiracles, Jobling suggests that part of tergum VII may enter into the forma- tion of the hypopygium. We are in no position to judge the merits or de- merits of these opposing points of view. Neither appears to be compatible with the facts.6 However, we believe Jobling has demonstrated that there has been rotation within the postabdomen, a fact not consistent with the in- terpretation of Zeve and Howell. For this reason, we have followed Job- ling's terminology, though with some reservations. Synsternum VII+VIII is quite variable in its structure. It may be di- vided into two clearly separated lateral plates, which lack any dorsal con- nection (e.g., Trichobius major) , or it may be continuous dorsally (Noctilio- strebla ?) or at least apically (Strebla). The suture between VII and VIII may be visible (Strebla) or not. Within the same genus (Trichobius), VII+VIII may be fused to tergum IX or clearly separated by a membranous suture. In the genus Nycterophilia, the abdomen differs markedly from that of other New World Streblidae. In the male, segments I-VI are represented by clearly defined, sclerotized sterna. The hypopygium consists of a conspic- uous synsternum (VII+VIII) whose apex is closed dorsally to form a ring; the apex articulates with a dorso-apical sclerite, tergum IX, which bears the terminalia and the anus. The cerci appear to be represented by very small setose flaps on each side of the anus. Excepting segment IX, the hypo- pygium is sclerotized ventrally for its entire length. Male terminalia. — The morphology of the male terminalia (fig. 45B) of the Streblidae has been discussed by Jobling (loc. cit.), Zeve and Howell (loc. cit.) Theodor (1954), and Theodor and Moscona (1954). The internal clasper-like structures were referred to as claspers by Jobling (1936) and later (1951) as gonapophyses. Zeve and Howell (1963) also use the term gonapophyses. Theodor (1954) has referred to them as parameres. In 5 The position of the spiracles does not in itself seem to us to be a valid criterion. In Nycterophilia, the first spiracle is so close to the upper margin of sternum I that it sometimes seems to be incorporated in the margin, and the second lies just above the posterior angle of sternum II. In all other New World Streblidae, they are clearly situated on tergum I+II. 9 Comparison of the structure of the New World Streblinae and Trichobiinae with that of the Old World genus Brachytarsina (= Nycteribosca) suggests that part of tergum VI may be jncjnded in the anterior part of the hypopygium. 420 ECTOPARASITES OF PANAMA fig. 45B we have not shown the U-shaped sclerites illustrated by Jobling and Zeve and Howell. In the Trichobiinae and Streblinae, each gonapophysis (gon.) typically bears a pair of distinctive ventral or ventro-lateral setae. These paired gonapophyseal setae (p.g.s.) consist typically of a macroseta (ms.) and a smaller accessory seta (a.s.). Other setae may also be present. The acces- sory seta is posterior to the macroseta in the uniformis, dugesii, and longipes groups of Trichobius, and in Speiseria, Pseudostrebla, Stizostrebla, Aspi- doptera, Mastoptera, Noctiliostrebla, Paradyschiria, and Exastinion. It is anterior to the macroseta in the major, caecus, and phyllostomae groups of Trichobius and in Joblingia, Megistopoda, Neotrichobius, Paratrichobius, Anastrebla, Metelasmus, Paraeuctenodes, and Strebla. In Joblingia schmidti, there are two pairs of macrosetae, and sometimes two pairs of accessory setae. In Trichobioides perspicillatus, the accessory seta is ventral or ventral and slightly posterior to the macroseta. In Eldunnia there are four well- developed setae in a row, the two posterior setae shorter than the two anterior ones. Generally, the paired setae are similar in position on the two gonapo- physes and are inserted along the ventral margin near base. However, they may be situated near mid-length or (as in Paratrichobius and related genera) near apex. In Trichobius phyllostomae and related species, their location may differ markedly between the two gonapophyses. In some (e.g., Trichobioides), the paired setae may be inserted on the lateral face of the gonapophysis. We have not studied the aedeagus (aed.) in detail, though its structure differs between genera and species. In many New World Streblidae, the extrusible portion of the aedeagus is dorso-ventrally flattened and ribbon- like, but it may be cylindrical or even flagelliform. In others (e.g., Noctilio- strebla) , it is blade-like in profile and may even have a dorso-apical spine. In most (except Nycterophilia, Noctiliostrebla, and Paradyschiria), its pos- terior portion is coiled, and extrusion is accomplished through straightening of the coil by contraction of muscles associated with the aedeagal apodeme (aed.a.). In Nycterophilia, the apodeme is short and straight. In Noctilio- strebla and Paradyschiria, it is feebly coiled. In the Trichobiinae and Streblinae, there is a pair of flap-like, usually setose structures in the wall of the genital pockets and closely associated with the dorsal connection of the membrane to the gonapophyseal sheath. These are referred to by Jobling (1951) as the surstyli. In Nycterophilia, unlike any other Streblidae except Ascodipteron, the terminalia are external, and consist of three principal elements (fig. 45A). There is an outer pair of "claspers," each consisting of a long cylindrical clasper shaft (cl. sh.) which is continuous with the body wall of tergum IX and not separated from it by a suture of any kind. Each of these bears a distal, movable harpago ( ha. ) . Internal to the claspers and articulated to the ventral apical margin of synsternum VII + VIII is another pair of broad clasper-like structures, apparently homologous with the gonapophyses of other New World Streblidae and, like them, each bearing a pair of ventral setae. Situated between the gonapophyses is the extrusible, ribbon-like WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 421 an. sp.VII sp.I st. I hyp- aed. gon Fig. 44. A, female terminalia, ventral view, Strebla christinae, new species. B, male abdomen, lateral view, Strebla sp. B adapted from Jobling (1951). See text for ex- planation. 422 ECTOPARASITES OF PANAMA ha. gon. aed. aed. a. p.g.s. gon. B gon. a. aed. Fig. 45. Male terminalia. A, Nycterophilia parnelli, new species, shaped sclerites omitted). See text for explanation. aed. B, Strebla sp. (U- aedeagus. This has a simple, short apodeme which cannot be coiled as it can in most other New World Streblidae. We believe the abdominal morphology shows that the species of Nyctero- philia— in spite of the remarkably specialized flea-like adaptions — are the most generalized New World Streblidae known. Interestingly, the only other Streblidae which, like Nycterophilia, seem to approach a generalized type, are the males of Ascodipteron. In these, the abdominal terga and sterna, though weakly sclerotized, are identifiable. The females of Asco- dipteron, on the other hand, are the most highly specialized, morphologically and biologically, of all the Streblidae. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 423 Abdomen. — Female: Jobling (1951) and Zeve and Howell (1963) agree in their interpretation of segments I-VII in the female abdomen. Although they differ in the terminology they apply to the apical structures, neither seems to have resolved the homologies. Our preliminary studies indicate that characters of the postabdomen are not only of value in identifying species, but may assist in an understanding of relationships within the family. Tergum VII varies greatly in extent, shape, and sclerotization. It is best developed in Strebla. Sometimes it is not at all pigmented but may be repre- sented by micropile and/or paired setae. Occasionally, it is united or even completely fused with the supra-anal plate to form part of the terminal cone (see below). In a very few instances (e.g., Noctiliostrebla, Joblingia, and Trichobius major) tergum VII may be divided into two tergites. Zeve and Howell (loc. cit.) have not discussed the segmental nature of the terminal cone (also called hypopygium and proctiger by Jobling). We believe this composite structure to be formed of elements of segments VIII -X (and, theoretically, XI). In some streblids, even tergum VII is fused with the supra-anal plate of the cone. Characteristically, the most conspic- uous structure of the cone is a sclerotized ring consisting of a dorsal supra- anal plate (sup.a.p.) and a narrow ventral arc (v.a.). The supra-anal plate may be longer than wide (fig. 43 A) or very short and scarcely longer than the ventral arc (fig. 44 A) . The ventral arc may be produced anteriorly as a median lobe (fig. 44 A) or on each side as a narrow strut (figs. 43 A, B) . Between the ventral arc and the anus is a small setose sclerite called the subanal plate (figs. 43B, 44A, sub.a.p.) by Jobling (1951). In Joblingia schmidti, the cerci are united with the ventral arc, which is separate from the supra-anal plate and so situated as to appear to be a sternum of that plate. This is also true in species of the Trichobius major group (fig. 43) and other streblids. The fusion of the cerci to a sternal structure raises a question as to their identity, since cerci are generally re- garded as tergal appendages. Perhaps the streblid cerci are paraprocts. In J. schmidti, there are also two U-shaped anal sclerites in the wall of the proctiger. One of these (supra-anal sclerite) lies above the anus, the other (infra-anal sclerite) lies below it. These are absent in many species and genera. In Strebla christinae n.sp., there is a single supra-anal sclerite (fig. 44 A, sa.s.). We restrict the term proctiger (pr.) to the membranous projection which bears the anus, anal sclerites, the subanal plate, and the cerci. In species of the Trichobius caecus group, and some other streblids, there is a small sclerotized strip that has an apical crescent-shaped thickening or rim which we refer to as the postgenital sclerite. It is situated between the vulva and the ventral arc. We interpret this as sternum IX. It is best de- veloped in the species of the Trichobius caecus group and is useful in identi- fying them. It is most easily observed in alcohol-preserved specimens and should be examined from a posterior view. Its shape cannot ordinarily be determined from slide preparations. In the abdomen of female Nycterophilia, syntergum I +11 and sometimes tergum VII are sclerotized. In non-gravid females, many of the segmental 424 ECTOPARASITES OF PANAMA margins are indicated by external indentations of the connexivum. The terminal cone (fig. 51B t.c.) is apically produced above the proctiger (pr.) and bears conspicuous distal macrosetae. The proctiger is free. The abdominal sterna, though indicated, are incompletely represented and very weakly, if at all, sclerotized. Below the proctiger is a small sclerite that is connected by a sclerotized strip to a small transverse sclerite lying posterior to the vulva. These appear to be comparable to the structures figured by Jobling (1951, fig. 1A, B) for Nycteribosca gigantea. He regarded the sclerotized strip as representing sternum X and the sclerite as sternum IX. The latter sclerite is similar in form and position to the postgenital sclerite (see above) of the Trichobius caecus group. Chaetotaxy. — We have used the term microseta for extremely minute se- tae. In some instances, these may be unusually minute and relatively dense, forming a micropile. They are usually visible only in slide preparations, and some can be distinguished only under oil immersion. The term macroseta has been used for setae that are conspicuously long or longer than other as- sociated setae. In general, the dorsal abdominal setae are well developed and indeed sometimes very long in brachypterous and wingless species, e.g., the female of Joblingia schmidti. In Nycterophilia, certain coarse abdominal setae, especially the connexival setae, may appear "braided" even under relatively low magnification. In some instances, their margins appear serrate. A aed. aed.a. an. ant. ar. a.s. a.sp. Bac. C ca. ce. cl.sh. conn. c.s. ct. Cu ex. 1 ex. 2 ex. 3 d.c.s. e. epm. 2 epm. 3 eps. 1 eps. 2 eps. 3 List of Abbreviations anal vein e.s. aedeagus fc. aedeagal apodeme f.l. anus /M.S. antenna g. arista gon. accessory gonapophyseal seta gon.a. anterior thoracic spiracle gon.s. basicosta H costa ha. calyptera hit. cercus h.s. clasper shaft hyp. abdominal connexivum Ibl. coxal spur l.l. ctenidium l.m.c. cubitus lv. procoxa ' M mesocoxa md.s. metacoxa metn. (paired) dorsal m.p.l. abdominal connexival setae m.scl. eye ms. mesepimeron mv. metepimeron oc. proepisternum oc.l. mesepisternum p. metepisternum p. 1 epi-anal sclerite frontoclypeus fracture line of radius furcasternal suture gena gonapophysis gonapophyseal apodeme gonapophyseal sheath humeral vein harpago halter hypo-anal sclerite hypopygium labella lateral lobe of tergum I+II lateral membranous cleft laterovertex medius median mesonotal suture metanotum median pleurotrochantinal lobe mesoscutellum (scutellum) gonapophyseal macroseta mediovertex occiput occipital lobe palpus (maxillary) pronotum WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 425 pd. pedicel stc.s. pg. postgena stpl. p.g.s. paired gonapophyseal setae sub.a.p. pltr. pleurotrochantine (s) sup.a.p. pr. proctiger t. prsc. mesoprescutum (prescutum) t.3 pv. postvertex R radius i.e. Rg radial sector th. r.m. rostral membrane tr. r.s. remiform scale tr.s. *a&. supra-anal sclerite v.a. sc. mesoscutum (scutum) v.g.m. scl. scutellum sp. spiracle v.m.c. st. sternum w.p. sternacostal suture sternopleuron (a) subanal plate supra-anal plate tergum laterotergite (metapleuron of Jobling) terminal cone theca trochanter transverse mesonotal suture ventral arc ventral gonapophyseal macroseta vertical membranous cleft wing process Biological Notes As indicated above, the Streblidae are obligate pupiparous ectoparasites of bats. There are few published studies or notes on their biology. The only paper of consequence that deals with the habits and life history of New World species is that of Ross (1961) who reported on several species of Trichobius that occur in Arizona. From unpublished field observations of the senior author and of Prof. Kenneth W. Cooper (pers. comm.), it appears that many species occupy rather specific niches on the host, such as the wing membranes, head, or trunk. Many of the modifications of size, form, and structure appear to be adaptations to these niches. Field and laboratory studies are needed to analyze them. The movements of some of these flies on the wing membranes of the bat are remarkable. Using a low power binocular microscope, we observed that Paratrichobius dunni (Curran), Trichobius longipes (Rudow), T. joblingi n. sp., and various species of Strebla run rapidly with equal facility, in for- ward, backward, lateral, and oblique directions. Their skittery movements reminded one of a bit of mercury rolling on a plane surface that was being tilted in various directions. The senior author has tried to keep colonies of Streblidae alive in the laboratory. He succeeded only when the flies were kept on the host. Four- teen Carollia p. azteca were "deloused" after being caged for several days in the laboratory. Only two were negative for parasites ; the remaining 12 had an average of 2.15 specimens of T. joblingi and 0.25 specimens of Strebla carolliae per bat. When removed, the streblids died in one or two hours or less. This agrees with the experience of Ross (op. cit.), working with Arizona Streblidae. At first, it was thought that the flies died of desiccation. However, they died as quickly when kept in ceramic pots with relatively high humidity. The species handled were Strebla carolliae n. sp. and Trichobius joblingi n. sp., on Carollia p. azteca, and Strebla mirabilis ( Waterhouse) , S. hertigi n. sp., and Trichobius longipes (Rudow), on Phyllostomus h. pana- mensis. All were collected in the Canal Zone. 426 ECTOPARASITES OF PANAMA These species rarely leave the host. They appear to feed constantly and have been observed "diving" in and out of the fur as they feed briefly and move from one site to another. Why they feed so often is an interest- ing question. Perhaps it is because rapid conversion and utilization of food are needed to nourish the developing intra-uterine larva. Only one larva matures at a time, and it may be that a short developmental period is required to maintain the population. Unfortunately, nothing is known about the population dynamics of these flies. Nor do we know if the males feed as frequently as the females. On the other hand, it may be that the continual rapid movements of these flies require a high energy intake. The very active species which were seen to feed constantly stay on the host most of the time. We suspect that the host bats preen themselves and that the movements of the flies are evad- ing actions. More generalized species like Joblingia schmidti Dybas and Wenzel and Trichobius yunkeri n. sp. do not stay on the host as much of the time nor do they feed as frequently as do the more specialized streblids. Many specimens of J. schmidti were found crawling in guano and on the walls of a bat cave. T. yunkeri and related species are much less rapid flyers than most Trichobius species and may be found in swarms in the roost- ing places of the hosts. Whatever the need, the suggestion that food is converted rapidly is sup- ported by the observations of Dr. Phyllis Johnson (pers. comm.). While at the Gorgas Memorial Laboratory, she dissected live specimens of Tri- chobius longipes and Strebla mirabilis. She noted that the blood in the gut appeared to have been broken down and absorbed more quickly in these flies than in any other blood-sucking insect she had dissected. This suggests that the short survival time of these flies off the host, might literally be due to exhausting their energy. Dr. Glenn Richards (pers. comm.) has suggested that a need for a high water intake rather than of food might be a more important limiting factor. The problem is an interesting one and warrants study. At best, the fecundity of these flies must be low. The females do not appear to have a well-developed spermatheca for sperm storage. The spe- cies observed by Ross (loc. cit.) mate on numerous occasions. Even if the mortality rate is low and the flies are long-lived, it must require a consider- able period of time for a population to achieve sizable numbers. This may explain why the flies are absent or the population density low in certain bat roosts, especially new or temporary sites. For example, an abandoned power house shelter at Madden Airstrip, built during World War II, had a large colony of Carollia p. azteca and Phyllostomus h. panamensis. No puparia could be found in this shelter nor were any flies seen on the walls or in the air. The bats had few flies. Conversely, population densities of flies like T. yunkeri were often very high on bats, like Pteronotus parnellii fuscus, which roosted in large colonies in caves or similar long established habitats. Crude experiments on "homing" and host selection were also undertaken in the laboratory of the Environmental Health Branch at Corozal, to de- termine the feasibility of future studies along these lines. Specimens of two hosts (Carollia p. azteca and Phyllostomus h. panamensis) were very WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 427 lightly anesthetized with ether. Their flies were removed, and the bats were returned to a screened cage. After a few minutes, the flies recovered from the anesthesia. They were released in the room containing the hosts' cage. Most of them flew quickly and directly to the cage and literally clawed and burrowed their way through the screen to return to the hosts. The parasites of these two species of bats always returned to their normal host. Carefully controlled multiple choice experiments along these lines may pro- vide insights into some of the problems of host selection and specificity. Ross (loc. cit.) observed that the Streblidae which he studied deposited their pupae off the host, sometimes in communal sites in crevices or on the walls and floor of the roosting site. This agrees in general with our obser- vations. We have occasionally found puparia in small numbers in the fur of the bats. We believe this to be abnormal. Ross (loc. cit.) also reported being bitten by Streblidae. We have never observed this, and it may be true only of certain species. Host-parasite specificity is discussed follow- ing the section on systematics. Systematics There are three described genera of New World Streblidae that have not been taken in Panama. These are Synthesiostrebla, Stizostrebla, and Paraeuctenodes. Synthesiostrebla (amorphochili Townsend) is known only from Peru. Tonatia silvicola (=amblyotis) the host of Stizostrebla longi- rostris occurs in Panama, and it is likely that the fly will eventually be collected there, too. We have seen specimens of Paraeuctenodes from Trinidad and from Guatemala. Thus, it seems likely that the genus is repre- sented in Panama. The following key is designed not to reflect relationships of genera, since these are largely obscure, but to facilitate identification. KEY TO THE GENERA OF NEW WORLD STREBLIDAE 1. Body strongly, laterally compressed, flea-like (fig. 46). Wings, if fully devel- oped, with most veins represented only by rows of setae (fig. 52) ; wings some- times reduced to inconspicuous strut-like structures. Male (figs. 49, 50) : Pre- abdomen with sterna I-VI sclerotized and distinct; terminalia (fig. 45 A) exter- nal, aedeagus lying between well-developed gonapophyses, these in turn lying between conspicuous claspers (Nycterophiliinae) Nycterophilia Ferris Body, if laterally compressed, never flea-like. Wings, if fully developed, with six longitudinal veins (fig. 42) ; wings sometimes reduced to oval or elongate struc- tures, or (rarely) completely absent. Male: Pre-abdomen (segments I-VI) with only sterna I and II sclerotized and distinct; I usually very small, some- times reduced to a very small sclerite on each side; terminalia internal, con- sisting of aedeagus and a pair of gonapophyses (fig. 45B) 2 2. Head with a ctenidium (fig. 39 ) 3 Head without a ctenidium (fig. 38) 7 3. Ctenidium consisting of only 18-19 spines, restricted to postero-ventral part of head (fig. 39C) ; palpi "free," with normal setae, not forming a triangular shield for the front of the head Eldunnia Curran Ctenidium consisting of numerous spines, extending around sides to dorso-lateral parts of head (fig. 39A, B, D) ; palpi very broad, festooned with numerous, heavy thorn-like setae, together forming a triangular shield for the front of the head (fig. 39A, B ; Streblinae) 4 428 ECTOPARASITES OF PANAMA 4. Wings brachypterous. Dorsal connexivum of abdomen covered with setae (fig. 140) . Prescutum with a complete median suture. Postgena, behind the ctenidium, with a remiform scale (fig. 39A, r.s.) Metelasmus Coquillet Wings macropterous. Dorsal connexivum of abdomen bare (but with micropile), except for segmentally arranged pairs of setae. Median suture of prescutum usually short, never complete 5 5. Hind tibiae long and slender, with numerous short setae, lacking conspicuous macrosetae on upper edge 6 Hind tibiae with at least two, sometimes six or eight macrosetae on upper edge; in some species as many as 12-13 setae conspicuously longer than the others Strebla Wiedemann 6. Postgenae, behind the ctenidium, each with a laterally directed remiform scale (fig. 39D, r.s.) Anastrebla n.g. Postgena without a remiform scale Paraeuctenodes Pessoa & Guimaraes 7. Wings normally developed and functional 17 Wings much reduced or absent 8 8. Wings absent, or represented only by a very minute, inconspicuous flap 9 Wings reduced, but conspicuous 10 9. Minute species, body 1.29-1.48 mm. long. Legs of approximately equal size. Scutum membranous except for a short sclerotized strip connecting the pre- scutum and scutellum on each side (fig. 114A) Paradyschiria Speiser Larger species, body 1.70-2.48 mm. long. Hind legs greatly elongated, at least twice as long as the front legs (fig. 101A). Scutum sclerotized, the suture be- tween it and the scutellum rigid Megistopoda Macquart 10. Venter of thorax shield-like (fig. 98), anterior and posterior margins broadly rounded, the anterior margin dorsally reflexed and runner-like. Hind legs elongated, conspicuously longer than the others 16 Venter of thorax not thus. Hind legs, if elongated, not conspicuously longer than the others 11 11. Median mesonotal suture extending posteriorly beyond the transverse suture to the scutellum (figs. 79, 83) 15 Median mesonotal suture not extending posteriorly beyond the transverse suture. . .12 12. Median and transverse mesonotal sutures united to form an inverted "Y" (fig. 108 A) ; longitudinal and vertical membranous clefts closed without evidence of a suture. Laterovertices of head each with a longitudinal pigmented suture .... Noctiliostrebla n.g. Median and transverse mesonotal sutures, united to form an inverted "T" (fig. 103A) ; longitudinal and vertical membranous clefts present (open) or, if closed, the line of fusion is marked by a heavily pigmented, rigid suture 13 13. Posterior margin of head rounded. Median mesonotal suture not bifurcate ante- riorly; both longitudinal and vertical membranous clefts open 14 Head with two oblique flap-like occipital lobes, whose truncated posterior margins are festooned with setae (fig. 105E). Median mesonotal suture bifurcate ante- riorly; longitudinal and vertical membranous clefts closed, the line of fusion marked by heavy pigmented sutures Exastinion n.g. 14. Palpi slightly longer than wide, nearly vertical. Dorsum of abdomen clothed with short setae. Male : Sternum V absent ; gonapophyses with accessory setae inserted posterior to the macrosetae. .' Aspidoptera Coquillet Palpi oval, nearly horizontal. Dorsum of abdomen clothed with long setae (figs. 76A, 78A). Male: Sternum V present, divided into two rounded sternites; gonapophyses with accessory setae inserted anterior to the macrosetae (fig. 78C ) Anatrichobius n.g. 15. Minute species, 0.73-1.29 mm. long, with short legs. Wings with indistinct vena- tion; with an (rarely, two) apical, setigerous, digitiform process (fig. 83) Mastoptera n.g. Large species, 3.96-5.55 mm. long, with powerful, elongated legs. Wings with distinct venation, without an apical digitiform process (fig. 79) Joblingia Dybas & Wenzel WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 429 16. Inner face of profemora with stout spines (fig. 97). Mesepisternum divided into dorsal and ventral parts by a horizontal, longitudinal, membranous cleft similar to that separating the prescutum from the mesepisternum. Wings with only three longitudinal veins (fig. 99C) Neotrichobius n.g. Inner face of profemora lacking stout spines ; mesepisternum not divided into two parts by a membranous cleft. Wings with four or more longitudinal veins (figs. 100D, 101E) Megistopoda Macquart 17. Middle of anterior prescutal margin (fig. 115A) with two closely placed, sharp teeth which fit into grooves on the posterior part of the head ; humeral calluses strong, flat projections which fit under posteroventral margin of head. Costal vein heavily sclerotized, wider and bearing strong setae from base to junction with Ri, beyond which it is narrower, less strongly sclerotized, and bears short setae ; r-m near fork of Rs (fig. 115A) Synthesiostrebla Townsend Anterior margin of prescutum often sinuate, with blunt median projections, but never with a median pair of sharp teeth. Costa usually rather uniform in width and sclerotization throughout its length, r-m situated some distance from fork of Rs (fig. 86A) 18 18. Upper surface of posterior tibiae with at least two, sometimes five or six, macrosetae which are conspicuously longer than the others 22 Upper surface of posterior tibiae with setae more or less uniform, never with con- spicuously longer macrosetae 19 19. Venter of thorax shield-like, as in Megistopoda and Neotrichobius (fig. 98), broadly rounded anteriorly and posteriorly, the anterior margin reflexed up- wardly. Inner surface of profemora with a diagonal row of heavy spines (represented by strong setae in one species) ; hind legs greatly elongated Paratrichobius Costa Lima Venter of thorax not thus, the lateral margins oblique anteriorly and posteriorly. Inner face of profemur without spines, though strong setae may be present (fig. 102) ; hind legs may or may not be greatly elongated 20 20. Palpi with setae along margins only, ventral surface bare, the dorsal surface with micropile but appearing bare under low magnifications. Ri united with costal vein opposite third crossvein, both with strong macrosetae to this point Trichobioides n.g. Palpi with setae on ventral surface and along margins. Ri united with costal vein at a point distinctly beyond level of third crossvein 21 21. Hind legs greatly elongated and twice as long as the forelegs Speiseria Kessel Hindlegs not twice as long as forelegs ; if elongated, then fore and middle legs are also elongated Trichobius Gervais 22. Occipital lobes produced posteriorly as broad flaps (fig. 119A) which overlap the antero-lateral margins of the prescutum, the latter lacking a median suture and produced anteriorly as a truncate median projection which fits between the occipital flaps; genae and postgenae evenly covered with numerous short, posteriorly directed setae of nearly uniform size Stizostrebla Jobling Posterior margin of occipital lobes rounded, not flap-like, though a short tubercle may be present; genae and postgenae with both long and short setae; anterior margin of prescutum sinuate, with a bilobed median projection, the median su- ture well developed, usually complete 23 23. Head distinctly broader than long, and as broad as thorax or nearly so ; postgenae and occiput (viewed from beneath) conspicuously expanded laterally; postver- tex sclerotized, separated from the occipital lobes by a dark suture. Female: Ventral abdominal connexivum without a subapical row of blunt spines; cerci not united with the ventral arc Pseudostrebla Costa Lima Head distinctly narrower than thorax; postgenae and occiput not conspicuously expanded; postvertex membranous as in Trichobius and Paratrichobius. Female: Ventral abdominal connexivum with a subapical row of stout spines (fig. 116B) ; cerci united with the ventral arc Parastrebla n.g. 430 ECTOPARASITES OF PANAMA Subfamily Nycterophiliinae Wenzel, new subfamily Until now, only one species of the genus Nycterophilia has been de- scribed. All specimens recorded by authors have been referred to the type- species, N. coxata Ferris, 1916. The senior author has segregated six new species in the collections of Chicago Natural History Museum. With- out doubt, additional species exist. Three new species have been identified in the collections from Panama. The species of this genus differ remarkably from other Streblidae, and one can argue that a new family should be created for them. However, this should not be done until detailed comparative studies are made of the morphology of both Old World and New World Streblidae (including the Ascodipterinae) and of the Nycteribiidae. For this reason, we only propose a new subfamily at this time. DIAGNOSIS: Body strongly, laterally compressed, flea-like. Thorax. — Short and very deep. Mesonotum without a median suture. Spiracles situated just above midpoint between dorsal and ventral thoracic margins. Wings. — Either macropterous or micropterous (strut-like) ; when fully developed, with much reduced venation, only a few anterior veins and remnants of vannal veins sclerotized. Legs. — Femora and tibiae with unsclerotized annuli similar to those of Nycteribiidae. Profemora greatly enlarged and strongly, laterally compressed. Tibiae with marginal incrassations (seen in slide preparations). Abdomen. — Seven pairs of spiracles present, the first spiracle lying along or above the upper margin of sternum I, the second in the connexival membrane between the posterior angles of tergum I and sternum I. Terga I and II separated ventrally by a membranous suture, fused dorsally. Sternum I longer than II. Female : Intermediate sterna not sclerotized or only very feebly sclerotized ; tergum VII a very small plate if sclerotized; terminal cone setose, dorsal to proctiger. Seventh sternites present, ventro-lateral in position. Also a small ventral sclerite and strip anterior to vulva. Male: Sterna III-VI well developed as distinct sclerites; VII and VIII united to form longitudinal portion of the hypopygium; tergum IX bearing a pair of claspers with a terminal movable harpago, an inner pair of blade-like gonapophyses, and a ribbon-like aedeagus with a short non-coiled apodeme (fig. 45 A). The strongly compressed, flea-like body, the distinctive wing venation, the external male terminalia, the claspers with movable harpago, and the full complement of pre-abdominal sterna in the male, clearly separate this from the other subfamilies of Streblidae. Further, the position of the first and second spiracles differs greatly from that of other New World Streb- lidae, in which both are situated in tergum I+II. Jobling (1951) did not discuss the systematic position of Nycterophilia, though he apparently regarded it as belonging to the Trichobiinae. Pessoa and Guimaraes (1940) assigned the genus to the Nycteriboscinae, but as noted earlier, they appear to have misinterpreted the structure of the head, probably because of a lack of material suitable for study. Nycterophilia does possess some features in common with the Nycteriboscinae, especially in the structure of the sterna IX and X in the female. However, similar sclerites are found in some of the Trichobiinae. The genus appears to be more closely related to the New World Streblidae. The presence of paired ventral setae on the male gonapophyses suggests this, as does the number and position of the abdominal spiracles in both sexes, the divided female seventh sternum, and the structure of the male hypopygium. It especially WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 431 appears to be related to Trichobius species of the caecus and pallidus groups, which, for the most part, occur on the same hosts as Nycterophilia. In these, the male cerci are very long and are connected by apodemes to the surstyli, suggesting a resemblance to the clasper shaft and harpago of Nycterophilia. On the other hand, the non-pigmented annuli of the legs, the abdominal segmentation of the male, and the external claspers suggest a relationship with the Nycteribiidae. Fig. 46. Nycterophilia parnelli, new species, holotype male; lateral view. We tentatively suggest that both Nycterophilia and Ascodipteron repre- sent specialized developments from a stock which gave rise to both Stre- blidae and Nycteribiidae. Ascodipteron should probably be regarded as belonging to a separate family, as it was by earlier authors. 432 ECTOPARASITES OF PANAMA Genus Nycterophilia Ferris Nycterophilia Ferris, 1916, Ent. News, 37: 436-437 (diagnosis). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., 5: 25 (keyed), 30 (cit.). Stiles and Nolan, 1931, Bull. Nat., Inst. Hlth., Wash., no. 155, p. 654. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed); 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 361, 362 (morph.). Bequaert, 1940, Rev. Acad. Colomb. Cien. Exact., Fis. y Nat., 3: 417 (keyed) ; 1942, Bol. Ent. Venez., 1: 85 (keyed). Jobling, 1949, Parasitology, 39: 321 (keyed); 1951, Trans. Roy. Ent. Soc. Lond., 102: 215 (morph.). Maa, 1965, Jour. Med. Ent., 1: 385 (cit.). Nycterophila Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 12 (keyed), 13. Pessoa and Guimaraes, 1940, Arq. Inst. Biol. Sao Paulo, 11: 421-424 (syst. pos.). Schuurmans Stekhoven Jr., 1941, Zool. Anz., 136: 80. Type-species : Nycterophilia coxata Ferris, 1916. DIAGNOSIS: Body form strongly, laterally compressed, flea-like. Head. — Vertex strongly projecting as dorsal lobes; occiput and post-occiput obliquely declivous, the posterior surface concave and apposed to the anterior margin of thorax. Laterovertices differentiated, bearing the eyes; occipital plates consisting of thin, feebly sclerotized transverse strips. Eyes not always differentiated, never with more than a single facet. Each postgena ventrally produced, below oral cavity, as a process bearing a stout seta; a series of two to five short spinelets present on each side immediately below oral cavity. Theca and palpi vertical in repose. Thorax. — Short, deep, anterior margin (figs. 46-48) inwardly declivous. Mesonotum without a median suture; transverse suture strong, complets. Scutellum very short and strongly convex, with a pair of closely appressed (partially fused) macrosetae which arise from the same theca. Longitudinal and vertical membranous clefts present, well- defined. Spiracles situated behind anterior margin about halfway between dorsal and ventral margins, or slightly above this. Pleurotrochantinal plate movable, separated from sternopleura by a membranous suture. Wings. — Emarginate distally in macropterous species ; costa, a longitudinal vein, and part of a third vein normally developed and sclerotized anteriorly, and with sclerotized rudiments of two veins in vannal region ; also with nine longitudinal rows of setae which may represent reduced veins; longitudinally folded in repose. Wings of micropterous species reduced to short strut-like structures without identifiable venation. Halteres large, conspicuous (fig. 50A, hit.). Legs. — Femora and tibiae each with a non-pigmented cuticular annulus similar to those of Nycteribiidae. Inner face of profemora with a few discal setae, bearing a pair of heavy spines near base and a blunt spine at about middle above ventral margin, as well as smaller psg-like submarginal spines antero-dorsally; with a ventral groove on inner face for reception of the protibia; outer face with more numerous setae; upper margin with both short setae and macrosetae; with an irregular and incomplete non- sclerotized annulus. Procoxae large, projecting dorsally; latero-ventral and ventral poste- rior margins armed with heavy or spiniform setae, which form a "pseudo-ctenidium". Middle and hind femora normal, spindle-shaped, clothed with setae and with a few macro- setae, the hind femora longer than the middle ones. Hind tibiae with a pair of conspic- uous subapical, bifid, spine-like setae; inner face with dense, short setae on apical half. Protarsi with minute setae ventrally, with spinelets dorsally and laterally. Middle and hind tarsi longer than tibiae ; inner face of first segment of hind tarsi with dense, short setae similar to those on inner face of hind tibiae. Tarsal claws with oblique parallel grooves on outer face, sometimes appearing pectinate, though the ventral margin does not seem to be incised. Abdomen. — Terga I and II separated ventrally by a vertical membranous suture, fused dorsally; a vertical row of marginal setae present on I along suture; lateral lobes of II with discal setae and, on posterior margin, three heavy "braided" spiniform setae, the dorsal one much smaller than the other two. Spiracles situated as follows : first pair just above middle of dorsal margin of sternum I; second pair just above dorsal angle of sternum I on line with interval between I and II; third pair in connexival membrane just WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 433 below posterior angles of tergum II and above interval between sterna II and III; fourth to sixth pairs situated more dorsally than the preceding, in the connexival membrane just above the lateral connexival setae and opposite intersternal membranes; seventh pair situated more ventrally than the three preceding, being just anterior to the hypopygium and dorsal to postero-dorsal edge of seventh sternites, in the female, and in the middle of the dorsal edge of synsternum VII+VIII, in the male. Female: Segments III-VI represented, at most, by paired median, dorsal con- nexival setae; VII represented by setae or by a small setose sclerite. Terminal cone prominent, dorsal to the proctiger, the latter separate and free (fig. 51). Lateral con- nexivum with micropile and coarse setae which are coarser than those on ventral half. Sterna III-VI at most very feebly sclerotized, if at all, but represented by vertical rows of coarse setae which become coarser dorsally. Ventral to the proctiger is a small sclerotized plate, connected by a sclerotized strip to a small sclerite in front of the vulva. Seventh sternites normally triangular, ventro-lateral in position. Male: Segments III-VI may also be represented by median paired dorsal connexival setae. Sterna I-VI well developed and sclerotized with coarse spiniform paired median ventral setae and, along sides, with marginal setae along posterior margin, these becom- ing coarser and "braided", dorsally, similar to the lateral connexival setae; sternum VI fused along median ventral line with anterior portion of hypopygium, the latter long, sclerotized, membranous dorsally except at apex where it is sclerotized and "closed" to form a "ring" together with the ventral portion. The apical portion of the hypopygium a separate sclerotized segment bearing the terminalia. These consist of a long clasper- like, ventral shaft, on each side, which distally bears an articulated "harpago"; a pair of conspicuous gonapophyses between the "claspers" and articulated basally with the ventral posterior margin of the hypopygium ; and, between the gonapophyses a ribbon-like (viewed dorsally) aedeagus. The aedeagal apodeme is relatively short and is not coiled. Tergum IX with dorso-apical macrosetae. Cerci a pair of small setose flaps on each side of anus. Spiracles situated very much as in female, but seventh pair situated just within dorsal margin, of hypopygial "ring" at mid-length. According to Ross (1961, p. 241), the greatly enlarged first pair of legs [profemora] "is not used in locomotion by N. coxata, but is held forward at the sides of the head in such manner as to part the fur as it progresses through it." Their well developed musculature suggests that these have other functions, too. These flies resemble fleas so closely in their movements that they deceive even the expert. When on the host, the wings are folded longitudinally and held over the upper sides of the abdomen. Thus, the similarity to fleas is enhanced. The forms with reduced strut-like wings must be even more flea-like in life. KEY TO THE SPECIES OF NYCTEROPHILIA 1. Dorso-apical spur of hind coxa long, conspicuous (fig. 50 A) 3 Spur short, inconspicuous, sometimes nipple-like (figs. 47A, B) 2 2. Antero-lateral angles of thorax sparsely setose ; mesepisternum with a row of sub- marginal bristles along margins of both the longitudinal and vertical mem- branous clefts and five or six other, discal setae7 (fig. 47A). Female: Terminal cone with two apical macrosetae (fig. 51C). Male: Gonapophyses (lateral view) strongly but evenly tapered from base to apex. Host: Pteronotus parnellii .... parnelli n.sp. 7 The discal setae are those in an imaginary quadrangular area bounded by the longitudinal and vertical membranous clefts and the inner edge of the anterior thoracic spiracle (figs. 47A, d.s.; 48). 434 ECTOPARASITES OF PANAMA Antero-lateral angles of thorax rather densely setose; mesepisternum with 13 or 14 discal bristles (fig. 47B). Female: Terminal cone with four apical macrosetae (fig. 51B). Male: Gonapophyses (lateral view) abruptly tapered apically. Hosts: Pteronotus psilotis, P. suapurensis fairchildi n.sp. 3. Each lateral lobe of tergum II with two large and one much shorter spinelets on posterior margin plus nine or ten other setae. Postgenae with two or three min- ute spinelets. Male: Paired dorsal connexival setae on abdominal segments III-V conspicuous (fig. 50A). Host: Natalus stramineus mexicanus. . . .natali n.sp. Each lateral lobe of tergum II with six to eight setae in addition to the posterior spinelets. Postgenae with four or five minute spinelets. Male: Paired dorsal connexival setae on segments III-V very fine, short, inconspicuous (fig. 50B). Hosts : Macrotus mexicanus, M. calif ornicus coxata Ferris Nycterophilia parnelli Wenzel, new species. Figs. 45A, 46, 47A, 49A, 51C. Nycterophilia coxata (not Ferris, 1916), Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (part.). Jobling, 1949 (part.), Parasitology, 39: 316 ff., 327. Bequaert 1940, Rev. Acad. Colomb. Cien. Ex. Fis. y. Nat., 3: 418. Closely related to fairchildi n. sp., and like it having a short metacoxal spur, but differing in thoracic chaetotaxy (fig. 47 A) , in that the female has only two (instead of four) apical macrosetae on the terminal cone and lacks a row of ventral abdominal spiniform setae anterior to the vulva, and the male gonapophyses are strongly but rather evenly tapered from base to apex (lateral view). DESCRIPTION: Head. — Eyes conspicuous, pigmented. Laterovertices with 10-11 setae of variable size, two or three pairs conspicuously longer than the others though they are not macrosetae ; occiput with a pair of dorsal macrosetae on each side ; posterior margins of head on each side with a vertical row of seven to nine short, distinct setae; side of head with a transverse row of five to seven short setae across middle; three or four postgenal spinelets present. Thorax. — Mesonotum covered dorsally with short setae. Chaetotaxy of sides as shown in fig. 47A. Scutum with six prominent setae anteriorly, consisting of two on each side and a median pair; the latter followed posteriorly by a median pair of shorter setae; scutellum with a median pair of closely appressed, often partially fused very long prominent macrosetae that arise from the same theca; lateral to the scutellum a macrossta and a short seta on each side. Wings. — As in N. coxata. Legs. — Profemora with five macrosetae alternating with short setae on upper margin; outer face with dorsal submarginal and numerous discal setae arranged in irregular longitudinal rows, inner face with three peg-like spines above, two of them more slender and situated near middle, a longer one near apical third; a ventral submarginal spinelet near base ; a large curved blunt spine above margin near middle and a short weak spine- let basal to it. Protibiae feebly tapered apically; incrassations of the margin not strong; in addition to marginal and submarginal setae of outer face there is a median longitudinal row, the uppermost setae of this row strong, coarse. Metacoxa with a short inconspic- uous dorsal spur (fig. 47A). Other tibiae and tarsi very similar to those of N. coxata. Abdomen. — Female: Tergum I with two setae, one above the other, along middle of antero-lateral face; posterior margin with. short fine setae. Tergum II at middle of posterior margin with four very slender long setae, two on each side; lateral lobes with two large stout spiniform setae at posterior margin, a shorter spiniform above them and fine setae anterior to these. Dorsal connexivum bare except as follows: segments III- V represented dorsally by paired macrosetae, those of VI and tergum VII shorter; III represented by two pairs, these "connected" to setae of lateral connexivum on each side by a row of setae slightly anterior to them; tergum VII small, feebly sclerotized, with four setae. Terminal cone with a single pair of dorso-apical macrosetae and ventral to these about eight setae of varying lengths, one of these (antero-median) much longer than the others. Posterior margin of sternum I with a median ventral pair of strong setae and dorsally with a row of setae, the more ventral ones very fine and short, the WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 435 d.s. A e. Fig. 47. Head and thorax, lateral view. A, Nycterophilia parnelli, new species, female paratype from Pteronotus pamellii fuscus (no. 7167), Madden Airstrip (Panama). B, N. fairchildi, new species, female paratype from Pteronotus suapurensis (no. 8777), Penonome Caves (Cocle) . See text for explanation. 436 ECTOPARASITES OF PANAMA setae becoming coarse, "braided", dorsally. Posterior margin of sternum II with braided setae, these becoming longer, stout and braided and similar to the connexival setae dorsally; ventro-median edge of sternum V with a fringe of somewhat longer setae; VI bare and connected with VII+VIII along ventral midline. Measurements: BL TL WL ww Male 1.35-1.40 0.42-0.46 1.15-1.32 0.55-0.58 Female 1.42-1.65 0.44 1.21-1.32 0.60-0.66 TYPE MATERIAL: Holotype male and allotype female (slides) from Pteronotus parnellii fuscus (host no. 3924) , railroad culvert, Paraiso (Canal Zone), C. M. Keenan and V. J. Tipton, 24 July 1959. In the collection of Chicago Natural History Museum. Paratypes. — 116 specimens (18 lots) all from Pteronotus parnellii fuscus as follows : 43 (1 lot) , railroad culvert, Paraiso (Canal Zone) , 24 July 1959 ; 8 (6 bats), same locality 15 and 16 September 1959, and 1, same locality, 1 December 1959 ; 2, railroad culvert east of Summit Golf Club (Canal Zone) , 26 October 1959; 12 (1 bat), Madden Air Field, Casa Larga (Panama), 23 May 1961 ; 12, Chilibrillo Caves (Panama) , L. H. Dunn no. 271 [MCZ] ; 3, same locality (labeled as from "Chilonycteris r. rubiginosa") [cu] ; 3 (3 bats), Guanico (Los Santos), 22 January and 16 February 1962; 1, Cerro Hoya (Los Santos), 14 February 1962; 7, Penonome (Code), L. H. Dunn [MCZ] ; 26, same locality, in cave, 11 March 1954, H. Trapido. Paratypes to be deposited in the collections listed on p. 410. OTHER MATERIAL EXAMINED: From Carollio, perspicillata azteca (!): 2, Buena Vista (Colon), 3 September 1959; 2 (from preserved bats in museum collection), Chilibrillo Caves (Panama), 30 August 1935. In ad- dition, we have seen numerous specimens which we tentatively assign to this species, from various hosts, taken in MEXICO, GUATEMALA, VENEZUELA, and TRINIDAD. Nycterophilia fairchildi Wenzel, new species. Figures 47B, 49B, 51B. ? Nycterophilia coxata (not Ferris, 1916), Pessoa and Guimaraes, 1940, Arq. Inst. Biol., 11: 421. Very similar to N. parnelli n. sp. and, like it, with a short inconspicuous dorso-apical spur on metacoxa; differing conspicuously from it in the thoracic chaetotaxy (fig. 47B) ; in that the female has four (instead of two) apical macrosetae on the terminal cone and has a ventral transverse row of about eight heavy spiniform setae anterior to the vulva; and in that the male gonapophyses are broad at base and abruptly tapered apically (in lateral view). DESCRIPTION : Very similar to parnelli in most characters. Head. — With three post genal spinelets. Thorax. — Chaetotaxy as in fig. 47B. Wings and legs. — As in parnelli. Abdomen. — Female: Lateral lobes of tergum II with eight setae in addition to the three coarse marginal spiniform setae. Chaetotaxy of dorsal connexivum very similar to that of parnelli, except that setae of segment VI are lacking; the setae on III are connected to lateral connexival setae by a group of five or six setae on each side. Terminal cone with four apical macrosetae (two each side) and five or six other setae on each side. Sterna I and II similar to those of parnelli, the lateral disc of II with a row of three to five setae in addition to the setae along posterior margin. Seventh sternites with about 12 setae, of which the ventral and dorso-apical ones are macrosetae, the others of varying lengths. Anterior to the vulva (ventral and between the seventh sternites) is a trans- WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 437 A l.m.c Fig. 48. Head and thorax, lateral view. A, Nyctcrophilia natali, new species, female from Natahis sp. (CNHM host nos. 64731-807) , Santa Clara, Sierra de las Minas (Zacapa) , GUATEMALA. B, N. coxata Ferris, from Macrotus mexicanus, La Galarza (Puebla). MEX- ICO. See text for explanation. 438 ECTOPARASITES OF PANAMA verse sclerotized strip which bears a row of eight conspicuous spiniform setae, four on each side appearing somewhat like a ctenidium. Male : Lateral lobes of tergum II each with 9-11 setae in addition to the very heavy spiniform setae on posterior margin. Dor- sal connexivum bare except for very fine, very short paired segmental setae on III-IV. Sternum VII+VIII with about 13 or 14 setae on each side. Upper portion of segment IX with about 14 setae in addition to the stout dorsal macroseta; shaft of claspers typically with a row of five, sometimes four, setae; harpago with a long basal ventral seta, a short very fine sub-basal dorsal seta, two short setae at about basal third, two very fine distal ventral setae and some apical microsetae that are visible only under oil immersion. Gonapophyses strongly but rather evenly tapered from base to apex. Measurements: BL TL WL ww Male 1.37 0.44 1.13-1.15 0.55-0.60 Female 1.59 0.47 1.29 0.55 TYPE MATERIAL: Holotype male and allotype female (slides) from Pteronotus suapurensis (host no. 8777), Penonome Caves (Code), 15 De- cember 1961, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — From Pteronotus suapurensis: 9(3 bats) , same data as the holotype; 6, no locality (CBK host no. 5014), 2 May 1957 [USNM]. From Pteronotus psilotis: 4 (3 bats), same locality, collectors and date as holo- type; and 1, same data but 24 January 1962. From Pteronotus sp. : 58, same locality as holotype, 11 March 1954, H. Trapido. From "Pteronotus davyi" : 3, same locality, without date, L. H. Dunn [MCZ] . Non-Panamanian paratypes. — From Pteronotus personatus: 47, Cartagena, COLOMBIA, 22 June 1965, C. J. Marinkelle. Paratypes to be deposited in the collections listed on p. 410, and in the collection of C. J. Marinkelle, Universidad de los Andes, Bogota, Colombia. OTHER MATERIAL EXAMINED : We have seen additional specimens, which we tentatively assign to this species, from MEXICO, GUATEMALA, and VENE- ZUELA. REMARKS : To our knowledge, Pteronotus davyi, listed as a host for the specimens collected by Dunn [MCZ], does not occur in Panama. N. fairchildi is named in honor of Dr. Graham B. Fairchild of the Gorgas Memorial Laboratory, Panama, Panama. Nycterophilia natali Wenzel, new species. Figures 48A, 50A. Nycterophilia coxata (not Ferris, 1916), Hoffmann, 1953 (part.), Mem. Congr. Cient. Mex., 7: 183, 187. N. natali n. sp. differs from both fairchildi and parnelli in that it has a conspicuous long (rather than a short) dorsal metacoxal spur. In this re- spect, it resembles N. coxata. However, the male holotype differs from coxata in that the pairs of dorsal segmental connexival setae on III-V are conspicuous rather than very fine and short; and the "harpago" of the male terminalia lacks a long, conspicuous, ventral, basal seta. N. natali also differs from coxata in the number of postgenal spinelets and in the number of setae on the lateral lobes of abdominal tergum II, as indicated in the key. In addition to characters natali has in common with fairchildi, parnelli, and coxata, the following may be noted : DESCRIPTION: Head. — Setae of posterior margin of head finer and shorter than in fairchildi and parnelli; three or four postgenal spinelets present. Thorax. — Ventral WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 439 MI sp.VII t.IX Fig. 49. Male abdomen. A, Nycterophilia pamelli new species, paratype from Pterono- tus parnellii fuscus (no. 7167) . B, N. fairchildi, new species, paratype from Pteronotus suapurensis (no. 8777). See text for explanation. 440 ECTOPARASITES OF PANAMA A B Fig. 50. Male abdomen, lateral view. A, Nycterophilia natali, new species, holotype. B. N. coxata Ferris (same specimen as 48B). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 441 A pr. c Fig. 51. Female abdomen, lateral view. A, Nycterophilia coxata Ferris, from Lep- tonycteris nivalis, Pima County, Arizona. B, N. fairchildi, new species, from Pteronotus suapurensis (no. 8777). C, N. parnelli, new species, from Pteronotus parnellii fuscus (no. 7167). See text for explanation. 442 ECTOPARASITES OF PANAMA setae of procoxae spine-like. Lateral lobes of tergum II with the usual heavy spine-like setae on posterior margin, and anterior to these, nine additional setae. Segmentally ar- ranged dorsal pairs of connexival setae on segments III-V moderately long, conspicuous. Hypopygial "ring" with eight setae on each side. Segment IX on each side above the ventral shaft, with nine or ten setae in addition to the dorso-apical macroseta; shaft with four setae along posterior edge; harpago without a long ventral basal seta, but with a fine rather short sub-basal seta dorsally and another ventrally, as well as micro- setae on apical half of inner and dorsal margins. Measurements: BL TL WL ww Male (Holotype) 1.32 0.44 1.21 Female (GUATEMALA and MEXICO) 1.52 0.47-0.48 1.36 0.66 TYPE MATERIAL: Holotype male (slide) from Natalus stramineus mexi- canus (host no. 6732) , San Lorenzo Caves, Fort San Lorenzo (Canal Zone) , 15 March 1961, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. OTHER MATERIAL EXAMINED : We have several specimens that appear to be natali, taken from Natalus sp. in Guatemala and Mexico. However, the Trichobius from Natalus s. mexicanus in Panama appears to be different from that (galei) taken from the same host in Guatemala and it is pos- sible that the Nycterophilia may be different, too. This may be easier to determine when series of both sexes are known from both localities. REMARKS : The females from Guatemala differ from those of coxata in the characters noted above. On the ventral margin anterior to the vulva there is a row of coarse setae set somewhat apart from the others. These may be homologous to the spiniform setae borne on a sclerotized strip in fairchildi n. sp. Subfamily Trichobiinae Genus Trichobius Gervais Trichobius Gervais, 1844, Atlas. Zool., p. 14, pi. 53. Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 62 (cit.), 65 (key), 33-37 (notes on morph.). Brues, 1904, Bull. Amer. Mus. Nat. Hist., 20: 131-134. Coquillet, 1910, Proc. U. S. Nat. Mus., 37: 616. Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Veter., Nictheroy, 5: 25 (keyed), 26-28 (cit., records). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 12 (keyed), 14-18 (synopsis). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 655. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; Bull. Amer. Mus. Nat. Hist., 66: 522 (keyed); 1935, Amer. Mus. Novit., no. 765, p. 6. Jobling, 1936, Parasitology, 28: 357 ff. (morph., syst. pos.). Guimaraes, 1937, Rev. Mus. Paul., 23: 653-666, 2 pis., (S. Am. spp.). Jobling, 1938, loc. cit., 30: 358-387, figs. 1-14 (revision of species). Bequaert, 1940, Rev. Acad. Colomb. Cien. Exact., Fis. y Nat., 3: 417 (keyed); 1942, 'Bol. Ent. Venez., 1: 86 (keyed), 87-88 (rec- ords, Venezuela). Schuurmans Stekhoven Jr., 1941 (& 1951), Beitr. Fauna Perus, 2: 95 (Peruvian species). Jobling, 1949, loc. cit., 39: 321 (keyed), 326 (records, Trinidad spp.). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 178 (keyed), 179- 182 (key, Mexican spp.), 182-183 (records, Mexican spp.). Maa, 1965, Jour. Med. Ent., 1: 384 (list of spp.) Trichobius Townsend, 1891 [with dugesii Townsend as type-species], Ent. News, 2: 105-106. Williston, 1896, Man. Fam. Gen. N. Am. Dipt., 2nd ed., p. 152 (keyed). Trichobia Guerin-Meneville, 1844, Icon. Regne Anim., 3: 556. Bigot, 1885, Ann. Soc. Ent. Fr., 5: 228. Strebla Kolenati, 1863, Hor. Soc. Ent. Ross., 2: 11 (not Wiedemann, 1824). WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 443 Kolenatia Rondani, 1878, Ann. Mus. Civ. Stor. Nat. Geneva, 12: 169 [type-species: Strebla wiedemannii Kolenati 1863, not Kolenati 1856]. Bigot, 1885, Ann. Soc. Ent. Fr., (6), 5: 232 (keyed). Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 33, 43 (synonymizes Kolenatia under Trichobius). Kesselia Curran, 1934, Bull. Amer. Mus. Nat. Hist., 66: 522 [type-species: Kes- selia pallida Curran]; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 359 (venation). Type-species : Trichobius parasiticus Gervais, 1844. Seventeen previously described species of this genus are considered by us to be valid. One of these is assigned to a new genus, Trichobioides. Twenty-seven species are treated in this paper. Of these, six previously described and 17 new species are recorded from Panama. T. caecus Ed- Fig. 52. Wing, Nycterophilia sp. From Jobling (1949b). wards and T. furmani n. sp. are included, in order to assist in defining closely related Panamanian species. T. pallidus Curran (in key) and T. diphyllae n. sp. are included, because their hosts have been taken in Panama and the flies may also be found there. The new species treated below bring the total number of described spe- cies of Trichobius to 34, making it the largest new world genus of Streblidae. In the collections of Chicago Natural History Museum, there are about a dozen additional species. Undoubtedly, many more will be discovered. The genus includes diverse types, representing at least six, and pos- sibly more, groups or species complexes. In the major group, we include T, major Coquillet, hirsutulus Bequaert, corynorhini Cockerell, pseudo- truncatus Jobling, sparsus Kessel, and several undescribed species from Central America and the West Indies; T. sphaeronotus Jobling and trun- catus Kessel appear to belong here. The species of the major group occur primarily on Vespertilionidae, though one may occur secondarily on Molos- sidae of the genus Tadarida. T. sphaeronotus occurs on Leptonycteris, a genus of Glossophaginae (Phyllostomidae). The caecus group includes T. caecus Edwards, three new species de- scribed below, and several additional undescribed species. All occur on bats of the families Natalidae and Chilonycteridae. T. pallidus Curran, which was taken on Furipterus (Furipteridae) superficially appears to be related to species of the caecus group. However, 444 ECTOPARASITES OF PANAMA it differs in several essential features and until a series can be studied in detail, we would place this species by itself. Two undescribed species from Peru, one from a furipterid (Amorphochilus schnablii) , seem to be related to pallidus but have a simple median suture. The relationships of T. adamsi Augustson are not clear. It may be a spe- cialized member of the major group. The species of the phyllostomae group, including T. phyllostomae Kessel, T. brennani n. sp., T. vampyropis n. sp. and an undescribed species from Brazil occur on fruit-eating bats of the subfamily Stenoderminae (includ- ing Sturnirinae of authors). Most of the remaining described species belong to the related dugesii and longipes groups. The species of the longipes group occur mostly on Phyllostominae of the genera Phyllostomus and Tonatia, although T. dunni n. sp. and several related undescribed species are apparently restricted to bats of the genus Molossus. The species of the dugesii complex are ap- parently mostly specialized types derived from species of the longipes com- plex and have radiated from the Phyllostominae to the Stenoderminae, Carolliinae, Glossophaginae and Desmodidae. KEY TO PANAMANIAN SPECIES OF TRICHOBIUS* 1. Sixth longitudinal vein without setae near basal angle (fig. 42) 2 Sixth longitudinal with setae at or near basal angle (fig. 61) 7 2. Median suture of prescutum bifurcate, short (fig. 56) 3 Median suture not bifurcate, usually extending to middle of prescutum or be- yond 10 3. Eyes with a single facet 4 Eyes with more than one facet pallidus Curran 4. A broad median area of mesonotum without micropile ; scutum at middle with only three transverse rows of bristles, including the antescutellars (fig. 54). Wing vein Rg twice as long as distance between fork and crossvein r-m. Female: Postgenital sclerite as in fig. 57K; seventh sternites with macrosetae along posterior margin, most discal setae well-developed ; chaetotaxy of terminal cone as in fig. 57 J. Male : Hypopygium with long, sparse setae dorsally and apically. Gonapophyses strongly curved, apices at approximately a right angle to base (fig. 57L). Host: Natalus stramineus mexicanus galei n. sp. Mesonotum and scutellum with micropile throughout ; scutum at middle with about four transverse rows of bristles including the antescutellars (fig. 56A). Rg slightly to no more than a third longer than distance between fork and cross- vein r-m. Female: Seventh sternites each with only three to five macrosetae along posterior margin, the rest of the setae small to minute. Male : Setae of hypopygium short except along apical margin 5 5. Chaetotaxy of lateral lobes of tergum I+II as in fig. 55A. Female : Postgenital sclerite as in fig. 57H; seventh sternites usually with three, sometimes four, macrosetae. Chaetotaxy of terminal cone as in fig. 57G. Male: Gonapophyses as in fig. 571. Hosts: Pteronotus suapurensis, P. psilotis johnsonae n. sp. Chaetotaxy of lateral lobes of tergum I+II as in fig. 55B. Host: Pteronotus par- nellii fuscus 6 8 Two of the included species, T. pallidus Curran and T. diphyllae n. sp. have not been recorded from Panama but may occur there. Two others, T. caecus Edwards and T. furmani n. sp., are included in the key in order to assist in differentiating between some closely related species that occur in Panama. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 445 6. Female: Inner pair of basal setae on tergum VII absent or very small (fig. 57A) ; seventh sternites each with three or four macrosetae near apical margin; sub- genital sclerite as in fig. 57B. Male: Gonapophyses as in fig. 57C caecus Edwards Female: The inner pair of basal setae on tergum VII well developed (fig. 57D) ; each seventh sternite usually with five or more macrosetae along apical margin ; postgenital sclerite as in fig. 57E. Male: Gonapophyses as in fig. 57F yunkeri n. sp. 7.9 Wing vein Ri strongly sinuate, the costal cell rather abruptly narrowed apically (fig. 59 A) ; Rs markedly longer than distance between fork and crossvein r-m, and the latter distance no more than twice the length of r-m. Female : With a distinct cluster of longer discal setae in posterior angles of sternum II. Male : Gonapophyses (fig. GOD), wedge-shaped in lateral view, ventral margin nearly straight, with a submarginal row of fine setae lonchophyllae n. sp. Wing vein Ri straight or only very feebly sinuate (fig. 61), the costal cell rather evenly tapered or nearly subparallel apically; length of Rs and distance be- tween fork and crossvein r-m subequal, the latter distance more than twice the length of r-m. Female : Marginal setae of sternum II longer toward posterior angles but discal setae not longer within the angles. Male: Gonapophyses (figs. 60A-C) ventrally curved and tapered, at most with one or two submarginal setae along ventral margin 8 8. Posterior margin of each occipital plate with a prominent posteriorly-directed tubercle which bears a very short spinelet-like seta; eyes separated by their width or more from lateral margin of head. Male: Gonapophyses as in fig. 60C uniformis Curran Posterior margin of each occipital plate with a short seta borne on an inconspic- uous tubercle, the seta not a spinelet; eyes extending to lateral margins of head or separated from margins by less than their width 9 9. Female: Four setae of tergum VII arranged in a transverse row. Male: Gon- apophyses as in fig. 60A. Hosts : Micronycteris megalotis, M. nicefori keenani n. sp. Female : Four setae of tergum VII arranged one pair behind the other, the anterior and posterior pairs widely separated. Male: Gonapophyses as in fig. GOB. Host: Lionycteris spurrellii lionycteridis n. sp. 10. Scutum and prescutum covered with setae 13 Scutum and prescutum with conspicuous areas devoid of setae (figs. 58B, 67B, 73B) 11 11. Prescutum with a longitudinal cluster of six to nine very short setae at middle anterior to the transverse suture (fig. 67B). Hosts: Phyllostomus spp costalimai Guimaraes Prescutum without such a cluster of setae 12 12. Size smaller, body 1.36-1.70 mm. long. Thorax distinctly flattened; prescutum with a transverse row of setae immediately anterior to transverse mesonotal suture and a median group of four to six setae, these sometimes separated from a lateral group on each side (fig. 73 A) ; W-shaped band of setae along posterior margin of scutum consisting of very short setae only. Hosts: Desmodus ro- tundus, Diaemus youngii parasiticus Gervais Size larger, body 1.65-2.25 mm. long. Thorax deeper, not flattened; row of setae anterior to transverse mesonotal suture broadly interrupted at middle (fig. 58B) ; band of setae along posterior margin of scutum with a row of very long antescutellar setae and anterior to these a transverse patch of very short setae. Host: Pteronotus parnellii fuscus sparsus Kessel 9 The species of the uniformis group are extremely difficult to identify without com- parative material, except by the structure of the male gonapophyses. 446 ECTOPARASITES OF PANAMA 13. Sternopleura strongly projecting between the procoxae; lateral margins of the projection subparallel apically (fig. 62D) 14 Sternopleura less strongly projecting between the coxae, the lateral margins oblique (fig. 62E ) 15 14. Size larger, body 2.20-2.65 mm. long. Prescutum two-thirds as long as broad, the anterior margin very strongly projecting at middle; mesonotal chaetotaxy as in fig. 74B. Wing vein Rs about as long as distance between fork and cross- vein r-m. Host: Vampyrops vittatus vampyropis n. sp. Size smaller, body 2.09-2.39 mm. long. Prescutum only slightly more than half as long as broad, the anterior margin projecting but less strongly so; mesonotal chaetotaxy as in fig. 74A. Wing vein Rs only slightly more than half as long as distance between fork and crossvein r-m. Host: Sturnira ludovici brennani n. sp. 15. Occipital lobes of head with 10-12 setae. Abdominal tergum I+II with a cluster of short setae on each side of middle of basal margin, a group of coarse long setae on lateral lobes, and a single seta (occasionally two) on anterior face, situated laterally and ventrally to median basal group 16 Occipital lobes of head with 17-19 setae. Abdominal tergum I+II with a cluster of about 10 short setae on anterior face on each side lateral and ventral to median basal cluster dunni n. sp. 16. Median pleurotrochantinal lobe present, reflexed dorsally and united with a descending process of the metepimeron so as to close the coxal cavities mesally (fig. 62A) 17 Median pleurotrochantinal lobe absent, or if one is present and reflexed, it is not united with the metepimeron so as to close the coxal cavities (figs. 62B, C) 20 17. Median anterior prescutal setae of about the same size as those along sides, and longer than median discal setae (fig. 66 A) ; prescutum with a distinct patch of micropile on each side in front of transverse suture, the patch extending mesad to or slightly beyond the second seta from lateral margin and anteriorly to second seta from transverse suture. Male: Gonapophyses as in fig. 63 A. Host: Tonatia minuta bequaerti n. sp. Median anterior prescutal setae distinctly shorter than lateral setae, often only slightly longer, if any, than dense short median discal setae (figs. 64, 65) ; scutum with a very narrow longitudinal band of micropile along inner margin of longitudinal membranous cleft, scarcely entering upon disk, if at all 18 18. Prescutal setae noticeably and rather abruptly denser on basal half of median discal area than they are laterally and anteriorly (figs. 64, 65B) 19 Prescutal setae gradually sparser anteriorly and laterally (fig. 65A). Male: Gonapophyses as in fig. 62B. Host: Tonatia silvicola dybasi n. sp. 19. Size larger, 1.76-2.35 mm. long. Genal-postgenal margins rather strongly convergent posteriorly. Male: Gonapophyses as in fig. 63F. Host: chiefly Phyllostomus hastatus longipes Rudow Size smaller, 1.59-1.73 mm. long. Genal-postgenal margins subparallel or feebly convergent for part of their length, then broadly rounded apically. Male: Gonapophyses as in fig. 63D. Host: Tonatia minuta mendezi n. sp. 20. Pleurotrochantinal lobe present (fig. 62B), this usually rounded, translucent, and feebly reflexed dorsally, though it may be long and strongly reflexed and superficially appear to unite with a ventrally directed spiniform process of the metepimeron. Male: Sternum VI present (except in macrophylli) 23 Posterior margin of pleurotrochantines without a median lobe (fig. 62C). Male: Sternum VI absent 21 21. Median discal mesonotal setae extremely minute and fine, sometimes almost in- distinguishable except in slide preparations at high magnification (fig. 73B). Host : Diphylla ecaudata diphyllae n. sp. Median discal mesonotal setae very distinct, easily visible even if minute 22 WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 447 22. Mesonotal setae gradually becoming longer posteriorly, the median antescutellar setae no more than twice as long as those immediately anterior to them (fig. 71). Male: Gonapophyses with scattered thorn-like setae (fig. 68D) dugesioides n. sp. Median setae of antescutellar row three to four times as long as the mesonotal setae anterior to them (fig. 72). Male: Gonapophyses with fine normal setae (fig. 68C) . Host doubtful furmani n. sp. 23. Median setae of antescutellar row scarcely longer than the discal scutal setae anterior to them (fig. 69) 24 Median setae of antescutellar row distinctly longer than the scutal setae anterior to them (figs. 66B, 70) 25 24. Prescutal setae rather evenly distributed and becoming gradually longer laterally and anteriorly, the minute discal setae extending anteriorly beyond the middle (fig. 69B). Male: Sternum VI absent. Host: Macrophyllum macrophyllum macrophylli n. sp. Prescutal setae becoming rather abruptly longer laterally and anteriorly, the median discal area of denser minute setae extending to about middle (fig. 69 A). Male : Sternum VI present. Host : Glossophaga soricina dugesii Townsend 25. Size larger, 1.46-2.17 mm. long. Antero-lateral setae of prescutum very long (fig. 66B). Male: Apices of gonapophyses distinctly curved ventrally (fig. 681). Host: Uroderma bilobatum urodermae n. sp. Size smaller, 1.21-1.7 mm. long. Antero-lateral setae of mesoprescutum long but not unusually so. Male: Gonapophyses (lateral view) nearly straight apically (fig. 68E). Hosts: Chiefly Carollia spp., joblingi n. sp. Trichobius pallidus group This group contains a single described species, T. pallidus, from British Guiana, taken from Furipterus horrens (fam. Furipteridae). Two ap- parently related, undescribed species are known to us from South America. One is from Amorphochilus schnablii, the only other species and genus of the family Furipteridae. In both of these undescribed species, the median mesonotal suture is simple, not bifurcate. We have seen only a single paratype of pallidus, and have not been able to satisfy ourselves that the median suture is truly bifurcate as in the caecus group, to which pallidus is obviously related. Interestingly, the Furipteridae are regarded as South American relatives of the Natalidae, which like the Chilonycterinae, are hosts of the caecus group. The pallidus group may be characterized as fol- lows: With the characters of the caecus group but differing in having multi-faceted eyes, the sixth sternum absent in the male, the tarsi shorter with the segments more compressed antero-posteriorly, and in lacking a ventral comb-like scale on the first tarsomere of the hind tarsi. Trichobius pallidus (Curran). Figure 53. Kesselia pallida Curran, 1934, Bull. Amer. Mus. Nat. Hist., 66: 522 — Kartabo, British Guiana, ex Furipterus horrens (American Museum of Natural History, N. Y.). Trichobius pallidus, Jobling, 1936, Parasitology, 30: 365 (keyed), 372 (redescr.), fig. 373; 1949, ibidem, 39: 317 (host). This species is known only from the type series, consisting of the holo- type and two paratypes, all males. The type host was collected in Panama, and it is possible that T. pallidus occurs there, too. 448 ECTOPARASITES OF PANAMA Trichobius caecus group The species of this group are apparently restricted to the Chilonycterinae (Phyllostomidae) and Natalidae. They probably should be regarded as a separate genus. The group may be characterized as follows : Head. — Dorsal surface of head not sub-divided into distinct laterovertices and oc- cipital plates; setae of occiput limited to lateral area; eyes a single large facet project- ing beyond lateral margins of head. Thorax. — High (deep), convex, anterior face Fig. 53. Trichobius pallidus (Curran). A, dorsal view. B, hindleg. C, wing. From Jobling (1938). oblique in profile, the upper edge projecting anteriorly above insertion of head. Meso- notum convex, the median suture short, bifurcate; anterior margin of prescutum rounded; setae of rather even size and distribution. Anterior margin of sternopleura angulate, feebly produced between the procoxae; posterior margin of pleurotrochantines straight; hind coxae nearly approximate. Wings. — Fourth and sixth longitudinal veins without setae near base. Legs. — First segment of hind tarsi with a ventral transverse comb-scale, inserted in a groove. Abdomen. — Female: Median dorsal connexivum not bare along entire mid-length, with at least two or more transverse rows of setae near apex; tergum VII and the "supra-anal plate" incompletely separated. Male: Sternum WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 449 V feebly sclerotized, transverse. Sternum VI very well developed, longest at middle and with distinctive marginal and submarginal setae along posterior edge. Gonapophyses symmetrical (in dorsal view), apices curved ventrally (in lateral view), paired ventral setae inserted at same level on each, the accessory seta of each pair situated anterior to the macroseta. The comb-scale on the first segment of the hind tarsi and the presence of marginal and submarginal setae on sternum VI (male) are characters not found in any other New World Streblidae. A well-developed male sixth sternum does not occur among any other Trichobiinae, but does in the Streb- linae, as defined here. It is of interest that the absence of setae along the base of the fourth longitudinal vein is also characteristic of the caecus group and of the Streblinae. Fig. 54. Thorax, dorsal view, Trichobius galei, new species, from Natalus stramineus mexicanus (no. 6731), San Lorenzo Caves (Panama). Trichobius galei Wenzel, new species. Figures 54, 57J-L. A small species, easily separated from the others of the group by the presence of only two rows of setae between the antescutellars and the transverse suture, the long, discal bristles on the seventh sternites of the female and on the hypopygium of the male, as well as by the short, very strongly bent male gonapophyses. DESCRIPTION: In general, with the characters of T. caecus excepting as follows: Thorax. — Anterior margin of prescutum scarcely projecting at middle, if at all; scutum at middle with only two transverse rows of setae between antescutellars and transverse suture. Wings. — Rs bare on basal half or slightly less, at least twice as long as distance from fork to crossvein r-m. Abdomen. — Lateral lobes of syntergum I+II with 9-13 coarse setae and below these usually several shorter, finer setae. Female: Tergum VII with only a pair of macrosetae; supra-anal plate with three much longer, apical macro- setae and a pair of lateral setae (one on each side) which are about as long as those 450 ECTOPARASITES OF PANAMA on tergum VII ; seventh sternites with all setae well developed, four or more macrosetae near apical margin ; postgenital sclerite as in fig. 57K. Male : Hypopygium covered with long setae dorsally and apically, with shorter setae ventrally ; gonapophyses as in fig. 57L. Measurements: BL TL WL ww Male 1.18-1.30 0.47-0.54 1.04-1.07 0.44-0.52 Female 1.32-1.65 0.48-0.55 1.04-1.37 0.52-0.60 TYPE MATERIAL : Holotype male and allotype female (slides) from Nata- lus stramineus mexicanus (host no. 6731), San Lorenzo Caves, Fort Sher- man (Canal Zone), 15 March 1961, C. M. Keenan and V. J. Tipton. In the Fig. 55. Lateral lobe of tergum I+II, lateral view, species. B, T. caecus Edwards. A, Trichobius johnsonae, new collection of Chicago Natural History Museum. Paratypes. — Approxi- mately 107 (27 bats), same data as the holotype. From Natalies sp. : 2, Camp Chagres, Madden Dam (Canal Zone), 19 June 1963, GML. Paratypes to be deposited in the collections listed on p. 410. REMARKS: This species is apparently restricted to Natalus stramineus mexicanus. A closely related species is known to us from the same host in Guatemala and Mexico. T. galei n.sp. is named in honor of Dr. Nathan Gale, Canal Zone Veterinarian. Trichobius caecus Edwards. Figures 55B, 57A-C. Trichobius caecus Edwards, 1918, Ann. Mag. Nat. Hist., (9), 1: 424. — Guacharo Caves, Trinidad, from unknown host (British Museum Natural History). — Good- win and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122: 223. Trichobius caecus Edwards (part.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 14 (keyed), 16 (records). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., no. 155, p. 655. Curran, 1935, Amer. Mus. Novit., no. 765, p. 8 (keyed). Jobling [as coccus'], 1938, Parasitology, 30: 371; 1949, ibidem, 39: 316, 326. This species is known to us only from Trinidad. Dr. Harold Oldroyd of the British Museum (Natural History) informed us that the type series WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 451 of T. caecus Edwards consists of three female cotypes mounted on one slide and an additional cotype preserved in alcohol. He kindly forwarded the slide containing the three females. The specimens had been mounted in euparol, still soft after many years. The mount was crushed in transit despite very careful packing. Although two of the specimens were obviously in poor condition when mounted and all three suffered some damage, the characters could still be clearly seen. All were remounted in euparol. One, which suffered little damage, was mounted separately on the original slide and has been designated as the lectoholotype. We have not seen the fourth specimen and do not know whether it is a male or female. Our description of the male is based on other Trinidad specimens from Tamana Caves. Dr. Oldroyd also forwarded to us the specimens which Kessel (1925, p. 16) recorded as caecus from Pteronotus davyi from Dominica. These speci- mens, preserved in alcohol, are in extremely poor condition. From speci- mens we mounted on slides, it is evident that they represent a species very closely related to T. johnsonae n. sp., but better preserved and geograph- ically representative samples from Pteronotus davyi and related hosts will be needed in order to determine their status. Dr. B. Jobling (in litt.) has informed us that in his revision of the genus Trichobius (1938, p. 371) he redescribed T. caecus "from two specimens which have been compared with the type of caecus. One of these was col- lected by Dr. L. H. Dunn from 'Chilonycteris rubiginosa rubiginosa' ( = Pteronotus parnellii fuscus) in Chilibrillo Caves, Panama, and the other by R. G. Hoves from Natalus stramineus in Jamaica." Jobling's illustrations (fig. 56) are obviously of the Panamanian specimen. This is evidenced by the chaetotaxy of the mesonotum and of the apex of the female abdomen, as well as by the host. These correspond with the characters and host of T. yunkeri n.sp. Jobling's Jamaican specimen from Natalus stramineus is almost certainly T. galei n. sp. or a closely related new species. Although Jobling's description of caecus seems to be based on both yunkeri and galei (or a related species), it could for the most part apply equally well to any species of the caecus group. DESCRIPTION: Head. — Vertical, dorsal surface elevated in middle; theca pyriform, palpi horizontal; laterovertices and occipital lobes not completely differentiated as separate sclerotized plates, though they are partially separated laterally by a trans- verse membranous strip; setae of occiput at sides only. Thorax. — High (deep), the anterior face oblique in lateral view, the upper edge (prescutum) projecting above insertion of head; anterior margin of prescutum rounded, narrowly incised at middle, the median suture short, bifurcate, the transverse suture straight, complete; mesonotal setae rather evenly distributed, the setae much longer anteriorly than posteriorly; scutum with three or four irregular, transverse rows of bristles between the long antescutellars and the transverse suture; scutellar bristles unusually long, the median pair nearly as long as mesonotum. Sternopleura angulate be- tween the front coxae, only feebly produced; pleurotrochantines together relatively nar- row, the posterior margin straight, hind coxae nearly approximate. Suture between sternopleura and pleurotrochantines strongly angulate. Wings. — Costa and Ri united at a point midway between second and third crossveins ; costa with well-developed macro- setae near base, these gradually smaller apically; fourth and sixth longitudinal veins bare basally ; bases of Ri and fifth longitudinal vein each with three or four macrosetae ; R only a little longer than distance between fork and crossvein r-m (14.5:13). Legs. — 452 ECTOPARASITES OF PANAMA Subequal, of moderate length; front tibiae only slightly shorter than the hind tibiae, none of them with macrosetae; basitarsus of hind tarsi nearly twice as long as broad, with a ventral comb-scale. Abdomen. — Dorsal connexivum with at least two transverse rows of setae near apex, anterior to tergum VII, never completely bare along entire midline, even in gravid females. Lateral lobes of tergum I+II with from 9-11 stout, long bristles and three to five finer, shorter setae below them (lectoholotype with a total of 16 setae). Female: Fig. 56. Trichobius yunkeri, new species. A, male, dorsal view. B, labium. From Jobling (1938, as T. caecus). Seventh tergum typically with a macroseta on each side and two, rarely four (holotype), short setae between them, these sometimes absent. Supra-anal plate with three apical macrosetae and two pairs of short, lateral setae (one pair each side). Seventh sternites usually with very short setae on outer half (except apical margin), the setae distinctly longer near mesal edge; apical margin with three, sometimes four, macrosetae (lecto- holotype with longar setae on outer half on one sternite, as in yunkeri) . Male : Sternum V feebly sclerotized, transverse, not divided, setae uniform throughout. Sternum VI present, relatively long, with a median, rounded bulge, with marginal and submarginal setae along posterior edge; dorsal surface of hypopygium with short, dense setae, these somewhat longer laterally and ventrally; setae also somewhat longer on tergum WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 453 IX, and becoming more so apically, those along apical margin being macrosetae; gon- apophyses as in fig. 57C. Measurements: BL TL WL ww Male 1.48-1.51 0.54-0.55 1.36-1.43 0.55-0.70 Female 1.24-1.84 0.52-0.66 1.43-1.60 0.55-0.71 MATERIAL EXAMINED: Lectoholotype female and two lectoparatype fe- males from Guacharo Cave, TRINIDAD, from the collection of British Mu- seum (Natural History). From Tamana Caves, TRINIDAD, 1 male reared from pupa, together with case, 20 November 1957, T. H. G. Aitken, col- lector [TVL] ; 2 males, 3 females, same locality, from "Chilonycteris rubi- ginosa" (Pteronotus parnellii), 11 November 1954, C. Colin Sanborn [CNHM] ; 11 males (4 too badly damaged to preserve, except genital mounts on slides) and 6 females, from same host, Guanapo Heights cave, TRINIDAD, 9 September 1954, C. C. Sanborn [CNHM]. Trichobius yunkeri Wenzel, new species. Figures 56, 57D-F. Trichobius caecus (not Edwards, 1918), Jobling, 1938, Parasitology, 30: 358, 364, 371 (part.), fig. 4. Bequaert, 194"0, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418. Jobling, 1949, loc. cit, 39: 316 (part.), fig. 1A. Closely related to T. caecus Edwards, but separated from it by the characters given in the key, notably the shape of the female postgenital sclerite (fig. 57E), the presence of five or six (instead of three or four) macrosetae on the female seventh sternites, and the shape of the male gonapophyses (fig. 57F) . DESCRIPTION: With the characters of T. caecus except as follows: Male: Gon- apophyses (fig. 57 F) longer, more slender, and more angulately bent at apex than in T. caecus (fig. 57C). Female: Median pair (sometimes three) of setae on tergum VII usually well developed (minute or absent in caecus), usually about half as long as the macrosetae lateral to them, sometimes with an additional seta lateral to one of the macrosetae ; supra-anal plate with three apical and two or three short, stout lateral setae on each side. Seventh sternites covered with short setae which become longer near mesal edge ; apical margin typically with five or six macrosetae, rarely four. Postgenital sclerite (fig. 57E) wider than in caecus (fig. 57B). Measurements: BL TL WL ww Male 1.51-1.59 0.52-0.59 1.37-1.43 0.55-0.66 Female 1.54-1.96 0.56-0.69 1.54-1.59 0.65-0.77 TYPE MATERIAL : Holotype male (slide) from Pteronotus parnellii fuscus (host no. 4437) from a railroad culvert at Paraiso (Canal Zone) , 15 Septem- ber 1959, C. M. Keenan and V. J. Tipton, collectors. Allotype female (slide) , same data as the type but from host no. 4386, 16 September 1959. In the col- lection of Chicago Natural History Museum. Paratypes. — 370 specimens from Pteronotus p. fuscus as follows: 46 flies (18 bats), same data as the allotype and 42 (14 bats), same data as the holotype; 135 (40 bats), same data as holotype, but 1 December 1959; 19 (4 bats) , railroad culvert east of Summit Golf Club, 26 October 1959 ; 15 (7 bats) , mine shaft, Coco Plantation, Gamboa (Canal Zone), 25 Novem- ber 1959; 2 (1 bat), Galeta Island (Canal Zone), 19 November 1959; 22 (1 bat) , Madden Airstrip (Panama) , 3 October 1961 ; and 22 (1 bat) , same data but 23 May 1961; 6 (2 bats), Chilibrillo Caves (Panama), 28 October 1959; 22 (4 bats), Almirante (Bocas del Toro), 28 January to 1 February 454 ECTOPARASITES OF PANAMA / \ V B D H K Fig. 57. Terminalia of Trichobius caecus group, including (from left to right) female terminal cone, female postgenital sclerite, and left male gonapophysis (lateral view). A-C, Trichobius caecus Edwards. D-F, T. yunkeri new species. G-I, T. johnsonae, new species. J-L, T. galei, new species. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 455 I960; 13 (1 bat), Rio Chucunaque (Darien), 17 February 1958; 35 (6 bats), Guanico (Los Santos), 22 January, 13 February and 16 February 1962; 7 (2 bats), Cerro Hoya (Los Santos), 18 and 21 February 1962; 3 (3 bats), Buena Vista Caves (Colon), 3 September 1959; 7 (5 bats), same locality, 15 September 1959 and 16 (1 bat), 24 November 1959. Paratypes to be deposited in the collections listed on p. 410. OTHER MATERIAL EXAMINED (50 specimens) : From Lonchorhina aurita: 6 (2 bats) , 17 July and 25 November 1959. From Pteronotus suapurensis: 1 (1 bat) , Fort Clayton (Canal Zone) , 19 October 1960 ; 27 (1 bat) , Natural Bridge, Madden Dam (Canal Zone) , 27 August 1959 ; 1 (13 bats) , Chilibrillo Caves (Panama), 28 October 1959. From Sturnira ludovici (mist net) : 1 (1 bat), 11 March 1962. From Carollia perspicillata azteca: 3 (12 bats), railroad culvert east of Summit Golf Club (Canal Zone), 26 October 1959; 5 (22 bats) , Buena Vista Caves (Colon) , 3 September, 15 September and 24 November 1959. From Artibeus lituratus palmarum: 6(1 bat) , Cerro Azul (Panama), 25 January 1958, GML. We have also seen hundreds of specimens from Pteronotus parnellii from GUATEMALA, MEXICO, COLOMBIA and VENEZUELA, which are tentatively assigned to this species. REMARKS: Pteronotus p. fuscus seems to be the primary host of this species. However, T. yunkeri was found together with T. johnsonae in equal abundance on P. suapurensis at Natural Bridge, Madden Dam, and there is no indication in our records that P. p. fuscus occurred there with that host, though this may have been so. In all other instances excepting two, where a bat other than P. p. fuscus was the host of yunkeri, P. p. fuscus was also known to be present. The small numbers of yunkeri taken on them could indicate that they were either strays or contaminations. T. yunkeri and related species tend to occur in swarms in the roosting sites of the host, thus greatly increasing the chances for such stray associations. We believe that the record from Artibeus lituratus palmarum represents an error in host identification or association. Virtually all the specimens received from this locality and collecting trip have incorrect host associations. In the case of the specimens from Sturnira ludovici, the host bat was collected in a mist net. When bats are thus captured, their streblids often leave and land on other nearby bats in the net. The species referred to by Cooper (1941, pp. 126, 127) as caecus is probably T. yunkeri. Trichobius yunkeri n. sp. is named in honor of Dr. Conrad Yunker of the Rocky Mountain Laboratory (Hamilton, Montana) and the Middle American Research Unit, Canal Zone, Panama. Trichobius johnsonae Wenzel, new species. Figures 55A, 57G-I. Very closely related to and generally with the characters of T. caecus Edwards and T. yunkeri n. sp., but differing from them in having only six to nine (rather than 11-16) setae on the lateral lobes of syntergum I +11, the very minute setae of the female seventh sternites, and especially in the shape of the female postgenital sternite and the relatively heavy male gonapophyses. 456 ECTOPARASITES OF PANAMA DESCRIPTION: Lateral lobes of syntergum I+II (fig. 55A) with only six to nine coarse setae and sometimes several finer setae ventral to these. Wings. — Rg bare on about basal fourth, approximately twice as long as distance from fork to crossvein r-m (11.5:7). Female: Seventh sternites with extremely fine, short setae outwardly, these longer toward mesal margin. Postgenital sclerite narrow and heavy (fig. 57H). Male: Gonapophyses as in fig. 57 I. Measurements: BL TL WL ww Male 1.59-1.70 0.45-0.58 1.45-1.48 0.71-0.73 Female 1.63-1.86 0.52-0.65 1.59-1.65 0.77-0.80 TYPE MATERIAL: Holotype male and allotype female (slides) from Pteronotus psilotis (host no. 8772) Penonome Cave (Code), 15 Decem- ber 1961, C. M. Keenan and V. J. Tipton, collectors. In the collection of Chicago Natural History Museum. Paratypes (278 specimens). — From Pteronotus psilotis: 66 (11 lots), same data as the holotype; 2, same data but 24 January 1962; 10 (1 lot), Armila (San Bias), 13 March 1963. From Pteronotus suapurensis: 26 (1 lot), Madden Dam (Canal Zone), 27 August 1959; 3 (1 bat), Casa Tilley, Cerro Punta (Chiriqui) , 6 March 1962 ; 81 (7 lots) , Penonome Cave (Code) , 15 December 1961; 62 (4 lots), Chilibrillo Caves (Panama), 28 October 1959, and 20 (1 lot), same locality, 8 March 1960; 8 (2 lots), Armila (San Bias) , 28 February 1963. From wall of cave: 461 (4 lots) , Penonome Cave (Code), 15 December 1961. The San Bias specimens were collected by C. 0. Handley, Jr. and F. M. Greenwell. Paratypes to be deposited in the collections listed on p. 410. OTHER MATERIAL EXAMINED: 1 specimen from Carollia perspicillata azteca, Penonome Cave (Code), 15 December 1961; 1 from Lonchophylla robusta, Chilibrillo Caves (Panama), 2 August 1960. REMARKS : We have seen a series of nine specimens, without host, labeled "Dominica, H. S. B.," sent to us by Dr. H. Oldroyd of the British Museum (Natural History). These had been determined by Kessel (1925) as T. caecus, but appear to be johnsonae or a new species closely related to it. They have an even heavier postgenital sclerite than do our specimens of johnsonae from Panama. We also have three lots, consisting of three males and two females, taken from Pteronotus davyi fulvus in Guatemala, which appear to be closely related to johnsonae but possibly represent a distinct new species. Trichobius johnsonae is apparently restricted to Pteronotus suapur- ensis and P. psilotis in Panama. In most instances, both hosts were collected at the same localities. Nycterophilia fairchildi n.sp. occurs on the same hosts. The single records of johnsor^ae from Carollia p. azteca and Loncho- phylla robusta are obviously of strays or contaminations. In both instances, the collections were from sites where Pteronotus were present. T. johnsonae is named in honor of Dr. Phyllis Johnson, formerly of the Gorgas Memorial Laboratory, Panama, and now at the University of Cali- fornia at Irvine. Trichobius major group As defined below, this group contains seven described species namely, T. major Coquillet, hirsutulus Bequaert, corynorhini Cockerell, truncatus WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 457 Kessel, pseudotruncatus Jobling, sparsus Kessel, and sphaeronotus Jobling. The first three are closely related, the others are provisionally assigned to the group. T. sparsus Kessel is the only species which has been collected in Panama. The following characters will serve to define the group. Head. — Laterovertices and occipital lobes either not differentiated or very weakly sclerotized and poorly defined. Abdomen. — Female: Cerci united with the ventral arc of the terminal cone. Male : Sternum V divided into two sternites or with a longitudinal median groove which appears to divide it. Sternum VI absent. Gonapophyses with the accessory setae inserted anterior to the macrosetae. Trichobius sparsus Kessel. Figures 42, 58. Trichobius sparsus Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 15 (keyed), 17 (descr.), pi. 1. (fig. 7), pi. 2 (fig. 10) — Chilibrillo River, Panama ex "Chilonycteris rubi- ginosa fusca" (U. S. National Museum). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth. Wash., no. 155, p. 655. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479, fig. (Streblidae II. — 6) ; 1935, Amer. Mus. Novit., no. 765, p. 9 (keyed). Jobling, 1938, Parasitology, 30: 373-375, fig. 6. Bequaert, 1940, Rev. Acad. Colomb. Cien. Exact., Fis. y Nat., 3:418. Trichobius sparsus is easily separated from the other species that occur in Panama by the distinctive mesonotal chaetotaxy (fig. 58B). To the characters given by Jobling (loc. cit.) the following may be added. Head. — Palpi with setae on basal two-thirds of ventral surface, with microsetae on upper surface. Thorax. — Microsetae present along lateral margins of prescutum, posterior half of scutum, and anterior third of scutellum. Abdomen. — Tergum I+II with a median basal group of setae which extends obliquely posteriorly and ventrally on each side to join a cluster of about 14-16 setae on face of lateral lobe, anterior to the posterior cluster of macrosetae. Connexivum with four sets of minute segmental setae, the three anterior sets consisting of two pairs each (a pair on each side of middle), the posterior set a single pair. Female: Supra-anal plate with two apical macrosetae, two macrosetae at middle, a pair of shorter setae on each side at mid-length, and a short seta each side of middle at base. Seventh sternites large, each with about 21 bristles, including one macroseta that is nearly twice as long as next longest setae; connexivum with a long seta on each side, but not on margin, anterior to the seventh sternites. Male : Sternum V divided into two large transversely oval plates. Sternum VI absent. Gonapophyses (fig. 58C) straight in dorsal view, blade-like and moderately curved ventrally in lateral view; ventral macroseta very long and extending beyond apex of gonapophysis, the accessory seta inserted anterior to the macroseta; a row of minute submarginal setae along ventral margin distal to accessory seta, two or three on lateral face, and one close to dorsal edge near apex. Measurements: BL TL WL ww Male 1.65-2.01 0.60-0.75 1.60-1.87 0.66-0.88 Female 1.95-2.25 0.71-0.77 1.90-2.09 0.82-0.88 PANAMANIAN MATERIAL EXAMINED: 29 flies (13 lots) from 51 bats. From Pteronotus parnelliifuscus, 20 flies (10 lots) from 49 bats : 3 (2 bats) , Almirante (Bocas del Toro), 29 January and 1 February 1960; 1, railroad culvert, Paraiso (Canal Zone), 16 September 1959; 4 (40 bats), same site and locality, 1 December 1959; 1, railroad culvert E. of Summit (Canal Zone), 26 October 1959; 1, Barro Colorado Island (Canal Zone), 12 Janu- ary 1957 [USNM] ; 3 (2 bats), Guanico (Los Santos), 13 and 16 February 1962; 8 (3 bats), Cerro Hoya (Los Santos), 14, 18 and 21 February 1962. From Carollia perspicillata azteca: 1, Almirante (Bocas del Toro), 31 Jan- uary 1960. From Artibeus lituratus palmarum: 7, Cerro Azul (Panama), 25 January 1958, Eustorgio Mendez [GML] . 458 ECTOPARASITES OF PANAMA OTHER MATERIAL EXAMINED : The only other specimens we have seen were collected from Pteronotus parnellii fuscus in GUATEMALA. The type of sparsus was collected from "Chilonycteris rubiginosa rubiginosa" (=P. parnellii fuscus of this paper) . We believe this to be the B D Fig. 58. Trichobius sparsus Kessel. A, underside of head and anterior part of thorax. B, dorsal view, female (wings removed). C, left male gonapophysis, lateral view. D, apex of abdomen, female, ventral view. A, B, D from Jobling (1938). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 459 principal if not the only host of this species. The specimens listed for Artibeus lituratus from Cerro Azul were certainly labeled incorrectly. The vial in which they were contained included flies from at least seven differ- ent host species, including Pteronotus p. fuscus. Field records show that six of these host species were collected at that locality on the same date. None of the flies were parasites of Artibeus. The record from Carollia probably represents a stray. Trichobius uniformis group The species of this group have the following characters in common : Small species, body 1.11-1.62 mm. long. Head. — With well-differentiated latero- vertices and occipital plates, the laterovertices transverse, the occipital plates oblique; setae of laterovertices, genal and postgenal margins, and occipital plates borne on tubercles, the tubercles along margins of postgenae and occipital lobes very distinct. Eyes small, with about six facets. Thorax. — Setae of mesonotum relatively uniform in size and distribution, somewhat larger and sparser anteriorly; median mesonotal suture complete or nearly so, usually united with the transverse mesonotal suture which is usually complete though sometimes indistinct at middle. Sternopleura bluntly, not strongly produced anteriorly ; posterior margin of pleurotrochantines slightly emarginate at middle, without a lobe. Wings. — All veins with macrosetae basally, those of Ri ex- tending a little more than halfway to apex of vein ; setae of costa and Ri borne on small tubercles, the setae becoming shorter apically toward junction; sixth longitudinal vein with setae nearly to base. Legs. — Short, the hind ones only slightly larger than fore- and midlegs. Abdomen. — Female: Tergum VII very feebly sclerotized, small, its setae minute. Cerci united with ventral arc of terminal cone. Male: Sternum V complete, undivided; VI present, very narrow. Gonapophyses with accessory seta inserted an- terior to the macroseta. Trichobius uniformis Curran. Figures GOB, 61B. Trichobius uniformis Curran, 1935 (part.), Amer. Mus. Novit., no. 765, pp. 9 (keyed), 10 (descrip.), fig. 8. Paraiso (Canal Zone), Panama, ex Glossophaga s. leachii (American Museum of Natural History). — Jobling, 1938 (part.), Parasitology, 30:364 (hosts), 365 (keyed); Bequaert, 1940 (part.), Rev. Acad. Colomb. Cienc. Exact., Fis. y. Nat., 3: 418. Jobling, 1949 (part.), Parasitology, 39: 316 ff. (hosts). Starrett and de la Torre, 1964, Zoologica, 49 : 57. Curran's type series of uniformis included the holotype, allotype, and three paratypes from Glossophaga s. leachii and two paratypes from Lonchophylla robusta. The paratypes from L. robusta represent a distinct species, described below as T. lonchophyllae n. sp. T. uniformis may be distinguished by the characters given in the key. To the characters given by Curran (loc. cit.) and above, the following may be added : Head. — Eyes distinctly removed from lateral margins of head. Innermost seta of posterior margin of occipital plate borne on a distinct, posteriorly projecting tubercle. Thorax. — Anterior margin of mesonotum straight, median suture usually indistinct posteriorly, often not united with the transverse suture. Wings. — Ri parallel to costa for most of its length. Length of Rs approximately equal to distance between fork and crossvein r-m. Abdomen. — Female: Four minute setae of tergum VII arranged with the inner two smaller setae (occasionally absent) only slightly posterior to the outer pair, appearing as a transverse row. Male: Gonapophyses as in fig. 60B. Measurements: BL TL WL ww Male 1.18-1.24 0.43-0.47 0.95-1.18 0.44-0.51 Female 1.24-1.46 0.47-0.50 1.00-1.22 0.46-0.52 460 ECTOPARASITES OF PANAMA PANAMANIAN MATERIAL EXAMINED : 31 specimens (19 lots) from (19) Glossophaga soricina leachii: 2, Fort Davis (Canal Zone), 18 November 1959; 4, Fort Clayton (Canal Zone), 14 September and 13 October 1959; 2, Galeta Island (Canal Zone) , 19 November 1959 ; 1, Fort Kobbe Beach (Canal Zone), 16 November 1959; 4 (holotype, allotype and 2 female paratypes), Paraiso (Canal Zone) , 17 October 1930, L. H. Dunn [AMNH] ; 3, San Lorenzo Cave, Fort Sherman (Canal Zone) , 29 March 1960, and 2, cativo trees, Fort Fig. 59. Trichobius lonchophyllae, new species. A, dorsal view, male. B, labium. C, apex of abdomen, female, ventral view. From Jobling (1938, as T. uniformis). Sherman, 4 December 1959; 4, Rio Hato (Code), 22 October 1959; 8, Santa Clara (Code), 27 October 1959; 1, Cerro Hoya (Los Santos), 17 February 1962. From Artibeus j. jamaicensis: 1, Fort Clayton (Canal Zone), 19 October 1960. From Desmodus r. murinus: 1, Almirante (Bocas del Toro) , 24 January 1960. The specimens from A. jamaicensis and D. rotundus were taken from bats collected in mist nets. OTHER MATERIAL EXAMINED : Numerous specimens from GUATEMALA, COSTA RICA, BRITISH GUIANA, and PERU, all from Glossophaga soricina except the British Guiana specimens which are without host. REMARKS : Curran's figure (loc. cit.) of this species is inaccurate in many WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 461 respects, including the length of Rs. However, the shape of RI is clearly that of uniformis and agrees with the holotype. This species appears to be limited to Glossophaga soricina, though it is possible that it may occur on other species of the genus Glossophaga. Published records from other hosts probably represent contaminations, errors of host identification, or other related species of the uniformis group. Trichobius lonchophyllae Wenzel, new species. Figures 59, GOD, 61A. Trichobius uniformis (part., ex Lonchophylla robusta) Curran 1935, Amer. Mus. Novit., no. 765, pp. 9, 10. Jobling, 1938, Parasitology, 30: 380 (descr., fig.). Bequaert, 1940, Rev. Acad. Colomb. Cien. Ex., Fis. y Nat., 3: 418. Jobling, 1949 (part.), Parasitology, 39: 316 (hosts). Although Jobling's (1938) records of T. uniformis refer in part to Cur- ran's species, his description and figures are obviously based on specimens of lonchophyllae n. sp. Mr. Jobling informs us that "T. uniformis has been redescribed and illustrated from the two paratypes, a male and a female lent to me from the collection of the American Museum of Natural History." Through the courtesy of Dr. Paul Arnaud, the senior author has been able to re-examine the female paratype and determine it to be lonchophyllae. It differs from uniformis and related species by the characters given in the key. The following characters may be added to Jobling's description : Head. — Relatively broad, the eyes conspicuously separated from sides. Innermost small seta of posterior margin of occipital plate borne on a distinct, posteriorly directed tubercle. Thorax. — Anterior margin of prescutum usually slightly projecting at middle; median suture connected with the transverse mesonotal suture, the latter straight or feebly anteriorly arcuate and not sharply defined on about middle third of width. Wings. — Ri distinctly sinuate, the costal cell rather abruptly narrowed, but Ri sub- parallel to costa from sinuation to juncture with costa; Rs distinctly longer (5.5:3.5) than distance between fork and crossvein r-m. Abdomen. — Female: Four minute setae of seventh tergum arranged in two transverse rows. Seventh sternite very broad, more than twice as long as broad. Male: Gonapophyses as in fig. 60D. Measurements: BL TL WL ww Male 1.18-1.21 0.44-0.49 1.09-1.15 0.52-0.57 Female 1.43-1.62 0.49-0.52 1.22-1.23 0.56-0.57 TYPE MATERIAL: Holotype male and allotype female (on slides) from Lonchophylla r. robusta (host no. 5081), Buena Vista Caves (Colon), 24 No- vember 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — From Lonchophylla r. robusta: 5, same data as the type, and 1, same data but 3 September 1949; 5, Chilibrillo Caves, Chilibre (Pan- ama), 27 March 1958; 2, same locality but 20 October 1961, C. M. Keenan and R. L. Wenzel; 1 (paratype of T. uniformis Curran), same locality, 9 March 1933 (no. 16), L. H. Dunn [AMNH] ; 2, La Laguna (Darien), 2900 feet elevation, 2 June 1963, C. O. Handley, Jr.; 1, Armila (San Bias), 14 March 1963, C. 0. Handley, Jr.; 16, Isla Bastimentos (Bocas del Toro), 30 January to 5 February 1963, C. O. Handley, Jr. ; 3 (2 bats) , 23 and 31 Janu- ary, C. O. Handley, Jr. Paratypes to be deposited in Chicago Natural History Museum, the United States National Museum, and the Environ- mental Health Branch, USAFSC, at Corozal (Canal Zone). 462 ECTOPARASITES OF PANAMA OTHER MATERIAL EXAMINED : 2 males f rom Artibeus lituratus palmarum, Cerro Azul (Panama) 2000 feet elevation, 25 January 1958, GML. REMARKS : The characteristic shape of wing vein Rx and the broad seventh sternites of the female are clearly shown in the illustration (fig. 59) of uniformis Jobling, 1938 (not Curran). Fig. 60. Trichobius uniformis group. A-C, left male gonapophysis. A, T. keenani, new species, paratype from Micronycteris megalotis (no. 9208), Guanico (Los Santos). B, T. uniformis Curran, from Glossophaga soricina leachii (no. 4908), Santa Clara (Chiriqui). C, T. lionycteridis, new species, holotype. D, right gonapophysis, T. lonchophyllae, new species, holotype. Thirteen of the 14 lots, including 36 of the 38 specimens, are from Lonchophylla r. robusta. However, we believe it may occur on other spe- cies of the genus Lonchophylla, too. The record from Artibeus is probably in error. Trichobius keenani Wenzel, new species. Figure 60A. Distinguished from related species by the characters given in the key. DESCRIPTION : With the characters of the group and also the following. Head. — Not as broad as in other species of the group, the eyes projecting to or nearly to lateral mar- gins of head; innermost small seta on posterior margin of occipital plates not borne on a conspicuous, posteriorly directed tubercle. Thorax. — Anterior margin of prescutum straight, median suture usually united with the transverse suture, sometimes slightly ab- breviated. Wings. — Rs approximately as long as distance between fork and crossvein r-m. Abdomen. — Female: Four minute setae of seventh tergum arranged in a transverse row, the outer pair usually slightly anterior to the inner pair. Seventh sternites reniform, scarcely twice as broad as long. Male: Gonapophyses as in fig. 60A. WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 463 Fig. 61. Wing. A, Trichobius lonchophyllae, new species. B, T. uniformis Curran. Measurements : Male Female BL 1.12-1.18 1.11-1.36 TL 0.38-0.42 0.41-0.46 WL 0.99-1.02 1.10-1.19 ww 0.48-0.50 0.48-0.51 TYPE MATERIAL: Holotype male and allotype female (on slides) from Micronycteris nicefori (host no. 5544) , Almirante (Bocas del Toro) , 12 Feb- ruary 1960, V. J. Tipton and C. M. Keenan. In the collection of Chicago Natural History Museum. Paratypes. — From Micronycteris megalotis microtis: 2, Camp Pina (Canal Zone) , 1 April 1960 ; 3, culvert under Borinquen Highway near Em- pire Range (Canal Zone) , 24 October 1961, C. M. Keenan and R. L. Wenzel; 3, same host, Guanico (Los Santos), 27 January 1962, C. M. Keenan and 464 ECTOPARASITES OF PANAMA V. J. Tipton. Paratypes deposited in the collections of Chicago Natural His- tory Museum and the United States National Museum. OTHER MATERIAL EXAMINED: From (2) M. megalotis microtis: 5 (de- stroyed accidentally), 24 October 1961, C. M. Keenan and R. L. Wenzel. From Uroderma bilobatum: 1, Fort Clayton (Canal Zone), 28 November 1960. epm.3 Fig. 62. A-C, posterior portion of venter of thorax showing condition in respect to median pleurotrochantinal lobe (m.p.l.) : A, lobe united with metepimeron (epm. 3) as in Trichobius longipes (Rudow) ; B, lobe short and blunt, not united with metepimeron, as in T. joblingi, new species; C, lobe absent, as in T. dugesioides, new species. D, sternopleura, Trichobius brennani, new species. E, same, T. longipes. Trichobius lionycteridis Wenzel, new species. Figure 60C. Differing from related species by the characters given in the key. DESCRIPTION : With the characters given for the group and also the following. Head. — Eyes only slightly separated from lateral margins of head. Innermost small seta on posterior margin of occipital plate not borne on a distinct, posteriorly directed tubercle. Thorax. — Anterior margin of prescutum slightly projecting at middle, median suture con- nected with the transverse mesonotal suture. Wings. — Length of Rs equal to or slightly shorter than distance from fork to r-m. Abdomen. — Female: Minute setae of seventh tergum arranged in two transverse rows ; seventh sternite about twice as broad as long. Male : Gonapophyses as in fig. 60C. Measurements : Male Female BL 1.27 1.40 TL 0.44 0.49 WL 1.17 1.29 ww 0.54 0.62 TYPE MATERIAL: Holotype male from Lionycteris spurrelli, Armila (San Bias) , 19 March 1963, C. O. Handley, Jr., collector. In the collection of the Chicago Natural History Museum. Paratype, a female from the same host. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 465 east slope of the Andes, Quincemil, 13°15/S-70°35'W (Cuzco?), PERU, 2400 feet, 25 April 1947, John C. Pallister, collector. In the collection of the American Museum of Natural History. Trichobius longipes group Six species — T. bequaerti n. sp., dybasi n. sp., mendezi n. sp., longipes (Rudow) (=mixtus Curran), costalimai Guimaraes, and dunni n. sp. — are included in this group. All except T. dunni n. sp. occur on species of Phyl- lostomus and Tonatia, related genera of the subfamily Phyllostominae (fam- ily Phyllostomidae) . T. dunni was taken on Molossus bondae (family Molos- sidae). Several undescribed species that are related to dunni are known to us from other bats of the genus Molossus. The longipes group may be characterized as follows : Head. — Laterovertices and occipital plates of head well differentiated. Median suture of mesonotum not forked. Eyes with 10-12 facets. Thorax. — Prescutal setae at middle of disk posterior to the median suture, denser and shorter than the lateral setae; ante- scutellar setae usually only slightly, if at all longer, than setae anterior to them. Pleurotrochantinal lobe dorsally reflexed and united with the metepimeron by a bar. Wings. — Sixth longitudinal vein lacking setae basally. Abdomen. — Male: Sternum V broad, undivided. Sternum VI very short, cord-like; gonapophyses curved to the left (in dorsal view), the accessory setae inserted posterior to the macrosetae. Female: Tergum VII very small, transverse, and feebly differentiated, if sclerotized at all; typically with four small setae arranged in one or two rows ; postgenital sclerite not visible. Cerci free, not united with the ventral arc. Trichobius longipes (Rudow). Figures 62A, E; 63F; 64. Strebla longipes Rudow, 1871, Zeitschr. Ges. Naturw., (n.s.), 37:121; 1872, Ann. Mag. Nat. Hist. (4), 9: 407 — Without locality, from "Phyllostoma hastatum" (Zoologische Staatsinstitut und Zoologische Museum, Hamburg). Speiser, 1902, Zeitschr. Hymen. Dipt., Heft 3, p. 159 (synonymized with T. parasiticus Gervais, in error). Trichobius dugesii (not Townsend, 1891), Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 16, pi. 1, fig. 1. Jobling, 1938, Parasitology, 30:383, fig. 12. Bequaert, 1940, Rev. Acad. Colomb. Cien. Exact., Fis. y Nat., 3:418 (part.). Cooper, 1941, Yearb. Amer. Phil. Soc., 1941: 124, 126 (part.). Trichobius mixtus Curran, 1935, Amer. Mus. Novit., no. 765, pp. 9 (keyed), 10 (descr.), fig. 10 — Chilibrillo Caves, Panama, from Phyllostomus hastatus pana- mensis (American Museum of Natural History). Guimaraes (part.), 1938, Rev. Mus. Paulista, 23: 654 (descr.), 662 (keyed), figs. 1,2; 1941, Papeis Avulsos, 1: 215. Bequaert, 1942, Bol. Ent. Venez., 1: 87. Guimaraes, 1944, loc. cit., 6: 189-191, figs. 18, 19, pi. 1, figs. 2,3 (puparium). Jobling, 1949, Parasitology, 39:316 ff. (hosts), 326 (records), fig. 2A. Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122, art. 3, pp. 238, 240. Starrett and de la Torre, 1964, Zoologica, 49 : 56. New synonym. The unique female type of Strebla longipes Rudow which was originally mounted dry on a slide was kindly lent to us by Dr. Herbert Weidner of the Zoological Museum at Hamburg with permission to remount it. The speci- men had been flattened from the pressure of the slip. We first treated it with trisodium phosphate, then with KOH, and remounted it in Canada balsam. Though it is still not an ideal study specimen, it is possible to establish its identity with Trichobius mixtus Curran, which was treated by 466 Jobling (1938) as T. dugesii. To the description given by Jobling, we add the following : Head. — Eyes with ± 11 facets. Thorax. — Microsetae present along upper edges of anepisterna, margins of prescutum along the longitudinal membranous cleft, the meta- notum and the laterotergite. Median pleurotrochantinal lobe reflexed dorsally and united Fig. 63. Left male gonapophysis, Trichobius longipes group. A, T. bequaerti, new species, holotype. B, T. dybasi, new species, paratype from Tonatia silvicola (no. 11903), Armila (San Bias). C, T. dunni, new species, holotype. D, T. mendezi, new species, holotype. E, T. costalimai Guimaraes. F, T. longipes (Rudow), from Phyl- lostomus h. panamensis (no. 4644), Chepo Road (Panama). by a bar to the metepimeron (fig. 62A). Abdomen. — Anterior face of tergum I+II with a single seta. Dorsal connexivum with two pairs of minute segmental setae. Female: Lateral connexivum with a cluster of six to eight setae coarser and longer than the other connexival setae, posterior to the lateral lobes of tergum I+II, and with a latero- ventral subapical macroseta on each side, anterior to the seventh sternites. Tergum VII with a transverse row of four minute setae, the middle pair slightly posterior to the outer. Supra-anal plate with four apical macrosetae and a short seta at mid-length on each side. Sternum I+II setose in a broadly triangular area and on each posterior angle; setae of apical margin slightly longer than the discal setae, increasing in length toward WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 467 the sides. Seventh sternites with about 11-12 setae, basal setae short, apical setae long, one a macroseta that is markedly longer than the others. Male: Gonapophyses slightly asymmetrical, both sinuate to left (in dorsal view), feebly curved apically in lateral view (fig. 63F). Measurements: BL TL WL ww Male 1.76-2.13 0.70-0.71 1.29-1.54 0.73-0.77 Female 1.87-2.35 0.78-0.82 1.59-1.92 0.73-0.85 PANAMANIAN MATERIAL EXAMINED: A total of 361 specimens (50 lots) from more than 90 bats. From Phyllostomus hastatus panamensis, 344 flies (48 lots) as follows : 35 (5 bats) , Fort Kobbe (Canal Zone) , 9 October 1959 ; 49 (19 bats), Fort Sherman (Canal Zone), 4 to 6 April 1960 and 30 July 1959; 3 (2 bats), Chagres River Cave, below Natural Bridge, Madden Dam (Canal Zone), 28 September 1959; 35 (6 bats), Natural Bridge, Madden Dam, 31 August 1959, and 9 (4 bats) , same locality, 18 September 1959 ; 13, Farfan (Canal Zone), 19 December 1915, T Hallinan [AMNH] ; 4, Culebra (Canal Zone) , 29 October 1930, L. H. Dunn [AMNH] ; 9 (1 bat) , Camp Pina (Canal Zone), 29 January 1960; 1, Rio Chilibrillo (Panama), 27 August 1957; 26 (20 bats), Chilibrillo Caves, 17 July 1957, and 43 (10 bats), same locality, 28 October 1959; holotype, same locality, 4 August 1931, L. H. Dunn [AMNH]; 2, Pacora (Panama), 24 April 1957 [USNM] ; 9, Tapia (Panama), J. R. Chapin [MCZ]. 1 (1 bat), Almirante (Bocas del Toro), 29 January 1960. From Phyllostomus sp. : 1, "Panama," 19 May 1957 [USNM]. From a mixed collection of Phyllostomus and Pteronotus spp. : 1, Rio Chilibrillo (Panama) , 27 August 1957, R. M. Altman. From Artibeus jamaicensis jamaicensis: 11 (3 bats), Natural Bridge, Madden Dam (Canal Zone). From Carollia perspicillata azteca: 1, Miraflores (Canal Zone), Kenneth W. Cooper [MCZ]. Without host: 2, "Chiriqui," 21 December 1961, C. Yunker; 1, Chilibrillo Caves, 1 January 1916, T. Hallinan [AMNH] ; 1, host K-16, Barro Colorado Island (Canal Zone), 2 December 1956 [USNM]. OTHER MATERIAL EXAMINED : We have seen about 70 specimens (25 lots) of this species from Phyllostomus h. hastatus and P. h. panamensis, from BOLIVIA, PERU, COLOMBIA, VENEZUELA, SURINAM, TRINIDAD, COSTA RICA, and GUATEMALA. REMARKS : Although Phyllostomus hastatus appears to be its principal host, T. longipes has also been taken from Phyllostomus discolor in Trinidad (Jobling, 1949, p. 317). Guimaraes (1938, p. 654) has recorded it from P. hastatus from Minas Gerais and Sao Paulo, Brazil, but his record (loc. cit.) from "Hemiderma perspicillatum" may be of a different species, perhaps Trichobius joblingi n. sp. Hoffmann's record (1953 : 186) of T. mixtus from Yucatan (from Artibeus jamaicensis yucatanicus) is probably of an un- described species that is known to us from Artibeus, in Mexico, Guatemala, and the West Indies. In this undescribed species, which is very similar in appearance to longipes, the pleurotrochantinal lobe is not united with the metepimeron and the transverse mesonotal suture is strongly angulate. One of the distinctive characters of T. longipes is the pronounced sexual dimorphism in mesonotal chaetotaxy (fig. 64) . The three following species resemble longipes very closely in most characters, except as indicated in the key, but the mesonotal chaetotaxy is nearly identical in both sexes. 468 ECTOPARASITES OF PANAMA Fig. 64. Trichobius longipes (Rudow), thorax, dorsal view. A, male. B, female. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 469 Trichobius mendezi Wenzel, new species. Figures 63D, 65B. Very similar to T. longipes (Rudow) but smaller and in general with shorter setae, and differing in lacking a cluster of coarse connexival setae on each side behind the lateral lobes of tergum I +11 in the female and in the distinctive male gonapophyses. DESCRIPTION: Head. — Eyes with about 10 facets. Thorax. — Long prescutal setae, limited to sides, not extending inwardly to median suture anterior to the denser median discal setae; discal setae only slightly denser at middle, becoming gradually sparser laterally and anteriorly and not noticeably shorter than the setae on anterior half of prescutum on inner half of width. Median antescutellar scutal setae at middle about twice as long as the shortest scutal setae anterior to them. Abdomen. — Dorsal con- nexivum with three sets of minute segmental setae, the posterior set usually consisting of two pairs rather than one; connexivum posterior to lateral lobes of tergum I+II without cluster of coarser, longer setae. Female: Tergum VII with four very short setae, the anterior pair more widely separated and a little longer than the posterior pair. Supra-anal plate with four apical macrosetae and at mid-length on each side a shorter seta. Sternum I+II noticeably emarginate at middle, setae at apical margin only slightly longer than discal setae and not noticeably longer toward sides, extending around sides to about halfway to base. Seventh sternites with about 14 or 15 setae, the basal ones short, the apical ones long, two macrosetae conspicuously longer than the rest. Lateral con- nexivum, anterior to seventh sternites, with a conspicuous macroseta (as in joblingi etc.). Male: Sternum V with apical setae only a little longer than the discal setae, not conspicuously longer laterally. Gonapophyses (fig. 63D) very similar to those of longipes but smaller, more slender and curved apically. Measurements: BL TL WL ww Male 1.63-1.73 0.54-0.55 1.13-1.20 0.55-0.63 Female 1.59-1.71 0.55-0.62 1.15-1.32 0.60-0.69 TYPE MATERIAL: Holotype male from Tonatia minuta (host no. 11764), Armila (San Bias), 20 March 1963, C. O. Handley, Jr. and F. M. Greenwell, collectors; allotype female, same data (host no. 11916), but 26 March 1963. In the collection of Chicago Natural History Museum. Paratypes. — 2 males (2 bats) , same host as holotype, Sibube (Bocas del Toro) , 17 January 1963, C. 0. Handley, Jr.; 1 male, same host, Puerto Obaldia (San Bias), 3 April 1963, C. 0. Handley, Jr. and F. M. Greenwell ; 2 females, same host, Cerro Hoya (Los Santos), 26 February 1962, V. J. Tipton and C. M. Keenan. Paratypes to be deposited in the collections of the Chicago Natural History Museum and the United States National Museum, Washington, D. C. REMARKS : These and other data indicate that mendezi is restricted to Tonatia minuta. This species is named after Mr. Eustorgio Mendez of the Gorgas Memorial Laboratory in recognition of his contributions to our knowledge of the ectoparasites of Panama. Trichobius dybasi Wenzel, new species. Figures 63B, 65A. Very closely related to T. longipes (Rudow) and T. mendezi n. sp., and differing from both in that the median discal mesonotal setae are not abruptly denser than the lateral and anterior setae. As in mendezi, the fe- male lacks a cluster of conspicuously longer or coarser connexival setae on each side posterior to the lateral lobes of tergum I+II. DESCRIPTION: With the characters of T. longipes except as follows. Thorax. — Pre- scutal setae becoming only slightly longer anteriorly along median suture, gradually longer laterally; median discal area not well denned. Abdomen. — Female: Connexivum 470 ECTOPARASITES OF PANAMA Fig. 65. Thorax, dorsal view. A, Trichobius dybasi, new species, allotype. B, T. mendezi, new species, female paratype from Tonatia minuta (no. 11916), Armila (San Bias). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 471 without cluster of coarser setae on each side behind lateral lobes of tergum I+II. Sev- enth sternites with 11 setae. Male: Gonapophyses as in fig. 63 B; sinuate to the left in lateral view and appearing very strongly, downwardly curved apically. Measurements: BL TL WL ww Male 1.46-1.63 0.54-0.58 1.15-1.26 0.60-0.66 Female 1.58-1.77 0.60-0.63 1.26-1.40 0.60-0.71 TYPE MATERIAL : Holotype male and allotype female from Tonatia silvi- cola (host no. 11903), Armila (San Bias), 25 March 1963, C. O. Handley, Jr. and Francis M. Greenwell, collectors. In the collection of Chicago Natu- ral History Museum. Paratypes. — 1 male, 3 females, same data as the holo- type ; 1 male, 3 females, same data, but 17 March 1963 ; 2 males, 1 female, same data, but 22 March 1963 ; 2 males, 1 female, same host and collectors, Puerto Obaldia (San Bias), 3 April 1963; 1 female, same host, Sibube (Bocas del Toro), 22 January 1963, C. O. Handley, Jr. Paratypes to be deposited in the collections of Chicago Natural History Museum; United States National Museum; the Gorgas Memorial Laboratory (Panama) ; and the Environmental Health Branch, USAFSC, at Corozal (Canal Zone). REMARKS : This species is named after Mr. Henry Dybas, Associate Cura- tor of Insects, Chicago Natural History Museum, who earlier collaborated with the senior author in studying these flies. Trichobius bequaerti Wenzel, new species. Figures 63A, 66A. Very similar to T. longipes (Rudow), dybasi n.sp., and mendezi n.sp., but the patch of microsetae on each side of prescutum is not restricted to extreme lateral margin, the antero-median prescutal setae are as long as the laterals, and the male gonapophyses are heavier (fig. 63A) . DESCRIPTION (Male) : With the characters of the group and the following. Thorax. — Discal mesonotal setae denser at middle, the mesonotal setae much longer apically and laterally. Microsetae of postero-lateral angles of prescutum extending inwardly from margin to second longitudinal row of setae (from margin), and anteriorly from trans- verse suture to second transverse row of setae (from suture). Abdomen. — With four pairs of minute dorsal segmental connexival setae. Gonapophyses (fig. 63 A) not as strongly curved in lateral view and more wedge-shaped than in T. longipes. Measurements: BL TL WL WW Holotype male 1.65 0.58 1.40 0.71 TYPE MATERIAL: A unique holotype male (slide) from Tonatia bidens (host no. 5489) Almirante (Bocas del Toro), 28 January 1960, collected by V. J. Tipton and C. M. Keenan. In the collection of Chicago Natural History Museum. REMARKS : This species is named after Dr. Joseph C. Bequaert, formerly Curator of Insects at the Museum of Comparative Zoology, Harvard Uni- versity, in recognition of his outstanding contributions to our knowledge of the pupiparous Diptera. Trichobius costalimai Guimaraes. Figures 63E, 67B. Trichobius costalimai Guimaraes, 1937, Revista Mus. Paul., 23: 660 (descr.), 662 (keyed), pi. 3, fig. 10 — from undetermined bat, Recife, Pernambuco (type deposi- tory not given). Jobling, 1938, Parasitology, 30: 368, fig. 14B. Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122, art. 3, p. 238. 472 ECTOPARASITES OF PANAMA T. costalimai is easily identified by the very distinctive mesonotal chaeto- taxy (fig. 67B) . It is the only species of the longipes group known to us in which the underside of the thorax and the hind coxae are microsetose. In addition to the characters listed in the original description, the following may be given : Thorax. — Microsetose area of prescutum extending inwardly along transverse suture almost to fourth seta from lateral margin ; elsewhere on basal half extending beyond the second row of setae, and on anterior half about two-thirds the distance to median suture ; sternopleura and pleurotrochantines microsetose ; sternopleural projection pointed. Legs. — Hind coxae microsetose below. Femora and tibiae, as well as procoxae and trochanters of middle legs with transverse lines ; and with base of each seta on coxae and surrounded by a slightly elevated ring. Abdomen. — Inner dorsal margin of lateral lobes of tergum I+II without a row or cluster of fine setae between apical coarse setae and base ; one or two short setae on an- terior face. Connexivum with two sets of dorsal segmental setae, the anterior set a single pair, the posterior set consisting of two pairs. Setae of sternum II subequal. Female: Connexivum without a cluster of coarser, longer setae on each side posterior to lateral lobes of tergum I+II. Tergum VII with two pairs of short subequal setae. Seventh sternites with about 11 setae. Supra-anal plate very short, with four apical macrosetae, as well as a short seta on each side and another at mid-length anterior to outer macroseta. Male : Sternum V with setae of apical margin only slightly longer than on disk. Gonapo- physes as in fig. 63E. Measurements: BL TL WL ww Male 1.72-1.76 0.63-0.66 1.29-1.43 0.58-0.66 Female 1.76-1.77 0.68-0.71 1.51-1.54 0.71-0.74 PANAMANIAN MATERIAL EXAMINED: From Phyllostomus d. discolor, 19 (4 lots) as follows: 10, Rio Tuira (Darien), 10 March 1958, P. Galindo [GML] ; 2, Rio Mandinga (San Bias), 29 May 1957, P. Galindo [GML] ; 6, Armila (San Bias) , 1 April 1963, C. O. Handley, Jr. and F. M. Greenwell ; 1, Almirante (Bocas del Toro) , 31 January 1960. From Uroderma bilobatum: 1, Puerto Obaldia (San Bias), 3 April 1963, C. 0. Handley, Jr. and F. M. Greenwell. From Desmodus rotundus: 2, Guanico (Los Santos) , 26 January 1962. From Pteronotus p. fuscus: 1, Guanico (Los Santos), 22 January 1962. From Carollia p. azteca: 1, Guanico (Los Santos), 27 January 1962. From Artibeus 1. palmarum: 1, Puerto Obaldia, 4 April 1963, Handley and Greenwell; 2, Cerro Azul (Panama), 25 January 1958, E. Mendez [GML]. OTHER MATERIAL EXAMINED: About 428 (24 lots) additional specimens, off Phyllostomus discolor, from GUATEMALA, EL SALVADOR, COLOMBIA, VENEZUELA, SURINAM, TRINIDAD, and PERU. Of these, 18 lots (402 flies) were from Phyllostomus d. discolor and P. d. verrucosus. One lot each was taken from Phyllostomus elongatus and P. h. hastatus. The remaining specimens were without host or dubious ones such as "Didelphis" ( !) and Sturnira. Other streblids from the -Surinam lot labeled as from Sturnira were Strebla consocius n. sp., a parasite characteristic of P. h. hastatus. REMARKS : Phyllostomus discolor appears to be the principal host of costalimai. It is our opinion that the records from other hosts taken at Guanico and Cerro Azul, in Panama, are in error. As pointed out elsewhere, a substantial percentage of the specimens collected at these localities were obviously incorrectly associated. We suspect that most of these, as well as the specimens of costalimai from other hosts, were from bats collected with mist nets (see Collecting hosts, above). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 473 Fig. 66. Thorax, dorsal view. A, Trichobius bequaerti, new species, holotype. B, T. urodermae, new species, female paratype from Uroderma bilobatum (no. 6958), Rodman Naval Base (Canal Zone) . 474 ECTOPARASITES OF PANAMA Trichobius dunni Wenzel, new species. Figures 63C, 67A. Superficially resembling T. phyllostomae Kessel and T. longipes (Ru- dow) , but differing from these and all other described species of Trichobius by the large number of bristles (17-19, as opposed to 10-12) on the occipital lobes of the head and by the distinctive male gonapophyses. The female is unknown. DESCRIPTION (Male) : Head. — Occipital plates each with about 15 conspicuous setae and on inner posterior margin about three additional short, fine setae. Palpi horizontal, their ventral surface densely clothed with short fine setae. Eyes conspicuous, with 11-12 large facets. Thorax. — Mesonotum distinctly convex, the anterior margin outwardly rounded, feebly emarginate at middle; median suture extending posteriorly to slightly beyond middle; setae longest and rather evenly distributed anteriorly and along sides, shorter posteriorly along sides; median discal setae shorter and denser posterior to the median suture and on scutum; setae of antescutellar row two to three times as long as setae immediately anterior to them ; scutellum with four setae. Wings. — Costa with both long and shorter setae, the long setae becoming shorter and of uniform size about halfway to junction with Ra, the junction situated beyond level of third crossvein but not as far as midpoint between second and third crossveins ; base of Ri with two or three long setae, base of fifth longitudinal vein with two. Legs. — Relatively short, the femora progressively longer, the hind femora from one-fourth to one-third longer than profemora; femora clothed with both long and short setae. Tibiae subequal, their setae short, dense, rather uniform. Tarsi rather strongly compressed antero-posteriorly, the claws relatively small. Abdomen: Lateral lobes of tergum I+II with numerous long setae, their inner dor- sal margins each with one or two short setae. Connexivum dorso-laterally with moder- ately long setae anteriorly, becoming shorter apically and ventrally, the underside densely clothed with relatively short setae; three pairs of minute, dorsal, segmental setae present. Sternum V weakly sclerotized, the discal setae similar to those of ventral con- nexivum, but those along apical margin distinctly longer. Sternum VI very narrow, inconspicuous. Hypopygium clothed with long setae, these longest dorsally and apically, the apical ones being macrosetae. Gonapophyses as in fig. 63C. Measurements: BL TL WL ww Holotype male 1.84 0.72 1.54 0.69 TYPE MATERIAL : Holotype male from Molossus bondae, Rio Chucunaque (Darien) , L. H. Dunn (no. 841) . In the collection of the Museum of Com- parative Zoology at Harvard University. REMARKS : T. dunni belongs to a small complex of closely related, un- described species that are found on species of Molossus. The record of Trich- obius phyllostomae (Curran, 1934, p. 523; Jobling, 1938, p. 377) from Molossus obscurus was probably based on one of these undescribed species. They appear to be the only New World Streblidae that occur on Molossidae. We have seen other undescribed species of this group from Mexico (Molos- sus nigricans) , Guatemala (Molossus sp.), Venezuela (no host), Trinidad (Molossus rufus), and Brazil (no host). Although the female of dunni is not known, it will very likely resemble those of the related undescribed species. In these, the seventh sternites are relatively much larger than in longipes and have numerous dense macro- setae. Although dunni has more numerous occipital setae than is usual in the genus, these setae are not as long and conspicuous as in the related, undescribed species. Trichobius dunni is named after Major Lawrence H. Dunn, in recogni- tion of his important contributions to medical entomology. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 475 A B Fig. 67. Thorax, dorsal view. A, T. dunni, new species, holotype. B, T. costalimai Guimaraes. 476 ECTOPARASITES OF PANAMA Trichobius dugesii group The species of this group are closely related to and probably derived from those of the longipes group. They differ in that the median pleuro- trochantinal lobe, if present, is not united with the metepimeron and ster- num VI is absent in some species. As in the longipes group, the female cerci are not united with the ventral arc. The species may be segregated into two complexes. The median pleurotrochantinal lobe is present in one, absent in the other. Trichobius dugesii complex In the species of this complex, a blunt pleurotrochantinal lobe is present but is not united with the metepimeron and sternum VI is present in the males of all excepting T. macrophylli n. sp. The species occur on bats of four subfamilies of Phyllostomidae : T. dugesii Townsend on Glossophaga sor- icina (Glossophaginae) ; T. joblingi n. sp. on Carollia perspicillata (Carol- liinae) and sometimes on Trachops cirrhosus (Phyllostominae) with which perspicillata often roosts; T. macrophylli n. sp. on Macrophyllum macro- phyllum (Phyllostominae) ; T. urodermae n. sp. on Uroderma bilobatum (Stenoderminae) . Trichobius urodermae Wenzel, new species. Figures 66B, 681. Superficially resembling females of Trichobius longipes (Rudow) but differing markedly in that the median pleurotrochantinal lobe is not united with the metepimeron as in that species. From other species of the dugesii group that have a median pleurotrochantinal lobe, it may be distinguished by its larger size, the very long antero-lateral prescutal setae, and the dis- tinctive male gonapophyses. DESCRIPTION : Head. — Slightly wider than long. Laterovertices and occipital plates well differentiated, the occiput slightly elevated above the rest of the head. Eyes with about nine large convex facets. Palpi rounded-oval, ventral surface with short setae, upper surface microsetose. Theca pyriform. Thorax. — Anterior margin of prescutum slightly projecting at middle, the median suture incomplete, usually extending less than half the distance to transverse suture, the latter slightly angulate, not well-defined at middle. Mesonotum as illustrated (fig. 66B) ; with a median area of dense finer setae, separated from anterior margin by three rows of very long setae that are two to three times as long as median setae, and separated from lateral margin by two longitudinal rows of long setae ; antescutellar setae at least twice as long as the scutal setae anterior to them; microsetae present along dorsal edge of anepisternum and along thickened pigmented lateral margin of prescutum and scutum. Median pleurotrochantinal lobe well developed, blunt, translucent, reflexed dorsally. Wings. — Costa with long macro- setae basally, these becoming progressively shorter apically, more or less uniform from about mid-length to apex; base of R! usually with three macrosetae, united with costa at a point approximately midway between second and third crossvein. Legs. — Fore- and midlegs typical of longipes group. Apical patch of dense fine setae on inner edge of hind tibiae not well defined. Tarsi compressed antero-posteriorly; hind tarsi (not including claws) about half as long as tibiae. Abdomen. — Lateral lobes of tergum I+II each with lateral cluster of 12-15 coarse setae, the inner (upper) margin of each lobe, anterior to this cluster, with a row of one to three fine setae. Sternum I+II setose in a broad triangular area and along lateral margins near apex; the marginal setae slightly longer toward sides; posterior margin feebly emarginate. Female: Tergum VII a narrow transverse strip with a longer seta WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 477 Fig. 68. Male gonapophyses (left, except as indicated), lateral view, Trichobius dugesii group. A, T. parasiticus Gervais. B, T. diphyllae, new species, holotype. C, T. furmani, new species, host unknown, Vitoc Valley (Junin), PERU. D, T. dugesioides, new species, holotype. E, T. joblingi, new species, composite, but based chiefly on holotype. F, T. macrophylli, new species, paratype from M. macrophyllum (no. 4741), Fort Davis (Canal Zone). G, left, and H, right gonapophyses, T. dugesii Townsend, from Glossophaga s. leachii (no. 5027) Fort Kobbe Beach (Canal Zone). I, T. urodermae, new species, para- type from Uroderma bilobatum, Almirante (Bocas del Toro). 478 ECTOPARASITES OF PANAMA on each side and a shorter pair medially and slightly posteriorly. Supra-anal plate with four apical macrosetae and two shorter setae on each side, one of these often inserted ventrad. Sternum I+II with weakly developed microsetae. Seventh sternites with about 17 setae, the apical setae longest, one conspicuously longer than the others. Subapical, ventro-lateral macroseta of connexivum absent. Male: Sternum I+II with distinct short transverse rows of microsetae. Discal setae of sternum V gradually longer toward apex, those of the apical margin considerably longer than the discals, those toward sides at least twice as long as the median setae. Sternum VI thread-like, barely visible. Hypo- pygium with numerous coarse setae. Gonapophyses as in fig. 681 ; ventral macroseta not reaching apex, the accessory seta inserted immediately proximal to the macroseta; two or three smaller setae inserted above ventral pair, a row of extremely fine setae along ventral margin; dorsal margin with a seta at angulation and another minute one near apex. Measurements: BL TL WL ww Male 1.46-1.84 0.62-0.65 1.32-1.39 0.58-0.71 Female 1.62-2.17 0.66-0.71 1.52-1.59 0.66-0.77 TYPE MATERIAL: Holotype male (slide) from Uroderma bilobatum (host no. 5528) , Almirante (Bocas del Toro) , 5 February 1960, C. M. Keenan and V. J. Tipton. Allotype female, same host (no. 6958) and collectors, Rodman Naval Base (Canal Zone), 27 April 1961. In the collection of Chicago Natural History Museum. Paratypes from Uroderma bilobatum, 22 (13 lots from 13 bats) as fol- lows: 2 (2 bats), type locality, 26 and 28 January 1960; 8, Fort Clayton (Canal Zone), 28 November 1960; 1, Fort Davis (Canal Zone), 22 Septem- ber 1960; 2 (2 bats), palm tree, Fort Randolph (Canal Zone), 6 October 1959; 1, Summit Gardens (Canal Zone), 11 September 1959; 2, Punta Pina (Darien), 24 March 1960; 6 (3 bats), Armila (San Bias), 18 March to 1 April 1963, C. 0. Handley, Jr. and F. M. Greenwell. Paratypes to be de- posited in the collections of Chicago Natural History Museum ; the United States National Museum; the Gorgas Memorial Laboratory, Panama; the Environmental Health Branch (U. S. Army) at Corozal (Canal Zone) ; Departamento de Zoologia, Secretaria de Agricultura, Sao Paulo ; and Facul- dad de Ciencias, Universidad Central de Venezuela, Caracas. REMARKS : This species is known to us only from Uroderma bilobatum and is one of the relatively few species of Trichobius known from bats of the subfamily Stenoderminae. Trichobius dugesii Townsend. Figures 68G,H ; 69A. Trichobius dugesii Townsend, 1891, Ent. News, 2: 106 — Guanajuato, Mexico, ex Glossophaga soricina (Snow Entomological Museum, University of Kansas, Law- rence) . Van der Wulp, 1903, Biol. Centr.-Amer., Ins. Dipt, 2: 432. Aldrich, 1905, Smiths. Misc. Coll., 46: 657 (part.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 14, 15 (part.). Stiles and Nolan, 1931, Bull. Nat. Inst.Hlth.Wash.,no. 155, p. 655 (part.). Guimaraes, 1938, Revista Mus. Paulista 23: 656 (part.); 1941, Papeis Avulsos, 1: 213-214 (part.). Schuurmans Stekhoven Jr., 1941, Zool. Anz., 136: 79. Bequaert, 1942, Bol. Ent. Venez., 1: 87 (part.). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 181, 186 (part.). Jobling, 1949, Parasitology, 39: 316 ff., 326 (part.). Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122: 245. Trichobius blandus Curran, 1935, Amer. Mus. Novit., no. 765, pp. 9 (keyed), 10 (descr.), fig. 11 — Paraiso [Canal Zone], Panama, ex Glossophaga soricina leachii (American Museum of Natural History). Jobling, 1938, Parasitology, 30: 358 (part., ex G. soricina). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 479 Fis. y Nat., 3: 318 (part.). Schuurmans Stekhoven Jr., 1951 (and 1941?), Beitrage Fauna Perus, 3: 95. At least six species have been confused under this name in the literature. Among these are T. joblingi n. sp., T. dugesioides n. sp., T. longipes (Rudow) (=: T. mixtus Curran) and several undescribed species. It is nearly impossible to determine which published references should be assigned to this name. As indicated elsewhere, the species treated as dugesii by Job- ling (1938) is T. longipes, while the species figured and described by him as T. blandus (not Curran, 1935) is T. joblingi n. sp. However, Jobling's records (loc. cit.) of blandus from Glossophaga soricina apply to dugesii. Although closely related to some of the species listed above, dugesii is easily distinguished by the combination of: extremely short antescutellar setae (fig. 69 A) ; its translucent, rounded, median pleurotrochantinal lobe; the distinctive gonapophyses and the presence of sternum VI, in the male. PANAMANIAN MATERIAL EXAMINED : A total of 109 specimens in 63 lots. From Glossophaga soricina leachii, 103 flies (59 lots) from 63 bats, as fol- lows: 1, San Lorenzo Cave, Fort Sherman (Canal Zone), 29 March 1960; 2, Fort Sherman, 4 December 1959; 1, Galeta Island (Canal Zone), 19 No- vember 1959; 4, France Field (Canal Zone), 30 November 1959; 5, Fort Kobbe Beach (Canal Zone), 16 November 1959; 8, Coco Solo ammunition area (Canal Zone), 20 October 1959; 3, Empire Range (Canal Zone), 30 September 1959; 5, Fort Clayton (Canal Zone) 14 September and 5, same locality, 13 October 1959; 4, Fort Davis (Canal Zone), 18 November 1959 and 2, same locality, 20 September 1960; 2, Camp Chagres, Madden Dam (Canal Zone), 25 June 1963; 1 [holotype of T. blandus Curran], Paraiso (Canal Zone), 17 October 1930, L. H. Dunn #1 [AMNH] ; 4, Chepo (Pan- ama), 24 May 1960; 2 (2 bats), Chilibrillo Caves (Panama), 28 October 1959 and 2 August 1960 ; 1, Huile (Panama) , ? 24 October 1960 ; 2, west side of Taboga Island (Panama), 24 September 1959; 3 [paratype females of T. blandus Curran], Bella Vista, near Panama (Panama), 25 June 1931, L. H. Dunn [AMNH] ; 9, Guanico (Los Santos), 28 January 1962; 1, Cerro Hoya (Los Santos), 10 February 1962; 1, Santa Clara (Code), 27 October 1959; 8, Rio Hato (Code), 22 October 1959; 18, Santa Clara (Code), 27 October 1959; 2, Rio Tuira (Darien), 10 March 1958, Pedro Galindo [GML] ; 7, Rio Mandinga (San Bias), 30 May 1957, P. Galindo [GML] ; 2, Almirante (Bocas del Toro) , 1 February 1960. From a mixed collection of (12) Carollia p. azteca and (6) Glossophaga s. leachii: 1, Huile (Panama), October 1960. From Carollia p. azteca (?) : 1, Gatun Tank Farm (Canal Zone), 19 November 1959. From Trachops cirrhosus: 6, Rio Mandinga (San Bias), 30 May 1957, P. Galindo [GML]. OTHER MATERIAL EXAMINED: From Glossophaga s. leachii: 2, Mineral Montechristo (Moragan), EL SALVADOR, 18 June 1953, H. Felten [SM] ; 2, near Santa Tecla (Libertad), EL SALVADOR, H. Felten [SM] ; 4, Finca El Zapote, Zapote (Escuintla), GUATEMALA, 7 July 1948, L. de la Torre and R. D. Mitchell, CNHM Guatemala Zoological Expedition (1948). From Glos- sophaga s. valens: 1, from preserved bat collected at Vitoc Valley, Tarma Province (Junin), PERU, by F. Woytkowski [CNHM]. 480 ECTOPARASITES OF PANAMA B Fig. 69. Thorax, dorsal view. A, Trichobius dugesii Townsend; B, Trichobius macro- phylli, new species; both from Macrophyllum macrophyllum (no. 4749), Fort Davis (Canal Zone). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 481 REMARKS: Trichobius dugesii appears to be restricted to Glossophaga soricina, though it is possible that it may occur on other species of Glos- sophaga. None were obtained from numerous specimens of G. commissarisi collected in Panama. It is possible that dugesii may occasionally transfer to other hosts, but there is no evidence that this is so. Those collected in Pan- ama from the mixed host series are probably from the specimens of Glos- sophaga. The specimens from Carollia p. azteca were doubtfully associated with the host by the collectors. Those taken from Trachops were a part of a collection that contained several obviously incorrect host associations, probably due to mislabeling. It is obvious that Guimaraes (1938), Bequaert (1942), Jobling (1938 and 1949), and Hoffmann (1953) treated dugesii, in part, but probably only the specimens they recorded from Glossophaga soricina. Guimaraes (loc. cit., pi. 2, fig. 6) figured as dugesii a species with long antescutellar setae, from Carollia perspicillata. It was probably T. joblingi n.sp. Trichobius joblingi Wenzel, new species, Figures 68E, 70. Trichobius dugesii (not Townsend 1891), Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 16, pi. 1, fig. 1 (part.?); Curran, 1935, Amer. Mus. Novit., no. 765, p. 9 (part.?). Bequaert, 1942, Bol. Ent. Venez., 1: 87 (part.?). Guimaraes, 1938, Rev. Mus. Paulista, 23: 656 (part.) pi. 2, fig. 6 (?). 1941, Papeis Avulsos, 1: 215 (part.). Cooper, 1941, Yearb. Amer. Phil. Soc., 1941: 124, 126 (part.?) Jobling, 1949, Para- sitology, 39: 316 ff., 326 (part.). Trichobius blandus (not Curran, 1935), Jobling, 1938, Parasitology, 30, pp. 365 (keyed), 385 (descr.), fig. 13. Bequaert, 1940, Rev. Acad. Colomb. Cienc. Ex., Fis. y Nat., 3: 418 (part.) Starrett and de la Torre, 1964, Zoologica, 49: 58. T. joblingi n.sp. is closely related to T. dugesii Townsend and T. mac- rophylli n.sp. From dugesii it may easily be distinguished by: the long antescutellar setae, which in joblingi are from two and one half to three times as long as the scutal setae immediately anterior to them (subequal to scutal setae in dugesii) ; the presence in the female, posterior to the lateral lobes of tergum I +11, of a cluster of three to five setae which are coarser and longer than the others (none conspicuously longer in dugesii) ; the shorter setae of tergum VII (female) ; and by characters of the male gonapophyses. From macrophylli it may be separated by its longer antescu- tellar setae (less than twice as long as anterior scutal setae in macrophylli) ; the much smaller eyes and relatively longer occipital plate, the eyes viewed from above being shorter than greatest length of occipital plate (as long as or longer than occipital plate in macrophylli) ; and by having only three to five coarser lateral connexival setae in the cluster behind tergum I +11 (10- 13 in macrophylli) . In the following description, we have added characters not given by Job- ling and emphasized others. DESCRIPTION: Laterovertices, occipital plates, upper surface of palpi, upper edges of mesepisterna, outer edges of prescutum on basal third, and outer edges of scutum microsetose, in addition to the metanotum and laterotergites, connexivum of abdomen, and the abdominal sclerites (excepting tergum I+II). Head. — Eyes relatively small, shorter (viewed from above) than greatest length of occipital plate; with ± 10 small facets. Thorax. — Median discal setae rather abruptly shorter and denser than those lateral and anterior to them, limited on prescutum to basal half or less and separated from the 482 ECTOPARASITES OF PANAMA lateral margins by about three rows of longer setae; antescutellar setae conspicuously and abruptly longer (usually two and one half to three times) than the scutal setae immediately anterior to them. Median pleurotrochantinal lobe rounded, translucent, slightly reflexed. Abdomen. — Inner (dorsal) margin of each lateral lobe of tergum I+II (anterior to the cluster of coarse setae) with two or three fine setae in a single row. Dorsal con- nexivum of abdomen with three pairs of minute "segmental" setae. Sternum I+II well developed, broader than pleurotrochantines, with nearly uniform setae arranged in a broadly triangular area and, along lateral margins on basal half. Female : Connexivum on each side posterior to lateral lobes of tergum I+II, with a cluster of from three to five setae which are conspicuously coarser and/or longer than the others; on each side anterior to the seventh sternite is a latero-ventral apically directed macroseta. Tergum VII feebly differentiated but represented by two pairs of minute setae, the anterior pair slightly more widely separated, the posterior pair close behind them. Supra-anal plate with four apical macrosetae and on each side at middle, a shorter seta; occasionally an additional short seta is present on each side medial to the lateral seta. Seventh sternites with about ten setae of varying lengths, the basal ones short, the apical ones macrosetae, one twice as long as next longest setae. Male : Sternum V feebly sclerotized, as wide as abdomen, its posterior margin broadly and feebly emarginate, the setae rather uniform throughout, except along posterior margin where five or six bristles on each side are about twice as long as the others. Sternum VI present, thread-like. Hypopygium bluntly coni- cal, with numerous macrosetae. Gonapophyses (viewed dorsally), asymmetrical and curved to the left; ventral margin appearing straight in lateral view (fig. 68E). Measurements: BL TL WL ww Male 1.21-1.48 0.47-0.49 1.00-1.07 0.45-0.55 Female 1.26-1.70 0.52-0.55 1.15-1.26 0.49-0.60 TYPE MATERIAL : Holotype male and allotype female (slides) from Carol- lia perspicillata azteca (host no. 4899) , railroad culvert east of Summit Golf Club (Canal Zone) , 26 October 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — 1870 specimens (334 lots, 614 bats) from Carollia perspic- illata azteca: 29 (10 bats) , Brazoa Brook (Canal Zone) , 6 October 1959 ; 56 (6 bats), Coco Solo (Canal Zone), 13 January 1960, and 2, same locality, 8 December 1960 ; 5 (8 bats) , Curundu (Canal Zone) , 14 September 1959 ; 105 (37 bats), Fort Davis, various dates, October and November 1959, and 14 (8 bats), same locality, various dates, September and November 1960; 2, Fort Clayton (Canal Zone) , 14 September 1959 ; 16 (28 bats) , Fort Gulick (Canal Zone), various dates, September and October 1959; Fort Randolph, 1 October 1959 ; 207 (174 bats) , Fort Sherman (Canal Zone) , various dates, March through December, 1959-61; 185 (67 bats), France Air Force Base (Canal Zone) , various dates, August to November 1959 ; 9 (3 bats) , Galeta Island (Canal Zone), 19 November 1959; 21 (8 bats), culvert, Gatun Tank Farm (Canal Zone) , 19 November 1959 ; 36, hollow tree, 1 mi. from Gatun- cillo (Canal Zone ?), 28 July 1960; 10, Farfan (Canal Zone), 19 December 1915, T. Hallinan ; 274 (51 bats) , mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 September and 25 November 1959; 12 (1 bat), Navy By- Pass Road (Canal Zone), 8 October 1959; 5 (2 bats), Navy Tank Farm, 6 October 1959 ; 6 (3 bats) , railroad culvert, Paraiso (Canal Zone) , 15 and 16 September 1959; 270 (17 bats), railroad culvert, east of Summit Golf Club (Canal Zone) , 26 October 1959 ; 46 (2 lots) , Summit (Canal Zone) , 17 April 1957, C. B. Koford [USNM] ; 106 (34 bats), Sardanillo Caves, Summit (Ca- WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 483 nal Zone) , 12 August 1961, C. Yunker [RML] ; 41 (9 bats) , culvert, Rousseau Road, 13 October 1959 ; 2, Pina Field range, 27 September 1960 ; 43, house, Camp Pina (Canal Zone), 26 October 1960; 17 (4 bats), Camp Chagres, Madden Dam (Canal Zone), 20 to 25 June 1963, GML; 41, Gamboa (Canal Zone), 1 May 1957 [USNM] ; 14 (3 lots), Barro Colorado Island (Canal Zone), 2 January, 19 February, and 5 April 1957 [USNM]; 2, Arraijan (Panama), 26 March 1961; 59 (27 bats), air-raid shelter, La Chorrera (Panama), 18 September 1959; 135 (17 bats), culverts, Chepo Road (Panama), 8 and 12 October 1959; 2 (2 bats), Chilibrillo Caves (Panama), 28 October 1959; 2, same locality, 2 August 1960; 17 (29 bats), same locality, 18 August 1959 ; 3, same locality, 27 August 1944 [USNM] ; 3, "Ft. Lorenzo, Panama" [USNM, det. as T. dugesii by Q. C. Kessel] ; 15 (5 bats), Isla Taboga (Panama), 24 September 1959; 61 (57 bats), various dates, September and November 1959, Buena Vista Caves (Colon) ; 1, culvert, Rio Hato Road (Code), 22 October 1959; 16 (4 bats), Puenta de Cocos, Isla del Rey (Darien), 22 and 23 March 1960; 16, Pina Point (Da- rien), 23 and 24 March 1960; 9, Rio Chucunaque (Darien), 19 February 1958, Pedro Galindo [GML] ; 9 (3 bats), Rio Tuira (Darien), 25 February to 2 March 1958; 1, Rio Seteganti, February 1961; 27 (11 bats), Guanico (Los Santos), 21 to 28 January 1962; 10 (8 bats), near Rio Guanico, Gua- nico, (Los Santos), 8 February to 1 March 1962; 6 (4 bats), Cerro Hoya, 10 to 25 February 1962 ; 80 (22 bats) , Almirante (Bocas del Toro) , 23 Janu- ary to 1 February 1960. Paratypes in the collections listed on p. 410. OTHER PANAMANIAN MATERIAL EXAMINED (272 specimens) : From Carol- lia castanea, 78 flies (23 lots from 27 bats) as follows: 25 (8 bats), Sibube (Bocas del Toro), 17 to 26 January 1963, C. 0. Handley, Jr.; 2, Rio Tuira (Darien) , 26 January 1958, P. Galindo [GML] ; 2, Rio Chucunaque (Darien) , 17 February 1958, P. Galindo [GML] ; 1, Tacarcuna (Darien), 19 June 1963, GML ; 4 (2 bats) , La Laguna (Darien) , 2900 feet elevation, 3 and 9 June 1963 [GML] ; 40 (15 bats), Armila (San Bias), 23 February to 29 March 1963, C. O. Handley, Jr. From Carollia subrufa, 44 flies (13 lots from 16 bats) as follows: 12 (2 bats), Sibube (Bocas del Toro), 22 and 23 January 1963, C. O. Handley, Jr.; 8 (1 bat), Isla Bastimentos (Bocas del Toro), 3 Feb- ruary 1963, C. O. Handley, Jr.; 26 (5 bats), Cayo Agua (Bocas del Toro), 12 and 16 February 1963, C. 0. Handley, Jr.; 19 (7 bats), Armila (San Bias), 25 February to 20 March 1963, C. 0. Handley, Jr.; 1, La Laguna (Darien), 2900 feet elevation, 29 May 1963, GML. From Carollia sp. : 1, Barro Colorado Island (Canal Zone) , 13 February 1957 [USNM] ; 7, Albrook Field (Canal Zone), 14 January 1963 [USNM]; 37, Cerro Tigre (Canal Zone), 15 January 1953 [USNM] ; 1, Rio Tuira (Darien), 25 February 1958, P. Galindo [GML]; 23, culvert, Chepo Road, near Pacora (Panama), 11 April 1961, C. Yunker [RML]. From mixed collections of Carollia p. azteca and Glossophaga s. leachii: 60, Huile (Panama), 24 October 1960; 21, San Lorenzo Cave, Fort Sherman (Canal Zone) , 15 March 1961. From Artibeus j. jamaicensis: 2, Almirante (Bocas del Toro) , 23 January and 18 February 1960 ; 3 (2 lots) , cave, west side of Taboga Island, 24 September 1959. From A. lituratus palmarum: 6, Cerro Azul (Panama), 25 January 1958, E. 484 ECTOPARASITES OF PANAMA Mendez [GML]. From Chiroderma villosum jesupi: 1, Almirante (Bocas del Toro), 23 January 1960. From Desmodus rotundus murinus: 1, Almirante (Bocas del Toro), 31 January 1960; 13 (2 bats), Guanico (Los Santos), 26 January and 1 February 1962. From "Dirias" : 2 (2 lots) , Summit (Canal Zone) 17 April 1957, C. B. Koford (USNM). From Glossophaga soricina leachii: 5 (4 bats), Almirante (Bocas del Toro), 26 to 30 January 1960; 9, Gamboa (Canal Zone), 17 April 1957, C. B. Koford [USNM]. From Lon- chophylla robusta: 4 (16 bats), Buena Vista Caves (Colon), 16 June 1960. From Lonchorhina aurita: 6(5 bats) , mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 September 1959; 19 (11 bats), same locality, 25 November 1959; 4 (2 bats), railroad culvert east of Summit Golf Club (Canal Zone) , 26 October 1959 ; 21, 2 miles north of Summit, 17 April 1957. From Macrophyllum macrophyllum: 6 (3 bats), Fort Gulick (Canal Zone), 15 September 1959; 1, Fort Davis (Canal Zone), dock 46, 13 October 1959. From Micronycteris nicefori [mist net] : 4 (2 bats), Armila (San Bias), 28 March 1963, C. 0. Handley, Jr. From Natalus stramineus mexicanus: 3, San Lorenzo Cave, Fort Sherman (Canal Zone), 15 March 1961. From Phyllostomus hastatus panamensis: 1, Natural Bridge, Madden Dam (Canal Zone), 31 August 1959; 9, Guanico (Los Santos), 26 January 1962; 3, Chilibrillo Caves (Panama), 17 July 1959; 1, Farfan (Canal Zone), 19 December 1915, T. Hallinan. From Pteronotus parnellii fuscus: 1, culvert, Paraiso (Canal Zone), 1 December 1959; 1, mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 25 November, 1959; 11 (5 bats), railroad culvert east of Sumit Golf Club (Canal Zone) ; 1 Buena Vista (Colon), 3 September 1959; 7 [ex "C.r. rubiginosa"], Chilibrillo River (Panama), 22 September 1923 [USNM]. From Tonatia silvicola: 1, Armila (San Bias), 22 March 1963, C. O. Handley, Jr. From Trachops c. cirrhosus: 14 (3 bats) , culvert, Chepo Road (Panama), 8 and 12 October 1959; 5 (2 bats), Rio Mandinga (San Bias), 30 May 1957, P. Galindo [GML]. From Uroderma bilobatum: 1, Fort Randolph (Canal Zone), 6 October 1959. From "bat": 3 (2 lots), Gamboa (Canal Zone), 12 December 1961; 3, Buena Vista (Colon), 3 September 1959; 5, Curundu (Canal Zone), 7 October 1960. Without specific host or locality: 15, from "culvert of RR" [USNM]. NON-PANAMANIAN MATERIAL EXAMINED : We have examined more than 500 additional specimens of joblingi, mostly from Carollia perspicillata, taken at various localities in GUATEMALA, BRITISH HONDURAS, EL SALVADOR, COSTA RICA, COLOMBIA, VENEZUELA, TRINIDAD, TOBAGO, SURINAM, BRAZIL and PERU. Only a few specimens were from hosts other than species of Carollia, including: 3 from Myotis sp. (( !) in Venezuela ; 1 from Rhogeesa io ( !) in Trinidad; 1 each from Lampronycteris platyceps, Glossophaga s. soricina, Desmodus r. rotundus and Phyllostomus h. hastatus in Trinidad ; and 3 obviously mislabeled specimens from Molossus major in Surinam. REMARKS: As noted above, T. joblingi n.sp. is the species figured by Jobling (1938) as T. blandus (not Curran 1935). Mr. Jobling informed us (in. litt.) that he redescribed blandus from two specimens collected by Dunn from Glossophaga soricina and Carollia perspicillata in the Canal Zone. His figure is of joblingi and not of dugesii Townsend (= blandus WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 485 Fig. 70. Trichobius joblingi, new species. Thorax, dorsal view. A, male from unidenti- fied discarded host (no. 4168), Buena Vista (Colon). B, female from Carollia p. azteca (no. 4628), Brazoa Brook (Canal Zone). 486 ECTOPARASITES OF PANAMA Curran) . However, his specimen from Glossophaga soricina was probably dugesii. Most published records of dugesii from Carollia perspicillata are actually based on specimens of joblingi. Jobling (op. cit.) described the posterior margin of sternum V as being "strongly emarginated in the middle." We have not found this to be true in the specimens examined by us, though idiosyncrasies of mounting may sometimes make it appear so in slide preparations. T. joblingi appears to be primarily a parasite of bats of the genus Carol- lia. Records from other hosts seem for the most part to represent transitory associations, disturbance transfers, and contaminations. Some represent errors of association or labeling. Temporary associations with a num- ber of hosts are to be expected of the parasites of a ubiquitous host like C. perspicillata which roosts with such a great variety of other bats. It is evi- dent from the field data that, in most instances, the other hosts from which joblingi was taken in Panama were roosting in the same sites as C. perspicillata. We do not believe that it normally occurs on these hosts (with the possible exception of Trachops) in the absence of Carollia. Carollia perspicillata is the most widely distributed, abundant and ubiq- uitous bat in Central and South America, and Trichobius joblingi n. sp. is the most abundant streblid parasite on that host. It constituted 74% of all Streblidae taken on perspicillata. It occurred on 76.7% of all individuals of this host collected and on 91.54% of all those parasitized by Streblidae. The average number of specimens taken per individual parasitized bat was 30 .o. Trichobius joblingi is named after Mr. B. Jobling, formerly of the Well- come Institute of Tropical Medicine, in recognition of his outstanding con- tributions to the systematics and morphology of blood-sucking Diptera, especially of the Streblidae. Trichobius macrophylli Wenzel, new species. Figures 68F, 69B. Very similar to T. joblingi n. sp. and T. dugesii Townsend and, like them, possessing a blunt translucent median pleurotrochantinal lobe, but differing in possessing a more convex and rounded mesothorax and much more evenly distributed mesonotal setae, which become longer much more gradu- ally towards sides and anterior margin. The antescutellar setae are inter- mediate in length between those of joblingi, and dugesii. In the female, posterior to lateral lobes of tergum I +11, there are 9-13 connexival setae that are coarser and longer than the others (three to five in joblingi, none conspicuously longer in dugesii) . DESCRIPTION : Nearly identical to joblingi but differing in the following characters : Head. — Eyes relatively large and conspicuous, as long as or longer than occipital plates (shorter in joblingi), with about 11 large convex facets. Thorax. — Mesonotal setae (fig. 69B), short at middle, becoming gradually longer toward sides and anterior margin; antescutellar setae less than twice as long as scutal setae immediately anterior to them. Median pleurotrochantinal lobe translucent, short, truncate or slightly emarginate at middle, feebly reflexed. Wings. — Rs, typically, slightly shorter than distance from fork to crossvein r-m. Abdomen. — Lateral lobes of tergum I+II with from two to five fine setae in a single row along inner (upper) margin. Dorsal connexivum with only one or two pairs of minute segmentally arranged setae. Female: Dorso-lateral and ventral WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 487 connexival setae subequal; immediately posterior to lateral lobes of tergum I +11, with a lateral cluster of from 9-13 setae which are conspicuously coarser and/or longer than the others ; a ventro-lateral subapical macroseta present. Tergum VII with one or two pairs of minute setae usually arranged in a transverse row, the inner pair sometimes slightly displaced posteriorly. Seventh sternites with 11-13 setae. Male: Dorso-lateral connexival setae longer than ventrals. Sternum V complete, the median apical marginal setae a little longer than the discals, those at sides conspicuously longer. Sternum VI absent. Gonapophyses as in fig. 68F. Measurements: BL TL WL ww Male 1.10-1.40 0.44-0.47 1.10-1.24 0.52-0.58 Female 1.10-1.57 0.47-0.55 1.26-1.40 0.55-0.66 TYPE MATERIAL (1185 flies in 46 lots from 79 Macrophyllum macro- phyllum) : Holotype, male and allotype female (slides) from host no. 4741, dock 45, Fort Davis (Canal Zone) , 13 October 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — 787 (33 bats), same data as the holotype; 295 (3 lots, 28 bats), same locality, 7 and 29 January 1960; 89 (14 bats), Natural Bridge, Madden Dam (Canal Zone), 31 August 1959; 12 (2 bats), Chepo Road, cul- vert, 12 October 1959; 2, Armila (San Bias), 23 February 1963, C. 0. Handley, Jr. and F. M. Greenwell. Paratypes deposited in the collections listed on page 410. OTHER MATERIAL EXAMINED: From Carollia perspicillata azteca: 1, Nat- ural Bridge, Madden Dam (Canal Zone) , 31 August 1959 ; 1, railroad culvert east of Summit Golf Club (Canal Zone), 26 October 1959. From Lon- chorhina aurita: 1 (9 bats), mine shaft, Coco Plantation, Gamboa Road (Canal Zone) , 25 November 1959. "Sobre Chiroptera" : 37, Lago de Valen- cia, VENEZUELA, 25 August 1948, J. Racenis [FCUCV] . REMARKS : This fly is apparently restricted to Macrophyllum macro- phyllum. The other two host records seem to represent strays or contamina- tions. Trichobius parasiticus complex In species of this complex, the pleurotrochantinal lobe is completely absent, sternum VI is absent in the male, and the male gonapophyses show greater modification in shape and chaetotaxy than in the dugesii complex. The mesonotal micropile is more extensive in several of the species than in any others of the dugesii group. Four species occur on Desmodidae, including T. diphyllae n. sp. from Diphylla ecaudata in Mexico and Guatemala and Venezuela ; an undescribed species from Diaemus youngii in Trinidad ; T. furmani n.sp., from Desmodus rotundus and Glossophaga soricina in Amazonian Colombia, Paraguay, and Peru ; and T. parasiticus, which appears to be primarily a parasite of Des- modus rotundus, from Mexico to Trinidad, Brazil and Peru. A fifth species, T. dugesioides — on Trachops cirrhosus, Chrotopterus auritus and Carollia perspicillata — is known to us from Guatemala to Panama. Within this complex, T. dugesioides has the least modified mesonotal chaetotaxy, while parasiticus and the undescribed species from Diaemus exhibit the greatest reduction of these setae. The male gonapophyses of parasiticus are the most highly modified of any known species of Trichobius. 488 ECTOPARASITES OF PANAMA Trichobius dugesioides Wenzel, new species. Figures 68D, 71. Very similar in apearance, though apparently not closely related, to T. joblingi n. sp., and often occurring together with that species on the same host. Unlike T. joblingi, it lacks a median pleurotrochantinal lobe and lacks sternum VI (in the male). It also differs in shape and chaetotaxy of the male gonapophyses and in the length of wing vein Rs in relation to the distance between fork and crossvein r-m. In dugesioides, the gonapophyses are straight (not sinuate) in dorsal view and have short thorn-like setae (very fine short hair-like setae in joblingi} . The mesonotal chaetotaxy dif- fers in that the antescutellar scutal setae are not as a rule abruptly longer than the scutal setae immediately anterior to them, as they are in joblingi. DESCRIPTION: Head. — Laterovertices and occipital plates well differentiated. Eyes with ± ten facets. Thorax. — Mesonotal chaetotaxy as shown in fig. 71, the antescutellar setae less than twice as long as the other scutal setae immediately anterior to them (two to three times as long in joblingi). Microsetae of prescutum extending inwardly from cleft to a little beyond first longitudinal row of setae along lateral margin. Pleurotro- chantines without apical median lobe, instead slightly emarginate at middle with a con- spicuous seta on each side of emargination. Wings. — Rs markedly shorter than distance between fork and crossvein r-m (subequal in joblingi). Abdomen. — Tergum I+II very similar to that of joblingi. Sternum I+II with setae of apical margin markedly longer toward the sides (only very slightly longer in joblingi). Dorsal connexivum with four pairs of minute dorsal segmental setae. Female: supra- anal plate with four apical macrosetae, without short marginal or discal setae (a short seta on each lateral margin in joblingi) ; seventh sternites with 13-15 bristles. Dorso- lateral and ventral connexival setae subequal, excepting a cluster of conspicuously longer setae behind the lateral lobes of tergum I+II. A long latero-ventral macroseta on each side anterior to the seventh sternites. Male : Dorso-lateral connexival setae longer than ventral connexival setae. Sternum V with apical setae longer than discals and becoming conspicuously longer toward sides. Sternum VI absent. Gonapophyses not sinuate in dorsal view, narrowly wedge-shaped in lateral view, the ventral margin nearly straight; with numerous short thorn-like setae ; accessory setae nearly half as long as macrosetae. Measurements: BL TL WL ww Male 1.26-1.40 0.49-0.55 1.07-1.15 0.52-0.60 Female 1.40-1.70 0.55-0.60 1.10-1.26 0.58-0.65 TYPE MATERIAL : 230 flies in 20 lots (35 bats), from Trachops c. cirrhosus. Holotype male, allotype female, Chepo Road (Panama), 12 October 1959, V. J. Tipton and C. M. Keenan. In the collection of Chicago Natural History Museum. Paratypes. — 105 (8 bats), same data but 8 and 12 October, and 94 (15 bats), 24 May 1960; 5 (3 bats), Cerro Hoya (Los Santos), 15 and 24 February 1962; 2, Rio Mandinga (San Bias), 30 May 1957, GML; 2 (2 bats), Armila (San Bias), 16 March 1963, C. O. Handley, Jr. and F. M. Greenwell. From Carollia perspicillata azteca* 124 specimens in 56 lots from 148 bats, as follows : 1, Camp Pina (Canal Zone) , 26 October 1960 ; 10 (6 bats) , air raid shelter, Fort Davis (Canal Zone), 6 October 1960, and 5 (7 bats), 29 October 1959; 1, same locality, 17 November 1960; 1, Fort Sherman (Canal Zone) , 18 October 1960 ; 17 (30 bats) , same locality, 2 and 4 Decem- ber 1959; 3 (2 bats), San Lorenzo Cave, Fort Sherman, 15 and 29 March 1960; 22 (16 bats), mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 September 1959; 16 (14 bats), same locality, 25 November 1959; 1, culvert, Brazoa Brook, Navy Tank Farm (Canal Zone), 6 October 1959; WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 489 B Fig. 71. Trichobius dugesioides, new species, thorax, dorsal view. A, male paratype (host no. 4639) and B, female paratype (host 4690), both from Trachops c. cirrhosus. Chepo Road (Panama). 490 ECTOPARASITES OF PANAMA 3, Juan Mina (Canal Zone) , 28 July 1960 ; 1, Pifia Field Range (Canal Zone) , 27 September 1960; 3 (2 bats), Camp Chagres [Boy Scout Camp], Madden Dam (Canal Zone), 20 June 1963 [GML] ; 1, Chilibrillo Caves (Panama), 2 August 1960 ; 6 (29 bats) , same locality, 18 August 1959 ; 1, culvert, Chepo Road (Panama), 12 October 1959; 4 (3 bats), Buena Vista (Colon), 3 and 15 September 1959; 11 (20 bats), same locality, 24 November 1959; 5 (3 bats), Cerro Hoya (Los Santos), 8, 18, and 23 February 1962; 2, Rio Seteganti (Darien), 5 February 1961; 2 (4 bats), Pina Point (Darien), 24 March 1960; 8 (3 bats), Almirante (Bocas del Toro), 28, 30, and 31 January 1960. Paratypes in the collections listed on p. 410. OTHER PANAMANIAN MATERIAL EXAMINED. — From Chrotopterus auritus: 5 (3 bats), Armila (San Bias), 7, 15 and 19 March 1963, C. O. Handley and F. M. Greenwell. From Lonchorhina aurita: 1, mine shaft, Coco Planta- tion, Gamboa Road (Canal Zone), 9 September 1959; 6 (19 bats), same locality, 25 March 1959; 4, Rancho Mojica (Bocas del Toro), 4800 feet elevation, 8 September 1961. From a bat doubtfully identified as Tonatia sp. : 6, Tacarcuna (Darien) , 4 September 1958, Pedro Galindo [GML] . From Natalus stramineus mexicanus: 1, Fort Sherman (Canal Zone), 17 March 1960. From mixed collections of (12) Carollia p. azteca and Glossophaga s. leachii: 1, Huile (Panama), 24 October 1960; of (3) C. p. azteca and (7) G. s. leachii: 2, San Lorenzo Cave, Fort Sherman (Canal Zone), 15 March 1961 ; of Natalus stramineus mexicanus and Carollia sp. : 1, same locality and date. We have also seen specimens from NICARAGUA and GUATEMALA, which we refer to as dugesioides. REMARKS : The records of dugesioides from hosts other than Trachops cirrhosus, Chrotopterus auritus, and Carollia perspicillata are probably in error or represent contaminations or transitory associations. In all likeli- hood, the true hosts are Trachops and Chrotopterus, closely related genera of Phyllostominae. T. dugesioides consistently occurs on Carollia perspicil- lata, which commonly roosts in the same places as Trachops, and it appears that these two bats "exchange" Trichobius to a considerable extent (see "Host-parasite Relationships," below.) On the other hand, dugesioides has not been taken from either C. castanea and only once from C. subrufa, species which are ecologically more restricted than perspicillata. The other streblids characteristic of perspicillata occur on all three hosts. Trichobius f urmani Wenzel, new species. Figures 68C, 72. Superficially very closely resembling joblingi, especially in mesonotal chaetotaxy ; differing from that species in that it lacks a median pleurotro- chantinal lobe, by the absence of sternum VI in the male, and the male gon- apophyses being straight rather than curved in dorsal view and long and feebly curved in lateral view rather than wedge-shaped. It may be easily separated from dugesioides, to which it is related, by the extremely long antescutellar bristles, some of which are three or four times as long as the scutal setae immediately anterior to them (less than twice as long in dugesioides) ; the extensive microsetose area on the prescutum; and the lack of thorn-like setae on the gonapophyses. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 491 Fig. 72. Trichobius furmani, new species, thorax, dorsal view. A, male. B, female. From Vitoc Valley (Junin), PERU (host unknown). 492 ECTOPARASITES OF PANAMA DESCRIPTION : Head. — As in joblingi and related species. Theca cordiform. Eyes with ± 11 facets. Thorax. — Prescutum, with anterior margin slightly projecting at mid- dle ; sides and anterior half with very long bristles, the discal setae short and denser and longer than discal scutal setae; microsetose area extending inwardly on prescutum to second (longitudinal) row of setae, excepting along transverse suture where it is feebly developed nearly to fourth row of setae; on anterior half of prescutum the microsetae extend inwardly about halfway to the median suture, but are so feebly developed as to be detectible only under oil immersion or very high "dry" magnification. Antescutellar scutal bristles very long, the longest ones two and one-half to four times as long as the setae anterior to them. Wings. — Rg shorter than distance from fork to crossvein r-m. Abdomen. — Tergum I+II with one to three short setae on anterior face on each side (between base and coarse setae of lateral lobe) ; inner dorsal margin of each lateral lobe with three to five fine setae in a single row. Abdominal connexivum with two pairs of dorsal segmental setae. Sternum I+II as in T. dugesioides. Female: Tergum VII with two pairs of setae; the outer, anterior pair a little less than half as long as apical macrosetae of supra-anal plate; the posterior pair close together, a little less than half as long as outer pair. Supra-anal plate with four apical macrosetae, without short setae on disk or lateral margins. Seventh sternites with about 15 setae. Connexivum with a ventro-lateral subapical macroseta on each side. Male: Sternum V as in T. dugesioides. Sternum VI absent. Gonapophyses (fig. 68C) long, straight in dorsal view, feebly curved apically in lateral view; ventral macrosetae very long, extending beyond apices of gon- apophyses; accessory setae very short. Measurements: BL TL WL ww Male 1.26-1.40 0.48-0.51 1.07-1.55 0.52-0.60 Female 1.40-1.70 0.52-0.56 1.10-1.26 0.58-0.65 TYPE MATERIAL: Holotype male and allotype female from Desmodus rotundus (Sanborn field no. 2942) , Segrario (Puno) , PERU, 13 October 1941, C. C. Sanborn. In the collection of Chicago Natural History Museum. Paratypes. — From Desmodus rotundus: 1, same data as the holotype. From specimens and wrappings containing a mixed collection of Desmodus rotundus, Phyllostomus hastatus, Carollia perspicillata, Glossophaga sori- cina, Tonatia silvicola and Vampyrops fumatus: 232, Vitoc Valley, Prov- ince Tarma (Junin), PERU, various dates, July to October 1940, F. Woytkowski. From Diphylla ecaudata: 4, Cerro de las Pinturas, upper Rio Iniride (Vaupes), COLOMBIA, 2-3 February 1957, Fred Medem. Paratypes to be deposited in the collections listed on p. 410. OTHER MATERIAL EXAMINED: From Glossophaga soricina soricina: 1, Riickenau, Friesland Colony near Itacurubi del Rosario, PARAGUAY, 14 June 1960, S. L. Loewen. REMARKS : Of the 232 specimens taken from the wrappings and speci- mens of the mixed lot of bats collected by F. Woytkowski, 220 are labeled as having been taken from the specimens of Desmodus rotundus. The records suggest that furmani may be a parasite of Desmodidae and may replace T. parasiticus on these bats in some parts of South America. Trichobius furmani is named after Prof. Deane Furman, University of California at Berkeley, in recognition of his outstanding contributions to the study of ectoparasites. Trichobius diphyllae Wenzel, new species. Figures 68B, 73A. Apparently related to dugesioides and furmani and like them, having the male gonapophyses straight in dorsal view. As in dugesioides the gon- WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 493 apophyses have short thorn-like setae in addition to the two pairs of ventral setae. Also distinguished from other species of the group by the very minute, fine, discal mesonotal setae as well as by the very extensive micro- setose area on the prescutum. DESCRIPTION: Head. — Broader than long (to base of occiput), with well-differen- tiated laterovertices and occipital lobes. Eyes with ± 10 facets. Palpi short, roundly- oval. Thorax. — Microsetose area of prescutum extending inwardly from longitudinal membranous fissure to third longitudinal row of setae on basal half, and about two-thirds the distance to median suture on anterior half. Disk of prescutum and scutum with extremely fine, short setae; with long sparse setae (fig. 73 A) laterally and anteriorly. Anterior projection of sternopleura truncate ; posterior margin of pleurotrochantines without a median reflexed lobe. Wings. — Length of Rs only slightly shorter than distance between fork and crossvein r-m. Abdomen. — Tergum I+II with a single short seta on anterior face; inner dorsal mar- gin with about five fine setae between base and cluster of coarse setae on lateral lobe. Sternum I+II, with setae of apical margin only slightly longer laterally than medially; postero-lateral angles with extensive setose area. Dorsal connexivum with four pairs of minute segmentally arranged setae. Female: Dorso-lateral connexival setae conspicu- ously shorter than the ventrals; lateral connexival setae behind the lobes of tergum I+II a little longer than those posteriorly but not forming a conspicuous cluster; a subapical ventro-lateral macroseta present on each side. Tergum VII variable in shape, longer than broad, with a longer seta along each lateral margin at about mid-length, and a pair of shorter subapical more closely spaced setae. Seventh sternites with about 15 setae, the apical ones macrosetae. Male: Lateral connexival setae of about same size as ventrals. Sternum V, at middle, with apical marginal setae distinctly longer than discal setae and becoming more than twice as long as the discals toward the sides. Sternum VI absent. Gonapophyses as in fig. 68B. Aedeagus rod-shaped, finely pointed at tip. Measurements: BL TL WL WW Male 1.48-1.62 0.48-0.54 1.17-1.32 0.60-0.63 Female 1.37-1.73 0.51-0.55 1.21-1.37 0.62-0.69 TYPE MATERIAL : Holotype and allotype from Diphylla ecaudata centralis (CNHM host nos. 73191-94, 73345-46), San Lorenzo, four miles northeast of Volcan de Jumay (Jalapa), GUATEMALA, elevation 5700-5750 feet, 15 February 1952, Luis de la Torre, CNHM Guatemala Zoological Expedition (1951-52). In the collection of Chicago Natural History Museum. Paratypes. — From Diphylla ecaudata centralis: 9, same data as the holo- type; 1, same data but 13 February 1952 ; 9, Finca Recreo, Municipio Yepo- capa (Chimaltenango) , GUATEMALA, elevation 4400 feet, 11 May 1948, R. L. Wenzel and Luis de la Torre, CNHM Guatemala Zoological Expedition (1948) ; 62, same data but 25 October 1948, R. D. Mitchell and L. de la Torre. From Diphylla ecaudata: 1, Pueblo Nuevo Xcan (Quintana Roo), Yucatan Peninsula, MEXICO, 29 July 1962, Ticul Alvarez ; 5, Felipe Carrillo Puerto (Quintana Roo), Yucatan Peninsula, MEXICO, 16 August 1962, J. Knox Jones, Jr. ; 7, El Limon, Rancho Grande (Aragua) , VENEZUELA, 30 March 1960, C. 0. Handley, Jr. [USNM]. Paratypes to be deposited in the collec- tions listed on page 410. REMARKS : This fly was not taken from the specimens of Diphylla ecau- data that were collected in Panama. However, it was taken in Venezuela, Guatemala, and Mexico, and thus it seems reasonable to expect that it will also be found in Panama, as was Strebla diphyllae n.sp. (q.v.) a character- istic parasite of the same host. 494 ECTOPARASITES OF PANAMA Trichobius parasiticus Gervais. Figures 68A, 73B. Trichobius parasiticus Gervais, 1844, Atlas Zool., p. 14, pi. 43. — "Guiane," from Des- modus rufus. — Costa Lima, 1921, Arch. Esc. Sup. Agric. Vet. Med., 5: 18-20 (part.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 14 (keyed), 15 (records). Speiser, 1900 (part.), Arch. Naturg., 66A, Bd. I, pp. 32 if. (morph., etc.), 59-60 (discuss., synonymy), 62 (cit), 65 (keyed), figs. 5-9; 1907, Ent. News, 18: 104 (cit.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., no. 155, p. 655. Curran, 1935, Amer. Mus. Novit., no. 765, pp. 8 (keyed), 11. Guimaraes, 1938, Rev. Mus. Paulista, 23: 657 (synonymy, discuss.), 662 (keyed), figs. 7, 8. Jobling, 1938, Parasitology, 30: 364 (hosts), 365 (keyed), 381 (redescr.), fig. 11. Mazza and Jorg, 1939, Mis. Estud. Pat. Reg. Argent., Publ. no. 41, pp. 87-93, figs. 1-3, 5-6 (morph.). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3:418. Jobling, 1949, loc. cit., 39: 316 ff. (hosts), 326 (records); 1951, Trans. Roy. Ent. Soc. Lond., 102: 215 (morph.). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 181 (keyed), 186 (rec- ords), pi. 4, fig. 4. Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122: 223, 269 (records). Trichobius kesseli Guimaraes, 1938, Rev. Mus. Paulista, 23: 660, pi. 3, fig. 9. Schuur- mans Stekhoven Jr., 1951 (and 1941?), Beitrage Fauna Perus, 3: 95 (part.?). Apparently the type of T. parasiticus no longer exists, but the interpreta- tion of Kessel (op. cit.), followed by Curran (1935), Guimaraes (1938), and Jobling (1938) is probably correct. The type and principal host, Des- modus rotundus (= Desmodus rufus) may carry another species, T. furmani n. sp. (q.v.) in some parts of South America, but the type of parasiticus was probably from a locality not far removed from the coast of "Guiane" and hence is almost certainly the species treated here. We have no speci- mens from French Guiana. When specimens are available, a neotype should be designated. As with several of the past authors, Speiser's concept (1900, p. 59) of T. parasiticus included several species, among them Strebla iviedemannii Kolenati 1862 ( =Kolenatia wiedemannii Rondani 1878, p. 169) and T. du- gesii Townsend. His specimens from Cuba could not possibly be parasiticus, nor could Trichobius parasiticus of Newstead (1909-1910, p. 468) nor Strebla wiedemannii Kolenati 1862, because neither parasiticus nor its hosts occur in the Greater Antilles. Speiser (1902, p. 159) later erroneously syn- onymized T. longipes (Rudow) under parasiticus. Costa Lima's figure (1921, pi. 1, fig. 1) of a wing of parasiticus probably applies to a species of the uniformis group, if the setae of the sixth longitudinal vein are correctly shown. The specimens identified as parasiticus by Bau (1929, p. 12) , Dunn (1929, p. 504) and Bequaert (1933) have not been re-examined. Bau's record is without locality or host data. Dunn's record does not include a host identifi- cation. Bequaert's specimens from Yocat, Yucatan were from Diphylla centralis and thus could have been either parasiticus or diphyllae n.sp., though we doubt that he could have mistaken diphyllae for parasiticus. T. parasiticus is easily recognized by the distinctive mesonotal chaeto- taxy (fig. 73B) and male gonapophyses (fig. 68 A) . To the characters given by Jobling (1938), the following may be added. Thorax. — Microsetose area of mesonotum extending from posterior margin of scutum to anterior margin of prescutum and inwardly to enclose the outer row of long prescutal setae and the setae of humeral callus. Abdomen. — Dorsal connexivum with four sets of WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 495 Fig. 73. Thorax, dorsal view. A, Trichobius parasiticus Gervais, male from Desmodus r. murinus (no. 9259) Guanico, (Los Santos). B, Trichobius diphyllae, new species, male paratype from Diphylla ecaudata centralis, Felipe Carrillo Puerto, Quintana Roo, MEXICO. 496 ECTOPARASITES OF PANAMA minute segmental setae, the anterior set consisting of two pairs, the three posterior sets of a single pair each. Upper inner margin of tergum I+II typically with two fine setae. Setae of sternum II extending anteriorly more than half the distance to base along lateral margins. Female: Tergum VII with two distal pairs of setae, the lateral pair coarse, about three times as long as inner pair and slightly anterior to them. Supra-anal plate with four apical macrosetae and, at mid-length, two to six short setae ; when only two short setae are present, they are situated on each side near lateral margin (six short setae noted in only one specimen, taken from Diaemus). Seventh sternites with 16-17 setae, two or three of them macrosetae, one much longer than the others. Connexivum pos- teriorly with a ventro-lateral macroseta on each side; a cluster of about 20 setae is present on each side behind lateral lobes of tergum I+II distinctly longer than other dorso-lateral and lateral connexival setae, which are shorter than ventral connexival setae. Male : Apical margin of sternum V with setae about three times as long as discal setae, except for median pair, which are about twice as long as discals. Measurements: BL TL WL ww Male 1.42-1.57 0.45-0.49 1.10-1.11 0.58-0.60 Female 1.52-1.70 0.52-0.55 1.22-1.29 0.60-0.63 PANAMANIAN MATERIAL EXAMINED. — From Desmodus rotundus muri- nus: 258 specimens in 41 lots from 74 bats, as follows : 24, Casa Lewis, near Cerro Punta (Chiriqui), 5 February 1960; 11 (8 bats), hollow tree, Casa Tilley, near Cerro Punta, 6 February 1962; 8 (24 bats), same locality, 23 January 1960; 23 (12 bats), same locality, 6 and 11 March 1962; 167 (11 bats) , Guanico (Los Santos) , 26 January to 25 February 1962 ; 13 (6 bats) , Almirante (Bocas del Toro), 23-31 January 1960; 11, Changuinola (Bocas del Toro), 27 February 1960; 25 (10 bats), Red Tank (Canal Zone), 27 January 1960; 1, Chilibrillo Caves (Panama), 2 May 1957 (USNM). From Diaemus youngii: 2, Armila (San Bias), 4 March 1963, C. 0. Handley, Jr. and F. M. Greenwell. From Phyllostomus hastatus: 1, Guanico (Los Santos), 26 January 1962. From "Vampire Bats": 1, Chilibrillo Caves (Panama), 14 July 1945. OTHER MATERIAL EXAMINED : Hundreds of specimens representing many lots from MEXICO, GUATEMALA, EL SALVADOR, COSTA RICA, COLOMBIA, VENE- ZUELA, SURINAM, TRINIDAD, BRAZIL and PERU. With few exceptions, these were taken from Desmodus rotundus, but parasiticus was taken together with T. diphyllae n.sp. on Diphylla ecaudata in Mexico and was found on Diaemus youngii in Panama. REMARKS : As noted, T. parasiticus is primarily a parasite of Desmodus rotundus though it may occur on other Desmodidae, too. Records from other bats almost certainly represent strays, contaminations, or transitory associations. Desmodus often roosts near other bats. Trichobius phyllostomae group We have assigned four species to this group. They are Trichobius phyllostomae Kessel, T. brennani n.sp., T. vampyropis n.sp., and an unde- scribed species. Of these, brennani and vampyropis occur in Panama. The other two are known to us only from Brazil. The group may be character- ized as follows : Head. — Laterovertices and occipital plates differentiated. Eyes multi-faceted, with from 26 to 32 facets. Thorax. — Anterior margin of prescutum distinctly projecting at middle. Median discal mesonotal setae dense, inserted in distinct thecae. Stenopleural WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 497 projection very strong, truncate. Median pleurotrochantinal lobe tapered and dorsally reflexed. Legs. — Relatively long; hind legs longest, the femora about as long as thorax; dorsal setae of tibiae conspicuous, two to three times as long as the other tibial setae. Abdomen. — Female: Cerci not united with the ventral arc. Male: Sternum V well de- veloped, with very long apical macrosetae, these progressively longer laterally. Sternum VI absent. Gonapophyses dissimilar, the macroseta and accessory seta inserted distinctly more distad on the left gonapophysis than on the right. Jobling (1938) referred T. phyllostomae to the group of species without well-differentiated laterovertices and occipital plates. In all our material of this group, these plates are differentiated, but are not as widely separated as in some other species. Trichobius brennani, new species. Figures 62D, 74A, 75A, B. Closely related to T. phyllostomae Kessel and differing from that species as follows : posterior margin of each occipital plate with only two minute setae rather than three or four; inner margin of lateral lobes of tergum I +11 with only one row of short setae instead of two irregular rows; tergum VII (female) with only a single pair of minute setae, rather than two pairs medial and apical to the macrosetae; male gonapophyses curved apically and with only a few submarginal microsetae (fig. 75 A, B), rather than wedge-shaped with a row of distinct marginal setae (fig. 75D). DESCRIPTION : Head. — Funnel-shaped, viewed from above, slightly elevated behind, with laterovertices and occipital lobes differentiated (sometimes not appearing so in slide mounts because of clearing and because of the presence of microsetae on the post- vertex and between the laterovertices and occipital lobes) ; laterovertices with seven bristles; occipital lobes with from 12-14 coarse setae and at middle of posterior margin two minute setae, one about twice as long as the other. Eyes prominent, extending slightly beyond margin, with 17-20 facets. Palpi horizontal, oval, microsetose on dorsal surface, ventral surface covered with setae, distal macroseta nearly twice as long as palpus. Theca cordiform, with about 20 setae, four to six along apical margin, four along basal margin, and ten on disk. Thorax. — Mesonotum and chaetotaxy as shown in fig. 74A. Sternopleural projection strong, wide, truncate or feebly emarginate (fig. 62D) , slightly reflexed dorsally. Median pleurotrochantinal lobe narrow, tapered distally, strongly reflexed dorsally. Wings. — As in T. phyllostomae; Hi united with costa at a point just proximal to level of second cross- vein; Rs shorter than distance between fork and crossvein r-m. Legs. — Relatively long, the fore-, mid-, and hindlegs, progressively longer. Femora and tibiae of each pair sub- equal, hind femora approximately as long as thorax. Femora nearly covered with setae, but bare along ventral tibial "groove" and basally on inner face of hind femora ; with long fringing setae along dorsal and ventral outer margins, and along inner ventral margin as well on midfemora; setae elsewhere short, becoming gradually longer toward apices. Tibial setae distinctly longer dorsally than they are laterally and ventrally; denser along inner edge of hind tibiae but not forming a well-defined patch. Hind tarsi slender, about half as long as tibiae, first segment longer than broad; second and third segments subequal ; fourth segment antero-posteriorly compressed dorsally ; last segment somewhat compressed laterally, elongate, narrow, sides subparallel. Abdomen. — Connexivum without setae along middle of dorsum except for four pairs of short, segmental setae ; with short setae laterally and ventrally. Each lateral lobe of tergum I+II with about 19 strong setae and, along lower margin, about five short setae; inner upper margin with a row of four or five fine setae extending medially and an- teriorly from the cluster of macrosetae. Sternum I+II slightly wider than pleurotro- chantines, the setae distributed in a roughly triangular area; posterior margin with longer setae. Female. — Tergum VII short, nearly as wide as supra-anal plate, feebly sclerotized, with a macroseta situated on each lateral margin and a pair of short setae 498 ECTOPARASITES OF PANAMA (one-third length of macrosetae) slightly medial and posterior to macrosetae. Supra- anal plate with four apical macrosetae, a short seta on each side, and another pair of widely separated short setae on disk at mid-length. Seventh sternites relatively small, nearly rounded, each with about 17—18 setae of varying lengths, one a conspicuously longer macroseta. Male: Sternum V well developed; setae progressively longer toward apex; with about 12 macrosetae along apical margin and usually two or more long sub- apical setae. Sternum VI absent. Gonapophyses (fig. 75 A, B) very slender; downwardly curved apically, right gonapophysis a little more strongly so than the left, its ventral macroseta inserted at about basal third, the accessory seta at about mid-length; left gonapophysis with macroseta inserted slightly beyond middle, the accessory seta im- mediately anterior to it. Aedeagus flagelliform. Measurements: BL TL WL ww Male 2.03-2.17 0.66-0.82 1.82-2.03 0.77-0.88 Female 2.07-2.39 0.77-0.78 1.98-2.25 0.85-0.91 TYPE MATERIAL (from Sturnira ludovici) : Holotype male and allotype female (slides) , from host no. 10469, Casa Tilley, 5300 to 5600 feet elevation, near Cerro Punta (Chiriqui) , 14 March 1962, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — 47, as follows : 7, same data and host specimen as holotype and allotype; 2, same locality, 15 February and 23 April 1960; 7 (6 bats), same locality 6 and 11 March 1962; 4 (4 bats), Casa Lewis, near Cerro Punta, 5600-5700 feet elevation, 5 (5 bats), Bambito, near Cerro Punta, 5000-6000 feet elevation, 7 March 1962; 2 (2 bats), Rancho Caballero (Bocas del Toro) , 5000 feet elevation, 11 September 1961 ; 5 (1 bat) , Cerro Mali (Darien), 4100-4800 feet elevation, 9 February 1964, C. 0. Handley, Jr.; 2, same locality and collector, 31 May 1963; 12 (3 bats), Cerro Tacar- cuna (Darien) , 4100-4800 feet elevation, 29 February, 4 and 7 March 1964, C. O. Handley, Jr. Paratypes to be deposited in the collections of Chicago Natural History Museum; the United States National Museum; The Mu- seum of Comparative Zoology ; the American Museum of Natural History ; Gorgas Memorial Laboratory; Environmental Health Branch at Corozal (Canal Zone) ; British Museum (Natural History) ; Departamento de Zoolo- gia, Secretaria da Agricultura, Sao Paulo, Brazil; and the Faculdad de Ciencias, Universidad Central de Venezuela, Caracas. REMARKS: This is one of the few Streblidae collected only above 4000 feet. It was taken only from Sturnira ludovici. The type of T. phyllostomae (Kessel, 1925: p. 17) is "a female in the British Museum, off Phyllostoma, Humboldt, Brazil, July, 1916, W. Erhardt— 1921-200." Through the cour- tesy of Dr. Harold Oldroyd, we received a female specimen from the same locality ("Humboldt, Sta. Cath., 7-VII-1918") from the collection of the British Museum (Natural History). ( It is a female and compares well with a male specimen in our collection taken from "Artibeus j. literatus," at Join- ville, Santa Catharina, Brazil. In both of these, as in the holotype, the upper inner margins of the lateral lobe of abdominal tergum I +11 have an irregu- lar double row of setae. In T. brennani n. sp. and in the undescribed species from Therezopolis (Rio de Janeiro) , Brazil, there is only a single row. The specimen from Therezopolis was taken from a species of Sturnira. It seems very doubtful that the type of T. phyllostomae was actually taken from a species of Phyllostomus. In one of the vials of specimens col- WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 499 B Fig. 74. Thorax, dorsal view. A, Trichobius brennani, new species, holotype male. B, T. vampyropis, new species, allotype female. 500 ECTOPARASITES OF PANAMA lected by Ehrhardt which contained T. phyllostomae, were examples of a species of Megistopoda and of Aspidoptera phyllostomatis (q.v.). These streblids appear to be restricted to Artibeus and Sturnira. The name Phyl- lostomus was loosely applied to phyllostomid bats until early in the Twen- tieth Century. The host of T. phyllostomae was probably a species of Artibeus or Sturnira. T. brennani n.sp. is named after Dr. James Brennan of the Rocky Mountain Laboratory in grateful appreciation of many courtesies and in recognition of his contributions to our knowledge of the chigger mites. Fig. 75. Male gonapophyses (left unless otherwise noted), Trichobius phyllostomae group. A, and B (right gon.), T. brennani, new species, paratype from Sturnira ludovici (no. 8028), Rancho Mojica (Bocas del Toro). C, T. vampyropis, new species, paratype from Vampyrops vittatus (no. 8115), near Mojica (Bocas del Toro). D, T. phyllostomae Kessel, from "Artibeus j. literatus," Joinville (Santa Catharina), BRAZIL. Trichobius vampyropis Wenzel, new species. Figures 74B, 75C. Distinctive in the very large number of eye facets (30-32) , as in brennani n. sp. and phyllostomae Kessel (26-30) ; the very strongly projecting ante- rior margin of the prescutum ; the markedly divergent fifth and sixth longi- tudinal wing veins ; the very small female seventh sternites ; the relatively narrow and long male sternum V with very long apical macrosetae; the distinctive male gonapophyses ; the truncato-emarginate projection of the sternopleura and the very long, tapered, strongly reflexed median pleuro- trochantinal lobe. Most nearly resembling brennani and phyllostomae, but in these smaller species the mesothorax is not as long and is more nearly quadrate; the anterior margin of the prescutum is less strongly projecting; the anterior margin of the sternopleura is truncate but not emarginate ; the WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 501 pleurotrochantinal lobe is short, translucent, and slightly bent but not as- cending ; the fifth and sixth longitudinal veins are not divergent ; the setae of the tibiae are conspicuously longer; and the tarsi are less compressed antero-posteriorly. DESCRIPTION : Head. — Funnel-shaped, convex. Eyes conspicuous, convex, projecting beyond margins of genae, with 30-32 facets. Laterovertices and occipital lobes well denned, separated; occipital lobes with 11 or 12 setae, the anterior ones conspicuously long macrosetae, the posterior setae shorter but coarse, excepting two minute setae near mesal edge of posterior margin. Palpi oval, horizontal, sparsely setose below. Theca pyriform. Thorax. — Sides distinctly converging anteriorly; anterior margin of pre- scutum strongly produced at middle, and distinctly emarginate at midline; median suture not reaching middle; mesonotal chaetotaxy as in fig. 74B. Sternopleura strongly pro- duced anteriorly, the anterior margin relatively broad and distinctly, angulately emar- ginate. Median pleurotrochantinal lobe long, tapered, strongly reflexed dorsally and slightly overlapping the metapleuron, so as to appear united with it, though it is not. Wings. — Long, costal setae rather long basally, becoming shorter apically, of uniform length to apex from about middle of costal cell; Rj united with costa opposite midpoint between second and third crossveins; Rs as long as or slightly longer than distance be- tween fork and r-m; base of R! with 'three macrosetae, base of fourth longitudinal vein with one, of fifth with five or six ; fourth and fifth veins diverging apically. Legs. — Fore- and midlegs subequal, the hind legs distinctly longer, though not markedly elongated. Tibiae uniformly clothed with short setae ; femora with long and short setae, the macro- setae best developed on upper edges. Tarsi antero-posteriorly compressed, claws stout. Abdomen. — Connexivum rather uniformly covered with relatively short setae, except along mid-dorsum, the setae borne on small, circular, sclerotized plaques. Lateral lobes of tergum I+II with numerous, heavy macrosetae. Female: Connexivum with four sets of minute dorsal segmental setae, the anterior three consisting of a single pair each, the posterior one of two pairs; without a ventro-lateral subapical macroseta. Tergum VII small, transverse, with four setae in a transverse row, the outer pair macrosetae, the inner pair a third or fourth as long as the outer. Supra-anal plate with four apical macrosetae and a single shorter seta on each side at mid-length. Seventh sternites small, each with about 13 setae of varying lengths, including several macrosetae, one of these twice as long as the next longest seta. Male: Sternum V relatively long, conspicuously narrower than abdomen, clothed with setae similar to those of connexivum, but with longer setae, including eight macrosetae, along apical margin; sternum VI absent. Ster- num VII+VIII not well developed, with about eight dorso-lateral setas, including one very long macroseta on each side. Tergum IX with two transverse rows of macrosetae dorso-laterally and a third row between these ventrad. Gonapophyses as in fig. 74B. Measurements: BL TL WL ww Male 2.20-2.40 0.77-0.91 2.06-2.19 0.93-1.07 Female 2.31-2.65 0.82-0.95 2.25-2.31 1.03-1.06 TYPE MATERIAL: Holotype male and allotype female (on slides) from Vampyrops vittatus (host no. 8115) near Rancho Mojica (Bocas del Toro), 13 September 1961. In the collection of Chicago Natural History Museum. Two male paratypes, same data as the holotype ; one male paratype, same host and collectors, Rancho Mojica, 12 September 1961 ; one male paratype, same host and collectors, Rio Changena Camp (Bocas del Toro) , 22 Septem- ber 1961 ; three male paratypes from Artibeus lituratus palmarum, 22 miles south of Changuinola (Bocas del Toro), 7 September 1961. All specimens collected by C. M. Keenan, V. J. Tipton, and other personnel of the Environ- mental Health Branch. One each deposited in the collections of the United States National Museum, the Environmental Health Branch at Corozal (Canal Zone), and the Gorgas Memorial Laboratory, Panama (Panama). 502 ECTOPARASITES OF PANAMA REMARKS: The fact that many Streblidae have been collected from Artibeus lituratus in Panama and elsewhere without otherwise recovering this species suggests that the paratypes from Changuinola may have been mislabeled as to host. The median pleurotrochantinal lobe is so long that in preserved speci- mens it reaches to or even slightly overlaps the metapleuron, and the abdo- men of alcohol-preserved specimens must be manipulated in order to determine that the two structures are not united. In slide-mounted speci- mens, it may appear to be united with the metapleuron, even when the slide is viewed from an angle using a binocular dissecting microscope. T. vampyropis is structurally intermediate between other members of the phyllostomae group and a complex of three genera — including Paratricho- bius, Neotrichobius, and Megistopoda — which, like the phyllostomae group occur on bats of the subfamilies Stenoderminae and Sturnirinae. The male gonapophyses are intermediate as regards the insertion of the paired ven- tral setae. Sternum V is similar in all of the above flies, in shape and chaeto- taxy ; it has exceptionally long macrosetae along apical margin. Genus Anatrichobius Wenzel, new genus Type-species: Anatrichobius scorzai Wenzel, new species. Closely related to Joblingia, with very similar head structure and like it, having the accessory setae of the male gonapophyses inserted anterior to the macrosetae. Differing from Joblingia in its much smaller size and relatively short legs and in the following characters: both the lateral and vertical membranous clefts of the thorax are open (longitudinal cleft closed and indicated by a dark suture in Joblingia) ; the median suture of the meso- notum does not extend beyond the transverse suture (to the scutellum in Joblingia) ; sternum V is present and divided into two sternites (absent in Joblingia) ; tergum VII absent or a single, median, very short, transverse, setose sclerite in the female (represented by a vertical, elongated, very feebly sclerotized, setose area with associatd spiracle on each side of hypo- pygium in Joblingia) ; female seventh sternum divided into two longitudinal sternites (a single median sclerite, though strongly emarginate anteriorly and posteriorly in Joblingia) ; finally, the thorax is very high and is dorsally strongly convex in Joblingia, whereas it is distinctly wider than high and only feebly convex above in Anatrichobius. Anatrichobius is also superficially similar to Aspidoptera but differs in that it has elongate, foliaceous palpi rather than transverse nearly vertical ones; the anterior thoracic spiracles are very small and inconspicuous (relatively large and conspicuous in* Aspidoptera) ; sternum V (male) is divided into two sclerites rather than undivided; and the accessory setae of the male gonapophyses are inserted anterior rather than posterior to the macrosetae. DIAGNOSIS : Medium sized species with dorsum of body and legs clothed with long setae. Head. — Convex, oval, sides subparallel. Laterovertices and occipital plates dif- ferentiated, the laterovertices subquadrate. Eyes small, faceted. Palpi leaf-like, longer than broad. Thorax. — Short above, wider than long. Anterior margin of prescutum broadly emarginate, pronotum narrowly emarginate behind the head. Spiracles small. WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 503 inconspicuous. Median mesonotal suture not extending beyond the transverse suture. Scutum very short and wide. Scutellum small. Vertical longitudinal cleft extending nearly to mesocoxal cavity. Venter elongated, slightly longer than wide, sternopleura only slightly longer than pleurotrochantines. Wings. — Brachypterous, venation essen- tially complete but variable. Legs. — Moderately developed, the middle legs shortest, the hind legs longest, metafemora about half again as long as mesofemora. Hind coxae large. Tarsi short; tarsomeres 1-4 strongly compressed antero-posteriorly, 5 large, conspicuous. Abdomen. — Dorsal connexivum densely clothed with long setae. Female: Tergum VII very short and wide, or absent. Supra-anal plate cordiform, pointed at apex. Cerci articulated to supra-anal plate by a short process, united ventrally with the ventral arc. Seventh sternites elongated, their inner apical margins flange-like and strongly sclero- tized, articulated with the median lobe of the ventral arc as well as with a pregenital sclerite (VIII ?). Sternum IX visible externally. Male: Hypopygium broadly conical. Sternum V divided into two transverse, oval, feebly sclerotized sternitas. Sternum VI absent. Gonapophyses blade-like, the accessory setae situated anterior to the macrosetae. Anatrichobius scorzai Wenzel, new species. Figures 76-78. DESCRIPTION : Chaetotaxy as illustrated. Micropile present on upper surface of palpi, head, mesonotum (except a broad median area), and abdomen including setose area of sternum II, and upper setose areas (at least apically) of femora. Intervals between femoral and some coxal setae may appear scabrous in slide preparations. Head. — Broadly pyriform, the dorsum convex. Laterovertices with about 20 setae, most of the anterior setae shorter; occipital lobes with about 25-30 setae. Eyes small, horizontal, with seven or eight small facets. Genae and postgenae densely setose, the dorsal and apical setae longest. Thorax. — Median suture usually united with the transverse suture, sometimes extending only a little beyond middle; transverse suture usually complete, occasionally poorly defined at middle. Prescutum with 38-40 setae on each side of median suture. Scutum with about 24-26 setae. Scutellum with four scutellar macrosetae in a transverse row and two additional ones at middle of apical margin. Mesepisterna clothed with dense long setae above. Wings. — Brachypterous, as shown in fig. 76, venation vari- able. Legs. — Hind tibiae with denser short setae along inner apical face, but not form- ing a dense patch. Last tarsomere about as long as 1-4 combined. Abdomen. — Con- nexivum in both sexes dorsally clothed with long setae; lateral connexival setae shorter, the ventral setae minute. Female: Lateral lobes of tergum I+II longer than in the male, the posterior part of the lobe (viewed from the side) bare; the dorsal margin (near mid- length) with a cluster of 12-14 longer setae which are continued anteriorly and across middle of margin by a row of short setae. Dorsal connexivum clothed with long setae, these more prominent laterally; setae of lateral connexivum shorter, those of venter very short, except for segmentally arranged pairs of long setae. Tergum VII curved pos- teriorly on each side, with from 12-20 setae, some of these often displaced and situated on connexivum. Supra-anal plate with from 10-12 setae. Seventh sternites with numerous macrosetae apically, shorter setae basally. Male: Lateral lobe of tergum I+II posteriorly with numerous discal and marginal setae, these shorter ventrally. Dorsal connexivum with a mixture of long and shorter setae, the long ones shorter than in female. Setae of sternum V similar to those of ventral connexivum, the anterior ones a little shorter. Hypopygium with numerous long macrosetae dorsally and short setae ventrally. Gonapophyses (fig. 78C) nearly alike. Measurements : BL TL Male 2.42-2.64 0.52-0.55 Female 2.99-3.30 0.60-0.77 TYPE MATERIAL: Holotype male (host no. 7459) and allotype female (host no. 7468), both in alcohol, from Myotis sp., Cueva Lara (Chiriqui), 5600 feet elevation, 3 May 1961, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. 504 ECTOPARASITES OF PANAMA Fig. 76. Anatrichobius scorzai, n. gen., n. sp. Female paratype, dorsal view, and right wing. Made from specimens taken from a mixed collection of Myotis n. nigricaws and M. chiloensis, at Cueva Lara, Casa Tilley, Cerro Punta (Chiriqui). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATPLIES 505 Fig. 77. Anatrichobius scorzai, n. gen., n. sp., female. Ventral view. From same speci- mens as fig. 76. 506 ECTOPARASITES OF PANAMA Paratypes. — From Myotis sp. : 1, same data as holotype; 1, Casa Lewis, Cerro Punta (Chiriqui), 5600 feet elevation, 4 May 1960; 1, same data but 12 March 1962. From Myotis nigricans: 4, Cave, Finca Lara, (Chiriqui), over 5800 feet elevation, 5 May 1961 ; 3, Lara's Cave, Cerro Punta (Chiri- qui), 6 March 1962 ; 1, Cerro Hoya (Los Santos) , 21 February 1962. From Myotis chiloensis: 1, Lara's Cave, Cerro Punta (Chiriqui), 5600 feet eleva- tion, 6 March 1962. From a mixed collection Myotis nigricans and M. Fig. 78. Anatrichobius scorzai n. gen., n. sp., male abdomen. A, dorsal, and B, ventral views. C, left gonapophysis, lateral view. Data as for female (fig. 76). chiloensis: 47 (3 lots), Lara's Cave, Cerro Punta (Chiriqui), 6 March 1962. From bat guano: 1, Lara's Cave, Cerro Punta (Chiriqui), 6 March 1962. Without host: 2, Cerro Azul (Panama), elevation about 2000 feet, 25 and 27 January 1958. From Myotis n. nigricans: 30, from a room of old building of Biological Station, Rancho Grande (Aragua), VENEZUELA, 7 July to 30 September 1962, C. and A. Machado, R. Antequera and M. Ramirez [FCUCV]. From Myotis nigricans: 3, San Antonio, San Agustin (Huila), COLOMBIA, 2350 meters elevation, 18 October 1951, P. Hershkovitz, CNHM Colombia Zoologi- cal Expedition (1948-52). From Myotis chiloensis oxyotis: 3, Hacienda Cadena, Marcapata (Cuzco), PERU, 1000 meters elevation, 19 February 1949, Celestino Kalinowski. From Lonchophylla robusta (!) : 1, Rancho WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 507 Grande (Aragua) VENEZUELA, Wm. T. Beebe [MCZ]. Paratypes to be de- posited in the collections of Chicago Natural History Museum ; the United States National Museum; Museum of Comparative Zoology at Harvard University; the Gorgas Memorial Laboratory, at Panama (Panama) ; the Environmental Health Branch (United States Army) at Corozal (Canal Zone) ; and the Universidad Central de Venezuela, Caracas. REMARKS : Anatrichobius scorzai is named in honor of Dr. Jose Vicente Scorza, Head of the Department of Parasitology, Escuela de Biologia, Fa- culdad de Ciencias, Universidad Central de Venezuela. The puparium of this species will be described later by Machado-Allison. The female speci- men illustrated (figs. 76, 77) was distorted by pressure on the slide. In gravid females, the abdomen is never quite as broad and rounded as shown. Tergum VII of the specimen illustrated has fewer setae than is typical. An undescribed species of Anatrichobius is known to us from Peru and Brazil. It differs strikingly from scorzai in that tergum VII is absent in the female and the chaetotaxy of the supra-anal plate is reduced to from four to six setae. Genus Joblingia Dybas and Wenzel Joblingia Dybas and Wenzel, 1947, Fieldiana, Zool., 31: 149. Jobling, 1949, Para- sitology, 39: 322 (keyed). Undescribed genus, Bequaert, 1942, Bol. Ent. Venez., 1: 86 (notes). Type-species: Joblingia schmidti Dybas and Wenzel, 1947. Until the Panamanian collections were obtained, the type-species of Joblingia was known only from the unique female holotype. Both males and females were collected in Panama, and it is now possible to give the characters of the male and to correct several errors in the original diagnosis of the genus and description of the species. The following may be added to the diagnosis of the genus. Head. — Laterovertices and occipital plates differentiated, but weakly sclerotized, connected by a narrow sclerotized strip along inner margins. Eyes small, vertical. Thorax. — Height greater than dorsal width. Mesonotum strongly convex ; median suture extending posteriorly to scutellum. Longitudinal membranous cleft closed, the line of fusion marked by a dark pigmented suture. Vertical membranous cleft reaching about halfway to coxal cavity. Abdomen. — Female: Seventh tergum represented by a vertical elongate setose area (which encloses the spiracle VII) on each side of the hypopygium. Sternum VII a single median plate which is emarginate along mid-line both anteriorly and posteriorly. Cerci united with the ventral arc, the latter not united with the supra- anal plate. Both dorsal and ventral anal sclerites present. Male: Sterna V and VI ab- sent, VII and VIII fused with IX. Hypopygium broadly conical. Gonapophyses with accessory setae inserted anterior to the macrosetae. Joblingia schmidti Dybas and Wenzel. Figures 79, 80. Joblingia schmidti Dybas and Wenzel, 1947, Fieldiana, Zool., 31: 152, figs. 25-27— Guatemala: Chocoyos (Chimaltenango), from Myotis n. nigricans (Chicago Natu- ral History Museum). Jobling, 1949, Parasitology, 39: 316 ff, fig. 3G, H. The following characters are added to the original description: Legs. — Proportionally shorter in males than in females. Abdomen. — Female: Ter- gum I +11 with dorsal and ventral groups of short bristles anteriorly and numerous very 508 ECTOPARASITES OF PANAMA Fig. 79. Joblingia schmidti Dybas and Wenzel, female. Dorsal view. From Dybas and Wenzel (1947), with corrections. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 509 short rather stout bristles on posterior margin of lateral lobe. Tergum VII represented on each side lateral to the terminal cone by a vertical elongate area (with shorter setae than connexivum) which encloses seventh spiracle. Seventh sternum not divided, though it is emarginate anteriorly and posteriorly and the setae are arranged as on two sternites, with a median smooth area between two extensive groups. Male : Lateral lobes of tergum I+II densely clothed with long and short bristles, these shorter along dorsal and ventral margins. Connexivum between lateral lobes with dense setae of moderate length, else- Fig. 80. Joblingia schmidti Dybas and Wenzel. A, left, and B, right male gonapophyses, lateral view. From Myotis nigricans (no. 7466), Cueva Lara, Casa Tilley, Cerro Punta (Chiriqui). where (dorsally and ventrally) except along mid-line near apex, with minute setae borne on small plaques. Hypopygium broadly conical, densely clothed with setae, these very long macrosetae posteriorly, shorter anteriorly and ventrally. Sternum II less strongly emarginate at middle than in female. Gonapophyses as in fig. 80. Measurements : Male Female BL 3.96-4.06 4.97-5.50 TL 1.04-1.07 1.03-1.16 MATERIAL EXAMINED : 31 specimens (10 lots) , all from Cueva Lara, Casa Tilley, Cerro Punta (Chiriqui), elevation 5600 feet. — From Myotis nigri- cans: 10 (4 bats), 3 May 1961. From Myotis sp. : 1, 5 May 1961. From Myotis chiloensis: 1, 3 May 1961. From mixed collection of M. nigricans and M. chiloensis: 1 (3 lots), 6 March 1962. From bat guano: 8, 6 March 1962. REMARKS: The original figures of Dybas and Wenzel (op. cit.), showed the laterovertices and occipital lobes as poorly defined areas. These were poorly pigmented and defined in the female holotype. The figure is repro- duced here (fig. 79) with corrections. 510 ECTOPARASITES OF PANAMA Trichobioides Wenzel, new genus Type-species : Trichobius perspicillatus Pessoa and Galvao, 1937. Trichobioides closely resembles Trichobius, but differs as follows: palpi without setae on ventral surface (present in Trichobius) ; upper surface of palpi with a longitudinal tuberculate ridge near mesal margin (absent in Trichobius) ; costa and RI united opposite third crossvein (beyond level of third crossvein in Trichobius) ; palpi and theca (when retracted) co- Fig. 81. Trichobioides perspicillatus (Pessoa and Galvao). Thorax, dorsal view, of male from Phyllostomus d. discolor (TVL nos. 1142-46), Santa Cruz, TRINIDAD. apted with apices of genae to form a vertical shield across the front of the head (not so in Trichobius) ; laterovertices with an oblique dark line which divides them into two parts (absent in Trichobius) ; and the female seventh sternites and the connexivum forming prominent apical lobes (not so in Trichobius) . DIAGNOSIS: Head. — Co-apted with and nearly as broad as the bisinuate anterior thoracic margin. Laterovertices and occipital lobes well denned, the laterovertices divided into two parts by an oblique, dark line (most conspicuous in alcohol specimens). Postvertex sclerotized, without setae, continuous with the occipital lobes. Palpi broad, with few setae, these restricted to margins; ventral surface bare, upper surface micro- setose; upper surface with a sharp longitudinal bituberculate ridge near mesal margin, each tubercle bearing a short seta. Eyes conspicuously separated from lateral margins of head. Thorax. — Broad, narrowing anteriorly; prescutum very long, more than twice as long as scutum. Wings. — Relatively short; setae of veins conspicuous; costa and Ri with macrosetae from base to juncture, opposite the third crossvein. Legs. — Short, WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 511 middle and hind legs subequal, the hindlegs slightly longer; femora stout, with both macrosetae and short setae ; tibiae with very short, fine setae only ; tarsi short. Abdomen. — Sternum I+II broad, as wide as abdomen. Female: Tergum VII very short. Cerci not united with the heavily sclerotized ventral arc. Seventh sternites long, produced apically as subconical lobes, which extend as far as or beyond apex of proctiger. Male: Hypopygium prominent, broadly conical. Sternum V short and broad, not divided. Sternum VI present as a very fine thread-like band. Gonapophyses with paired setae in- serted on lateral face, the accessory seta below or slightly posterior to macroseta. Fig. 82. Left male gonapophysis, lateral view. A, Trichobioides perspicillatus (Pessoa and Galvao), from Phyllostomus d. discolor (no. 6348), Rio Mandinga, (San Bias). B, Mastoptera minuta (Costa Lima), from Tonatia silvicola (no. 9477) Guanico (Los Santos). C, Mastoptera guimaraesi, new species from Phyllostomus h. panamensis (no. 4644), Chepo Road (Panama). Trichobioides perspicillatus (Pessoa and Galvao), new combination. Figures 81, 82A. Trichobius perspicillatus Pessoa and Galvao, 1937, Folia Clin. Biol., Sao Paulo, 9: 1, figs. 1-3 — "Bahia," from undet. host and Hemiderma perspicillatum (Lab- oratorio de Parasitologia, Faculdade de Medicina, Universidade de Sao Paulo). Guimaraes, 1938, Rev. Mus. Paulista, 23: 655 (notes), 662 (keyed), fig. 4. Jobling, 1938, Parasitology, 30: 386, fig. 14A; 1949, ibidem, 39: 316 ff. (hosts), 326. This remarkable species is easily recognized by the very distinctive mesonotal chaetotaxy (fig. 81). The female is also distinctive in the lobes formed by the seventh sternites. Measurements: BL TL WL ww Male 1.00-1.54 0.49-0.51 0.93-1.10 0.49-0.55 Female 1.59-1.84 0.53-0.58 1.10-1.15 0.55-0.60 PANAMANIAN MATERIAL EXAMINED : 12 specimens in (5 lots) from 6 bats. From Phyllostomus d. discolor: 8 (2 bats), Rio Mandinga (San Bias), 18 and 29 May 1957, E. Mendez [GML] ; 1 (probably this host), Cerro Azul (Panama), about 2000 feet elevation, 25 January 1958, E. Mendez [GML]. From Sturnira lilium parvidens: 1, Rio Tuira (Darien), 25 February 1958, 512 ECTOPARASITES OF PANAMA P. Galindo [GML]. From Desmodus rotundus murinus: 2 (2 bats) , Guanico (Los Santos) , 26 January 1962. OTHER MATERIAL EXAMINED : More than 100 specimens from GUATEMALA, EL SALVADOR, COSTA RICA, COLOMBIA, VENEZUELA, SURINAM, TRINIDAD, and PERU. Nearly all of these are from Phyllostomus d. discolor and P. d. verru- cosus, except one record from Phyllostomus elongatus (COLOMBIA) and one from a bat doubtfully identified in the field as Phyllostomus h. hastatus (SURINAM). REMARKS : The primary host of this fly appears to be Phyllostomus dis- color, though it probably occurs on other species of Phyllostomus. The Panamanian records from Sturnira and Desmodus may represent transi- tory associations or errors in field labeling. As noted elsewhere, a number of the flies collected in Los Santos were incorrectly associated with the hosts. Mastoptera Wenzel, new genus Type-species : Aspidoptera minuta Costa Lima, 1921. The species of Mastoptera superficially resemble those of the genus Aspidoptera but differ in having mouthparts and genae co-apted (in re- pose) becoming a frontal shield, as in Trichobioides ; the posterior ex- tension of the median mesonotal suture nearly to the scutellum ; the closure of the vertical membranous cleft without evident suture ; the reduced wing venation and the apical digitiform process at apex of the wing; the pres- ence of flanges on the ventral arc (female) and numerous other characters. Actually, Mastoptera is closely related to the macropterous Trichobioides and possesses many characters of that genus, including the co-apted mouth- parts ; the palpal flange ; the posteriorly strongly widened thorax ; similar male gonapophyses (though the paired ventral setae are inserted more distally) ; the broadly conical hypopygium, the presence of flanges on the ventral arc of the female, and the conspicuously long macrosetae of the apex of the abdomen. Easily separated from all other genera by the extra- ordinarily widely separated hind coxae, and by the digitiform process at the apex of the wings. DIAGNOSIS: Minute dorso-ventrally somewhat flattened brachypterous species, 0.73- 1.29 mm. long. Head. — Sides of head converging posteriorly, the posterior margin and anterior edge of the prescutum co-apted. Laterovertices and occipital plates distinct. Theca short, wider than long, vertical when retracted and co-apted with palpi and genae to form a frontal "shield." Palpi vertical, subtriangular, apices broadly truncate; ventral surface with setae on basal half only; upper surface microsetose on distal half, with a longitudinal submarginal flange near inner edge. Eyes minute, with one or two facets. Thorax. — Anterior margin of prescutum strongly emarginate, on a little more than middle third of width. Median suture extending posteriorly to or near the scutellum; transverse suture absent or indistinct, represented by pigmentation only, though some- times distinct at middle. Posterior margin of scutellum emarginate. Longitudinal membranous cleft open, short; vertical membranous cleft closed, without evidence of a suture. Spiracles small, rosette-like. Mesepisterna broad and setigerous dorsally, widest posteriorly. Wings (fig. 83C). — Brachypterous, venation indistinct; each wing with a digitiform process (rarely two ?) at apex. Anterior margin of sternopleura trun- WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 513 cate, feebly projecting. Posterior margin of pleurotrochantines nearly straight, very broad, approximately as wide as hind femora are long. Legs. — Short, subequal. Hind coxae widely separated. Femora with long sparse setae antero-dorsally, in addition to short ones on antero-ventral or posterior faces. Tibiae with very short setae arranged in rows, tibiae and tarsi subequal. Tarsomeres 1-4 progressively shorter, 4 rather strongly compressed antero-posteriorly; sides of last tarsomere subparallel. c Fig. 83. Mastoptera guimaraesi, new species. A, dorsal view, male. B, labium. C, wing. From Jobling (1957, as Aspidoptera minuta) . Abdomen. — Tergum I+II membranous along mid-line, nearly to base; lateral lobes conspicuous, usually about half as long as abdomen and bearing conspicuous macrosetae. Connexival setae borne on conspicuous plaques. Female: Tergum VII and supra-anal plate together forming a broad terminal cone; tergum VII feebly sclerotized, but ap- parently subquadrate and about twice as broad as long; supra-anal plate short and transverse; seventh sternites small, slightly produced posteriorly as lobes. Cerci free, not united with the ventral arc, the latter with flange-like projections. Male: Sternum V present, short and broad. Sternum VI absent. Hypopygium very broadly conical. Gonapophyses curved; paired setae inserted near mid-length, the macroseta anterior to the accessory seta; gonapophyseal apodeme trough-like. Adeagus flagelliform. At present we are able to distinguish two species of this genus: M. minuta (Costa Lima) from bats of the genus Tonatia and from Phyllos- 514 ECTOPARASITES OF PANAMA tomus hastatus, and M. guimaraesi n.sp. from Phyllostomus hastatus pana- mensis. It is possible that careful study of adequate material may show that each species of Tonatia harbors a separate species of Mastoptera, as is true for the streblids of the genera Pseudostrebla, Trichobius, and Strebla. KEY TO SPECIES OF MASTOPTERA MALES Tergum I+II with seven to nine conspicuous, long to moderately long bristles, and, along lateral margin, five to seven short ones; longest seta followed distally by one long bristle, lateral to which is a very short bristle (fig. 84A). Gonapophyses more strongly curved, with dorsal margin feebly sinuate before apex (fig. 82C). Host: Phyllostomus hastatus panamensis guimaraesi n.sp. Tergum I+II with nine to twelve conspicuous bristles and, along lateral margin, two or three very short ones; longest seta followed distally by two shorter but con- spicuous, long bristles (fig. 85B). Gonapophyses with dorsal margin evenly curved before apex (fig. 82B). Hosts: Tonatia spp., Phyllostomus hastatus (? subsp. hastatus) minuta (Costa Lima) FEMALES Lateral lobes of tergum I+II very long, nearly as long as hind coxa and femur com- bined. Dorsal abdominal connexivum with from one to three setae which are con- spicuously longer and coarser than the others, on each side medial and slightly posterior to apices of lateral lobes of tergum I+II (fig. 84A). Sternum II nearly as long as venter of thorax; seventh sternites with setae along inner and distal margins only, elsewhere microsetose guimaraesi n.sp. Lateral lobes of tergum I+II only slightly longer than hind femora. Dorsal ab- dominal connexivum lacking conspicuously coarser and longer setae medial to apices of lateral lobes of tergum I+II. Sternum II only a little more than half as long as venter of thorax; seventh sternites with setae throughout, in addition to microsetae minuta (Costa Lima) Mastoptera guimaraesi Wenzel, new species. Figures 82C, 83, 84. Aspidoptera minuta (not Costa Lima, 1924), Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417; 1942, Bol. Ent. Venez., 1: 88 (records). Jobling, 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 142 (part.), fig. 4; 1949, Parasitology, 39: 316-317 (part.), fig. II. Very similar to M. minuta (Costa Lima) , but differing by the characters given in the key (and noted below). As indicated above, the description and figure given by Jobling (1949) apply to this species, except for the record from Tonatia amblyotis (— T. silvicola) . In addition to characters given by Jobling, the following diagnostic features may be noted. DESCRIPTION: Head. — Eyes with two indistinct facets (one in minuta). Abdomen: Male. — Tergum I+II with seven to nine conspicuous long bristles and five to seven short ones along the lateral margin; longest macroseta followed distally by one shorter but long, conspicuous seta lateral to which may be a short seta. Gonapophyses with upper margin slightly sinuate before apex (fig. 82C). Female: Tergum I+II very long, nearly as long as hind femur and coxa combined. Connexivum dorsally with from one to three coarse bristles longer than the others, medial and slightly apical to apex of lateral lobes of tergum I+II. Sternum II nearly as long as underside of thorax. Seventh sternites with setae restricted to distal and inner margins, the rest of the surface microsetose only. Measurements : BL TL Male 1.00-1.10 0.27-0.29 Female 1.23-1.29 0.29-0.33 TYPE MATERIAL: Holotype male (slide) from Phyllostomus hastatus WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 515 panamensis (host no. 4644), hollow tree, Chepo Road (Panama), 8 October 1959, C. M. Keenan and V. J. Tipton ; allotype female (slide) , same data but host no. 4641 ; both in the collection of Chicago Natural History Museum. Paratypes. — From Phyllostomus hastatus panamensis, 284 in 30 lots from 68 bats, as follows: 1, Madden Dam (Canal Zone), 31 August 1959; 75 (19 bats), Fort Sherman (Canal Zone), 4 and 6 April 1960; 2, Fort Clayton (Canal Zone) , 20 June 1960 ; 2, Farfan (Canal Zone) , 19 December 1915, T. Hallinan [AMNH] ; 161 (18 bats), same data as the holotype; 2 (2 bats) , Chilibrillo Caves (Panama) , 28 October 1959, and 12 (20 bats) , same locality, 17 July 1959; 1, same locality, 28 May 1931, L. H. Dunn no. 22 [AMNH] ; 2, same locality, without data [MCZ] ; 2, Tapia (Panama) , 25 July 1923, J. P. Chapin [MCZ]; 1, "Panama," [USNM] ; 5 (1 bat), Almirante (Bocas del Toro). From Phyllostomus sp. : 6 (2 bats), Camp Pina (Canal Zone), 26 October 1960; 1, Guanico (Los Santos), 22 January 1962. Para- types to be deposited in the collections listed on p. 410. OTHER MATERIAL EXAMINED: PANAMA. — From Carollia p. azteca: 1, culvert, Chepo Road (Panama). Without host: 1, Chilibrillo Caves (Pan- ama), 17 July 1959; 2, same locality [MCZ] ; 1 (slide), "Canal Zone, Busck, April 30" [USNM, det. as A. minuta by Q. C. Kessel] . COLOMBIA. — Several lots taken from Phyllostomus hastatus in the Departments of Huila and Bolivar, by P. Hershkovitz, CNHM Colombian Zoological Expedition 1948-52. REMARKS: Jobling indicated that this species had a single eye facet, but our material indicates that there are two indistinct facets. In minuta, there appears to be only one. Jobling's figure (see fig. 83) is of a specimen from Phyllostomus h. panamensis taken at Chilibrillo Caves (Panama) . It is somewhat inaccurate or atypical in several respects. The anterior mar- gin of the prescutum is not broadly emarginate, as shown, but straight on each side and emarginate at middle only. The wing is shown with two setigerous digitiform processes at apex. We have not seen more than one such process in the specimens examined by us. This species is named for Dr. Lindolpho Guimaraes of the Departa- mento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil, in recog- nition of his many valuable contributions to our knowledge of ectopara- sites, and especially his outstanding studies on the New World batflies, both Streblidae and Nycteribiidae. Mastoptera minuta (Costa Lima), new combination. Figures 82B, 85. Aspidoptera minuta Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 21-22 (descr., keyed), 29 (cat.), pi. 2, fig. 2— Brazil: Matto-Grosso, from Tonatia amblyotes [sic!] (? Museu Nacional do Rio de Janeiro). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 26, pi. 2, fig. 13 (? part.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (fig., Streblidae II.— 4). Jobling, 1949, Parasitology, 39:316-317 (part); 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 136 (keyed), 142 (part., record from Tonatia only). This species may be separated from guimaraesi by the characters given 516 ECTOPARASITES OF PANAMA A B Fig. 84. Mastoptera guimaraesi, new species, base of abdomen, dorsal view. A, male (host no. 4644) and B, female (host no. 4641) paratypes from Phyllostomus h. panamen- sis, Chepo Road (Panama) . ws=dorsal connexival macroseta. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 517 B Fig. 85. Mastoptera minuta (Costa Lima). A, female, base of abdomen, dorsal view, paratype from Tonatia silvicola (no. 11698) Armila (San Bias). B, abdomen, male, dorsal view, paratype from T. silvicola (no. 9477), Guanico (Los Santos). 518 ECTOPARASITES OF PANAMA in the key. As can be seen from the following measurements, it is dis- tinctly smaller than that species. Measurements : BL TL Male 0.73-0.96 0.19-0.30 Female 0.83-1.06 0.24-0.29 PANAMANIAN MATERIAL EXAMINED: 58 specimens (21 lots from 21 bats). From Tonatia minuta, 22 (6 bats) as follows: 3, Fort Sherman (Canal Zone), 2 December 1959; 2, Almirante (Bocas del Toro), 26 February 1960; 12 (2 bats), Sibube (Bocas del Toro), 17 and 24 January 1963, C. O. Handley, Jr.; 1, Las Palmitas (Los Santos), 24 January 1962; 5 (2 bats), Cerro Hoya (Los Santos), 24 January and 26 February 1962. From Tonatia silvicola, 35 (14 bats) as follows: 2, Sibube (Bocas del Toro), 22 January 1963, C. O. Handley, Jr.; 11 (5 bats), Guanico (Los Santos), 27 January to 1 March 1962; 21 (6 bats), Armila (San Bias), 1 to 29 March 1963, C. 0. Handley, Jr. and F. M. Greenwell ; 3, Puerto Obaldia (San Bias) , 3 April 1963, C. 0. Handley, Jr.; 1, Cerro Mali (Darien), 8 February 1964, C. O. Handley, Jr. From Phyllostomus sp. (!):!, Chilibrillo Cave, 2 May 1957, Koford and Handley [USNM]. OTHER MATERIAL EXAMINED : About 250 specimens, mostly from various species of Tonatia, from COLOMBIA, SURINAM, ECUADOR, PERU and BOLIVIA are referred to this species. Five lots (28 specimens) taken from Phyl- lostomus hastatus in Amazonian COLOMBIA, by Kjell von Sneidern, were also examined. REMARKS: We suspect that the specimens taken on Phyllostomus has- tatus (subsp. ? hastatus) in Amazonian Colombia represent a third species, but to date we have not been able to distinguish between these and speci- mens from bats of the genus Tonatia. If these actually are minuta, then we must conclude that this species is able to live on bats of the genus Tonatia throughout the range of that genus, but on Phyllostomus hastatus only in Amazonian South America, and that it is replaced by guimaraesi on P. h. panamensis in northern South America and Central America. Genus Paratrichobius Costa Lima Paratrichobius Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Veter., Nictheroy, 5: 20 (diagn.), 25 (keyed), 28 (cat.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 12 (keyed), 19 (diagn.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 655. Curran, 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed), 7 (discuss.). Jobling, 1936, Parasitology, 28: 359 ff. (morph.). Pessoa and Guimaraes, 1937, Ann. Fac. Med. Sao Paulo, 12: 265, 267 (discuss.). Jobling, 1939, Parasitology, 31: 490 (diagn.). Bequaert, 1940, Rev. Acad. Colomb., Cienc. Exact., Fis. y Nat., 3: 417 (keyed) ; 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Parasitology, 39: 321 (keyed). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 177 (keyed), 187. Maa, 1965, Jour. Med. Ent., 1: 389 (cit., cat.). Type-species: Trichobius longicrus Miranda Ribeiro, 1907. Paratrichobius, and the related genera, Neotrichobius and Megistopoda appear to have been derived from the phyllostomae group of the genus Trichobius. All of these occur on fruit-eating bats of the subfamilies Steno- derminae or Sturnirinae. They share a number of characters, including WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 519 the large multi-faceted eyes ; the presence of a female postgenital sclerite, similar to that found in the caecus group of Trichobius; the fusion of the cerci to the ventral arc in the female ; and the distal insertion on at least one gonapophysis (male) of at least one of the ventral, paired gonapophyseal setae. In addition, Paratrichobius, Neotrichobius, and Megistopoda have a very similar shield-like thoracic venter. Paratrichobius appears to be the least, and Neotrichobius the most specialized genus. The species of Paratrichobius have presented us with some of the most difficult taxonomic problems in the current study. Unfortunately, the paucity of material has prevented us from resolving many of these prob- lems. Only two species, P. longicrus (M. Ribeiro) and P. dunni (Curran), have been described, but it is obvious from our material that a number of species, or other segregates, occur on various genera and species of the host bats. In Panama, the most confusing complex, taxonomically, is a series of segregates related to P. longicrus that occur on Artibeus toltecus, A. aztecus, A. jamaicensis, A. lituratus palmarum, and Vampyrops vittatus. They are very similar in chaetotaxy, but they differ, some conspicuously, in relative and absolute length of the wings and hind femora, and some in the number of setae present on sternum II and on the female seventh ster- nites. Our series are too small to permit significant analysis of these differ- ences. Further, it appears that there may have been some contamination, temporary transfers, and misidentifications of hosts, which complicate anal- ysis of the data. The Paratrichobius from A. jamaicensis and A. lituratus (fig. 88 A) differ from the others in having less strongly curved male gonapophyses, which are nearly identical in both samples. Their measurements, too, are very similar, though the specimens from A. jamaicensis appear to be slightly smaller and to have shorter wings in the male than do those from lituratus. The segregates from A. aztecus, A. toltecus, and Vampyrops vittatus have more strongly curved male gonapophyses (fig. 88B-D). As can be seen in fig. 89, the specimens from V. vittatus have longer wings than do any others excepting those from A. aztecus, and they differ significantly from all others in the small number of discal and submarginal setae on sternum II. The specimens from A. toltecus have shorter hind femora than do any others of this complex, and in this respect they approach dunni (Curran), salvini n. sp., lowei n. sp., sanchezi n. sp., and species A and B from Chiroderma villosum and Vampyrops helleri, respectively. The least reliable measurements are those of the hind femora. These appear to be bowed more than normally under certain conditions of preservation, espe- cially after treatment with KOH, and mounting on slides. It is our opinion that at least some of these segregates are either cryptic species or populations that are now speciating on the Stenoderminae. In- terestingly, the taxonomic complexities in the longicrus complex roughly parallel those of their hosts. It would seem that the hosts and their para- sites are at the various levels of taxonomic differentiation to be expected in groups that are rapidly evolving. However, it is also possible that the segregates of the longicrus complex represent different developmental re- sponses to different hosts or ecological situations, or represent local popu- 520 ECTOPARASITES OF PANAMA B Fig. 86. Paratrichobius longicrus (M. Ribeiro), male from Artibeus j. palmarum, TRINIDAD. A, dorsal view. B, underside of head. C, anterior portion of venter of thorax. From Jobling (1939a). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 521 lation differences in gene frequency. Much more material and biological information are needed in order to pursue these problems further. The altitudinal distribution of the hosts should be noted (fig. 89) in evaluating the differences between the flies. Paratrichobius longicrus (Miranda Ribeiro). Figure 89. Trichobius longicrus Miranda Ribeiro, 1907, Arch. Mus. Nac., Rio de Janeiro, 14: 236, pi. 25 — Quinta da Boa Vista [Rio de Janeiro], Brazil, from Artibeus jamai- censis (? Museu National de Rio de Janeiro). Paratrichobius longicrus Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 20, pi. 1, fig. 3 (wing). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 656 (cat.). Pessoa and Guimaraes, 1937 (part. ?), Ann. Fac. Med. Sao Paulo, 12: 265 (cit. host records). Jobling, 1939, Parasitology, 31: 492 (part., record from A. jamaicensis, Brazil only). 1949, ibidem, 39: 327 (part., records from A. jamaicensis, Brazil only). We provisionally accept the determination of the type host of P. longi- crus as A. jamaicensis, though it may have been incorrect.10 The type host was later given by Jobling (loc. cit.) following Stiles and Nolan (1931), as A. j. jamaicensis. This is an obvious error. The nominate race of jamaicensis does not occur in southern Brazil. We assign the Panamanian specimens taken from A. j. jamaicensis to P. longicrus. However, it is possible that A. jamaicensis is parasitized by a different species of Paratrichobius in southern Brazil than it is in Panama. It is our opinion that this bat does not normally harbor a species of Paratrichobius, but instead harbors Megistopoda aranea (Coquillet). Of 103 Panamanian specimens of A. j. jamaicensis that were parasitized by Streblidae, 56 (54.47e) carried M. aranea and only 4 (3.8%) carried a Paratrichobius sp. (a total of 5 specimens). Other typical parasites of jamaicensis, such as Aspidoptera busckii and Metelasmus pseudopterus occurred on 49.5% and 20.4% of the parasitized hosts, respectively. Only parasites that were obvious transients or contaminants, like Trichobius joblingi n.sp. or Strebla vespertilionis showed as low an incidence of parasitization as did Paratrichobius longicrus. This suggests that A. jamaicensis is acquiring its Paratrichobius from another lowland host, possi- bly A. lituratus palmarum. The only specimens which show close concor- dance in their measurements with those from jamaicensis are those from A. lituratus. However, in spite of the small number of specimens measured, the differences between individuals from the two hosts lead us to treat them separately until further data permit clarification of their status. The average number of discal setae on sternum II was 14 in both sexes, with a range of 12-16. PANAMANIAN MATERIAL EXAMINED: From Artibeus j. jamaicensis, 5 (4 bats) , as follows : 2, Almirante (Bocas del Toro) , 28 January 1960 ; 1, Fort Kobbe (Canal Zone), 13 October 1960; 1, Guanico (Los Santos), 12 Janu- ary 1962; 1, La Laguna (Darien), 17 June 1963, GML. 10 The taxonomy of the genus Artibeus has been much confused. A revision by C. O. Handley, Jr. is in press. 522 ECTOPARASITES OF PANAMA B Fig. 87. Thorax, dorsal view, Paratrichobius spp. (longicrus complex). A, male from Vampyrops vittatus (no. 8612) Rio Changena Camp (Bocas del Toro). B, male from Artibeus aztecus (no. 10344), Casa Tilley, Cerro Punta (Chiriqui). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 523 Paratrichobius sp. (? longicrus M. Ribeiro) from Artibeus lituratus palmarum. Figures 86, 88A, 89, 91B. Paratrichobius longicrus (? not M. Ribeiro, 1907). Jobling, 1939, Parasitology, 31: 490-92, (part., from A. jamaicensis palmarum), fig. 2. Bequaert, 1942, Bol. Ent. Venez., 1: 88. Jobling, 1949, loc. cit., 39: 316-317 (part., from A. jamaicensis palmarum) , fig. IB, C. Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122: 261. Starrett and de la Torre, 1964, Zoologica, 49: 61. Fig. 88. Male gonapophyses, Paratrichobius spp. (longicrus complex). A, P. longi- crus (M. Ribeiro), left gonapophysis, specimen from Artibeus lituratus (CNHM no. 94666), Morretinho (Sao Paulo), BRAZIL. B, left gonapophysis, specimen from Artibeus aztecus (no. 10344), Casa Tilley, Cerro Punta (Chiriqui). C, right (upper) and left (lower) gonapophyses of specimen from A. toltecus (no. 8116), near Mojica (Bocas del Toro). D, same of specimen from Vampyrops vittatus (no. 8612) Rio Changena Camp (Bocas del Toro) . • Mr. B. Jobling informed us (pers. comm.) that the specimen figured by him (1939, 1949) as P. longicrus is from Artibeus jamaicensis palmarum (= A. lituratus palmarum). The small numbers of A. lituratus palmarum and its parasites that were collected and examined in Panama make it difficult to analyze the parasites of this host. It may be that palmarum is normally parasitized only by Paratrichobius. These were recovered from 12 (70.6%) of 17 host bats parasitized by Streblidae. A single specimen of Aspidoptera phyllostomatis was also taken from them as were two Megistopoda aranea. 524 ECTOPARASITES OF PANAMA Although the sample is small, the wing measurements (fig. 89) of the specimens from A. lituratus palmarum suggest that two categories of wing lengths are represented. The "long-winged" males and females are from Bocas del Toro (22 mi. S. of Changuinola) , while all of the "short-winged" forms except one (from Sibube, Bocas del Toro) are from San Bias. The short-winged forms have fewer discal setae on sternum II, 9-11 in the male, as opposed to 16-18 in the long-winged forms. Among various possibilities, it may be suggested the long and short- winged forms could represent contaminations, geographic segregates, di- morphism, or different host segregates. The latter should not be overlooked in view of the complex taxonomic problems in the genus Artibeus. Ex- tensive and geographically representative collections are needed to clarify the status of the Paratrichobius on A. lituratus. PANAMANIAN MATERIAL EXAMINED : From Artibeus lituratus palmarum (18 flies in 9 lots from 7 bats) as follows : 7, 22 miles south of Changuinola (Bocas del Toro), 7 September 1961 ; 1, Sibube (Bocas del Toro), 27 Janu- ary 1963, C. O. Handley, Jr.; 2 (1 bat), railroad area south of Paraiso (Canal Zone), 5 November 1959; 5 (1 bat), Kobbe Beach (Canal Zone, 12 November 1956; 3 (3 bats), Barro Colorado Island (Canal Zone), 11 and 12 December 1956 and 12 January 1957. Paratrichobius sp. (longicrus complex), from Vampyrops vittatus. Fig- ures 87A, 88D, 89. Specimens from this host differ rather markedly from the others in the small number of discal setae on sternum II. The number ranged from 7-13, with a mean of 10, in 22 specimens checked. PANAMANIAN MATERIAL EXAMINED: 22 males and 10 females in 22 lots from 22 bats, as follows: 1, Rancho Caballero (Bocas del Toro), 11 Sep- tember 1961; 2 (2 bats), Rancho Mojica (Bocas del Toro) and 1, near Rancho Mojica, 9 to 13 September 1961; 13 (9 bats), Rio Changena Camp (Bocas del Toro), 28 September 1961; 6, Cerro Mali (Darien), elevation 4800 feet, 1 May and 1 to 6 June 1963, GML ; 3, same locality, 4 February 1964, C. O. Handley, Jr. ; 5, Cerro Tacarcuna (Darien) , 4 and 7 March 1964. REMARKS : Males were present on 20 of the 22 Vampyrops vittatus para- sitized by Paratrichobius, whereas females were present on only eight. Seventeen bats had only a single fly ; in 13 instances, these flies were males. Most of the hosts were collected in mist nets. We are not in a position to explain these sex ratios. It is possible that the females may leave the host more readily when disturbed, than do -the males, or that a smaller percent- age of them stay with the host bats when they leave the roost. It hardly seems likely that this differential represents the number of females that are off the host depositing pupae at any one time. Paratrichobius sp. (longicrus complex) from Artibeus aztecus. Figures 87B, 88B, 89. The wings of this form are the longest, in relation to the length of the hind femora, of any examined. The number of discal setae on sternum II WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 525 Artibeus toltecus A. aztecus A. j. jamaicensis A. I . palmarum Vampyrops vittatus ELEVATION: 32OO FEET 53OO FEET SEA LEVEL -3200 FEET SEA LEVEL-32OO FEET 24OO - 53OO FEET - LENGTH --OF WING - THORAX o o 0° • MALE O FEMALE Fig. 89. Paratrichobius longicrus complex. Measurements of length of wing, hind femur, and thorax of specimens from hosts indicated at head of columns. All measure- ments are for specimens collected in Panama. 526 ECTOPARASITES OF PANAMA Fig. 90. Paratrichobius dunni (Curran). A, dorsal view, female. B, underside of head and anterior portion of sternopleura. C, apex of abdomen, male, ventral view. From Jobling (1939a). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 527 was 11, 12, 16, and 17 in the four males, and 15 and 18 in the two females. MATERIAL EXAMINED: 5 (1 bat), Casa Tilley (Chiriqui), 11 March 1962; 1, same locality, 1 May 1960. Paratrichobius sp., from Artibeus toltecus. Figures 88C, 89. Measurements could be made on only four of the six specimens taken from A. toltecus. These flies are smaller than any of the others, but have proportionately longer wings. The number of discal setae on sternum II ranged from 11-15 (12 on the single female), with a mean of 13 for the four males. MATERIAL EXAMINED: 6 (6 bats) : 2, Rancho Mojica (Bocas del Toro), 13 September 1961 ; 1, Rio Changena Camp (Bocas del Toro) , 20 September 1961; 2, Casa Lewis (Chiriqui), 1 and 5 February 1960; 1, Casa Tilley, Cerro Punta (Chiriqui), 24 April 1961. Paratrichobius dunni (Curran). Figures 90, 91 A, 92A. Speiseria dunni Curran, 1935, Amer. Mus. Novit., no. 765, p. 7, fig. 6 — El Real, Panama, from either "Pteropteryx [sic!] canina or Uroderma bilobatum" (Ameri- can Museum of Natural History). Paratrichobius dunni Jobling, 1939, Parasitology, 31 : 492-494, fig. 3A-C. Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418. This distinctive species appears to be restricted to Uroderma bilobatum. It is related to salvini, but differs markedly in that the males possess spine- like setae along the ventro-apical margins of tergum IX (fig. 91A) . It also differs in having more extensive prescutal chaetotaxy. There can be little doubt that the type host of P. dunni was Uroderma bilobatum, since the species of Paratrichobius are apparently restricted to Stenoderminae. Peropteryx is a genus of Emballonuridae. The only streblid we have seen from Peropteryx is an undescribed species of Tricho- bius. Curran's two paratypes of dunni were taken from U. bilobatum at Summit (Canal Zone). Measurements: BL TL WL ww FL Male 1.54-1.76 0.56-0.62 1.21-1.22 0.60 0.48 Female 1.59-1.76 0.59-0.63 1.35-1.59 0.63-0.69 0.49 PANAMANIAN MATERIAL EXAMINED : From Uroderma bilobatum, 146 flies in 41 lots (57 bats), as follows: 32 (17 bats), Summit Gardens (Canal Zone), 11 September 1959; 8, same locality, 31 May 1960; 17 (4 bats), palm trees, Fort Randolph (Canal Zone), 6 October 1959; 5, France Field, 3 August 1960; 9 (2 bats), Fort Davis (Canal Zone), 14 and 22 September 1960; 24 (11 bats), Fort Clayton (Canal Zone), 28 November 1960; 1, Almirante (Bocas del Toro) , 28 January 1960 ; 1, Sibube (Bocas del Toro) , 26 January 1963, C. 0. Handley, Jr.; 1, Escobal (Colon), 28 September 1960; 9 (4 bats), Punta de Cocos, Isla del Rey (Darien), 22 and 25 March 1960; 7 (5 bats), Punta de Pina (Darien), 24 March 1960; 8 (7 bats), Puerto Obaldia (San Bias), 3 and 4 April 1963, C. O. Handley, Jr. and F. M. Greenwell; 24, Armila (San Bias), 29 March and 1 April 1963, same collectors. REMARKS : A single specimen was labeled as from Vampyressa sp., Rio Mandinga (San Bias), GML, 29 May 1957. Uroderma bilobatum was col- 528 ECTOPARASITES OF PANAMA lected on the same day and it is possible that the specimen of dunni was a stray from Uroderma. Paratrichobius lowei Wenzel, new species. Figures 92B, 93. Superficially resembling P. dunni (Curran) in size and mesonotal chaeto- taxy and, like it, having spine-like setae along ventro-apical margin of tergum IX. It differs in having an irregular oblique double row of setae, the upper two or three minute but distinct, medial and parallel to the row of Fig. 91. Left side of tergum IX (of male hypopygium), ventral view of ventro-apical margin. A, Paratrichobius dunni (Curran) , from Uroderma bilobatum, Summit Gardens (Canal Zone). B, P. longicrus (M. Ribeiro), from Artibeus lituratus (same data as fig. 88A). heavy profemoral setae in dunni this row is incomplete and two or three of the upper setae are short spines. It also differs from dunni, and from all other known species of the genus, in that the usual profemoral spines are only strong setae. DESCRIPTION : Head. — Eyes with 25-27 facets. Laterovertices with six setae, all strong, except for a very short weak seta in postero-lateral angle. Occipital lobes with about six strong long marginal setae plus a. very long one (about as long as head) on antero-lateral lobule; posterior and ventral to these a row of six to eight shorter but strong subspiniform setae; one or two of marginal occipital setae are conspicuously longer macrosetae, one at medial edge, another one-third from inner margin. Postgenae on each side behind oral cavity with four or five short spinelets in a transverse row; posterior to the outer spinelets are two or three shorter, finer ones. Thorax. — Anterior margin of prescutum biconcave for recsption of occipital lobes, projecting at middle; median suture extending to about middle, transverse suture inter- rupted at middle; a spiracular and prespiracular bristle present; anterior half of prescutum with 13-14 additional stronger setae; basal half with similar setae laterally, setae becoming fine and dense at middle. Scutum with long strong setae along sides, WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 529 these continued medially as a W-shaped antescutellar row whose bristles are slightly shorter and finer medially, two or three at middle conspicuously shorter; scutum else- where with dense fine short setae. Mesepisternum above with 16-20 bristles. Median pleurotrochantinal lobe long, apically narrowed, sharply reflexed dorsally but not united with the metepimeron. Wings. — Rs varying in length from slightly less to slightly more than twice distance from fork to crossvein r-m; Ri with setae to base. Legs. — Inner (anterior) face of profemora with a diagonal row of six stout bristles rather than spines, ventral to these an irregular row of six or seven short setae, the basal one minute; ventral half of inner face with numerous short bristles. Bristles of sub- marginal row on outer face of hind femur distinctly projecting beyond ventral margin of femur throughout its length. Fig. 92. Male gonapophyses of species of Paratrichobius. A, left gonapophysis, P. dunni (Curran), from Uroderma bilobatum (no. 6591), Fort Clayton (Canal Zone). B, same, P. lowei, new species, paratype from Artibeus cinereus (no. 11428), Armila (San Bias). C, right (upper) and left (lower) gonapophyses, P. sanchezi, new species, paratype from Enchisthenes hartii (MA no. 389), Rancho Grande (Aragua), VENEZUELA. Abdomen. — Lateral lobes of tergum I+II each with three or four fine shorter bristles along inner dorsal margin; posterior margin with a double row of about 12-13 coarse bristles, those of anterior row longest; setae becoming shorter ventrally. Sternum I with three spine-like bristles on each lateral lobe. Sternum II with 30-34 discal setae in a median triangular area; posterior margin with 20-22 bristles, those near middle of about same length as discals, the outer ones a little longer. Dorsal connexivum bare, with four pairs of minute segmental setae; lateral and ventral connexivum covered with nearly uniform, short, conspicuous setae, these slightly longer basally on ventral con- nexivum. Mesepimeron (above) with 16-20 bristles. Female: Tergum VII transverse, its anterior margin slightly emarginate at middle ; with a pair of lateral macrosetae and between and slightly posterior to these a pair of short setae. Supra-anal plate with four apical macrosetae and on each side near mid-length, a short seta. Seventh sternites with 9—12 setae, one a conspicuously long macroseta. Male : Sternum V with setae of apical margin mostly of about the same size as discals, excepting a pair of macrosetae on each side. Hypopygium dorsally with a postero-lateral macroseta and a shorter seta on each side of sternum VII+VIII; tergum IX apically with a row of about six fine macrosetae, and on each inner ventro-apical margin a row of about five stout setae, the three middle ones spine-like; a short seta anterior to these. Gonapophyses as in fig. 92B. Measurements : Male Female BL 1.48-1.73 1.76-1.95 TL 0.36-0.38 0.41-0.44 WL 1.50-1.58 1.59-1.74 ww 0.63-0.66 0.66-0.71 FL 0.82-0.85 0.80-0.91 530 ECTOPARASITES OF PANAMA TYPE MATERIAL : Holotype male (slide) from Artibeus cinereus watsoni (host no. 11964), Armila (San Bias), 27 March 1964, Charles O. Handley and Francis M. Greenwell. Allotype female, same locality, collectors and host (no. 11678), 16 March 1963; in the collection of Chicago Natural Fig. 93. Thorax, dorsal view, Paratrichobius lowei, new species, holotype. History Museum. Paratypes. — 5, same data as the holotype, but 28 Febru- ary to 4 March 1963; 2, same host, Rio Tuira (Darien), 8 March 1958, P. Galindo [GML]. Paratypes in the collections of Chicago Natural History Museum, the United States National Museum, and the Gorgas Memorial Laboratory, Panama, Panama. REMARKS : This species is named for Mr. Wilbur Lowe, in appreciation of his valuable assistance in the laboratory of the Environmental Health Branch, at Corozal. Paratrichobius sanchezi Wenzel, new species. Figures 92C, 94. Similar in size and appearance to P. dunni but differing markedly in having a row of seven to eight short setae parallel to the stout spines on the inner face of the prof emur and in lacking stout spine-like setae along inner ventro-apical margins of tergum IX. Also resembling P. lowei n.sp., but WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 531 in lowei the usual prefemoral spines are represented by strong setae; as in dunni, the ventro-apical margins of tergum IX have spinelike setae. DESCRIPTION : Head. — Eyes large, with ±25 facets. Laterovertices with six setae, the most anterior ones very short, the most postero-medial seta as well developed as most others, but the seta immediately anterior to it a conspicuously long macroseta. Oc- cipital lobes narrow, posterior margin festooned with six stout marginal setae and, postero- ventral to these, five or six short spine-like setae; antero-laterally is a much Fig. 94. Thorax, dorsal view, Paratrichobius sanchezi, new species, paratype female from Enchisthenes hartii (MA no. 326), Rancho Grande (Aragua), VENEZUELA. longer macroseta that is conspicuously longer than the head. Postgenae, posterior to oral cavity, with one or two short black thorn-like setae on each side, and posterior to these, two others, followed by four or five fine stout setae. Thorax. — Prescutum about as long as broad; anterior margin biconcave for recep- tion of occipital lobes, slightly but distinctly produced at middle. Median suture extend- ing posteriorly beyond middle; transverse suture indistinct at middl2. Prescutum with eight or nine setae on each side of anterior half, the three antero-lateral bristles coarser and longer, one small bristle each side of median line on anterior projection; posterior half with two or three bristles along margin of longitudinal cleft, and about 10 in front of transverse suture, these becoming shorter and finer medially; anterior to these an irregular transverse band of 15-22 fine shorter setae. Scutum with a W-shaped row of 12-14 coarse bristles and anterior to these an irregular W-shaped row of 14-18 shorter bristles, similar to median setae in front of transverse suture. Mesepisternum with about 21 bristles in an irregular double row along dorsal and posterior margin, these coarser and longer along dorsal edge. Venter with scattered fine setae; a short angu- late pleurotrochantinal lobe present. Wings. — Rs about twice as long as distance from fork to r-m. All longitudinal veins except costa with extensive bare areas basally, those of fifth and sixth veins twice as long as the others. Profemora as in dunni but with a row of seven or eight short setae medial to and paralleling the oblique row of six stout spines. Hind femora and tibiae slightly bowed, the femora clothed with short semi- 532 ECTOPARASITES OF PANAMA recumbent setae except along inner face on basal third; about six conspicuously longer setae on dorsal margin and a lateral one near apex; ventral submarginal row projecting below margin and visible in profile for entire length. Abdomen. — Inner dorsal margins of lateral lobes of tergum II each with three or four fine setae, posterior margin with 9-12 coarse setae, the most dorsal of these longest. Connexivum bare dorsally along middle except for four pairs of well-developed segmental setae; lateral connexivum with very short setae, the ventral setae conspicuously longer. Sternum I with two or three spines on each side. Sternum II with 10-12 discal and ±14 marginal setae. Female: Tergum VII with a macroseta on each side and medial and posterior to these a pair of short setae. Supra-anal plate with four apical macro- setae and a short seta on each side near mid-length. Seventh sternites with 10-13 setae, most of them rather short, two or three much longer, one a conspicuously long macroseta. Male : Hypopygium with a macroseta on each side on sternum VII+VIII ; tergum IX fringed apically with nine or ten macrosetae; IX also with six or seven shorter setae ventro-laterally on each side. Gonapophyses as in fig. 92C, slender, sinuate above near apex. Measurements: BL TL WL ww FL Male 1.54-1.99 0.66-0.68 1.81-1.99 0.71-0.73 1.07-1.15 Female 1.70-1.78 0.60-0.67 1.90-2.03 0.71-0.80 1.08-1.17 TYPE MATERIAL: PANAMA. — Holotype male from Enchisthenes hartii (GML host no. 400,370), Cerro Mali (Darien), elevation 4800 feet, 6 June 1963, GML; allotype female same host (no. 9807-B), Cerro Hoya (Los San- tos), 8 February 1962, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratype. — A female, same data as the allotype, but 9 February 1962. VENEZUELA. — Paratypes, from Enchisthenes hartii: 2, Biological Sta- tion, Rancho Grande (Aragua), 1090 meters elevation, 10 August 1962, C. Machado and R. Antequera; 4, same locality but 21 August 1962, J. V. Scorza and C. Machado ; 8, same locality, 29 August 1962, J. V. Scorza and C. and A. J. Machado. Paratypes in the collections of the United States National Museum ; Chicago Natural History Museum ; Faculdad de Ciencias, Universidad Central de Venezuela; and the Gorgas Memorial Laboratory, Panama, Panama. REMARKS : This species is named for Mr. Pantaleon Sanchez, of the En- vironmental Health Branch at Corozal, in appreciation of his valuable as- sistance in the field work in Panama. Paratrichobius salvini Wenzel, new species. Figures 95, 96C. Similar to P. dunni (Curran) ; like it having only a couple of spinelets and a bristle medial and parallel to upper end of oblique row of profemoral spines, rather than a complete row of setae ; differing from dunni in having a longer thorax and less dense and extensive mesonotal chaetotaxy ; and in having normal rather than spine-like setae along ventro-apical margins of tergum IX. DESCRIPTION : Head. — Dorsum as in lowei; the seta in postero-median angle of latero- vertices very short, weak. Postgenae with a row of about five spinelets on each side behind oral cavity, the outer ones shorter; above and posterior to these another small spinelet and several fine short setae. Thorax. — Prescutum on anterior half with spiracular and prespiracular and about 12 other semi-erect or erect bristles, separated from semi-recumbent setae of posterior half by a bare transverse area ; setae at sides of posterior half longer and stronger than those at middle, which form a small dense patch. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 533 Scutum with an irregular W-shaped row of minute setae in front of the antescutellar row. Mesepisternum with about 14 bristles along dorsal margin. Pleurotrochantinal lobe long, tapered, sharply reflexed dorsally, not united with metepisternum. Wings. — Rs about twice as long as distance from fork to crossvein r-m. Legs. — Inner (anterior) face of profemora with a minute bristle and two short spinelets medial to upper end of Fig. 95. Thorax, dorsal view, Paratrichobius salvini, new species, female from Chiro- derma salvini (GML no. 300,079), Tacarcuna (Darien). oblique row of seven spines and another one or two very short but stout setae near apex of row; last spine of oblique row curved; inner face medial to these otherwise without setae except four or five very short ones near base. The hind femora not markedly swol- len basally; setae of ventral submarginal row very short and inconspicuous apically, not projecting beyond lower edge of femur. Abdomen. — Lateral lobes of tergum I+II with about three or four very fine setae on inner dorsal margin and a double row of about 12-14 coarse setae on posterior margin, these shorter ventrally. Connexival setae short, slightly longer ventrally; mid-dorsum bare except for four pairs of minute segmental setae. Female: Tergum VII transverse, narrowly elliptical; chaetotaxy of VII and supra-anal plate as described for lowei and sanchezi, the macrosetae fine. Sternum II with about 23-27 discal and 14-16 evenly dis- tributed marginal setae, these shorter posteriorly, both on disk and along margin; 534 ECTOPARASITES OF PANAMA seventh sternites with about 10 setae, one or two conspicuously long. Discal and lateral marginal setae a little heavier, a pair of long slender setae each side of middle, separated by a pair of short setae. Male: Sternum II with 14-28 discal and 18-26 marginal setae. Sternum V feebly sclerotized, only about half as wide as sternum II ; setae rather uni- formly short except for setae of apical margin which are slightly longer and become Fig. 96. Male gonapophyses of species of Paratrichobius. A, right (upper) and left (lower) gonapophyses of specimen from Chiroderma villosum jesupi (no. 12211), Puerto Obaldia (San Bias). B, same, of specimen from Vampyrops helleri (no. 10004), Cerro Hoya (Los Santos). C, right gonapophysis, Paratrichobius salvini, paratype from Chiroderma salvini (no. 12990), Cerro Tacarcuna (Darien). longer laterally, two or three long setae on each side near outer margin, one a conspicu- ously long macroseta. Hypopygium with a dorso-lateral macroseta and a short seta on each side of sternum VII+VIII ; tergum IX with an apical row of about eight macrosetae, four or five shorter setae along ventro-apical margin, and two others anterior to these. Measurements: BL TL WL ww FL Male 1.61-1.93 0.67-0.77 1.92-2.16 0.84-0.85 1.04-1.15 Female 2.06-2.32 0.63-0.80 2.19-2.32 0.85-0.88 1.26-1.32 TYPE MATERIAL: Holotype male and allotype female from Chiroderma salvini (host no. 12990), Cerro Tacarcuna (Darien), 8 March 1964, C. 0. Handley, Jr. In the collection of Chicago Natural History Museum. Para- types. — 1 male, same data as the holotype; 1 male, same data but 6 March 1964. In the collections of the United States National Museum and Chicago Natural History Museum. OTHER MATERIAL EXAMINED: 1 female, from Chiroderma salvini (?), Tacarcuna (Darien), elevation 2000 feet, 15 July 1963 [GML no. 300,079]. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 535 REMARKS : The female taken on 15 July 1963 is only tentatively associ- ated with this species, because we were unable to verify the identification of the host. The specimen differs from the allotype in possessing a pair of discal macrosetae at mid-length on the supra-anal plate. We have not seen females from Chiroderma villosum (see below) to determine whether or not they possess such setae. If the host is correctly identified, then this character is probably variable as it is in some other streblids. Paratrichobius sp. A. Figure 96A. This appears to be a distinct species with shorter male gonapophyses than those of P. salvini n.sp., but we are unwilling to name it until addi- tional material, including females, is available for detailed comparison with salvini. MATERIAL EXAMINED (From Chiroderma villosum jesupi) : 3 males, Puerto Obaldia (San Bias) , 3 and 4 April 1963, C. O. Handley, Jr. and F. M. Greenwell; 1 male, Armila (San Bias), 17 March 1963, same collectors. Paratrichobius sp. B. Figure 96B. This appears to be a distinct species, similar to species A, but with gonapophyseal macrosetae inserted farther distad. Again, we are unwill- ing to name this in the absence of more extensive comparative material. The occurrence on Vampyrops of a species related to dunni and salvini is in- teresting. All other specimens from Vampyrops (sp., dorsalis, and vittatus) that we have examined are related to Paratrichobiits longicrus. MATERIAL EXAMINED (From Vampyrops helleri) : 1 male, Cerro Hoya (Los Santos), 24 February 1962. PROVISIONAL KEY TO THE PANAMANIAN SPECIES OF PARATRICHOBIUS 1. Inner face of profemora with only two or three short spinelets and/or setae medial and parallel to upper end of oblique row of spines or heavy setae (figs. 90, 95) ... .2 Inner face of profemora with a complete row of setae medial and parallel to the oblique row of spines or strong setae (figs. 86, 93) 3 2. Mesonotal chaetotaxy as in fig. 90A. Male: Inner ventro-apical margins of tergum IX (of hypopygium) with spine-like setae (fig. 91A). Host: Uroderma bilo- batum dunni Curran Mesonotal chaetotaxy as in fig. 95. Male: Inner ventro-apical margins of tergum IX with normal setae as in P. longicrus (fig. 91B). Hosts: Chiroderma spp., Vampyrops helleri salvini n.sp. (and species A, B) 3. Inner face of profemora with seven spines or stout setae in a diagonal row, the apical one curved. Hind femora short, no more than 1.26 mm. long 4 Inner face of profemora with six stout spines, the apical one not curved. Hind femora longer, 1.32-1.77 mm. long longicrus complex 4. Inner face of profemora with stout spines. Mesonotal chaetotaxy as in fig. 94. Male: Inner ventro-apical margins of tergum IX with normal setae. Host: Enchisthenes hartii sanchezi n.sp. Inner face of profemora with strong setae, not spines, in diagonal row on inner face. Mesonotal chaetotaxy as in fig. 93. Male: Inner ventro-apical margins of tergum IX with stout spine-like setae as in dunni (fig. 91A). Host: Artibeus cinereus lowei n.sp. 536 ECTOPARASITES OF PANAMA Neotrichobius Wenzel and Aitken,11 new genus Type-species: Neotrichobius stenopterus Wenzel and Aitken, new species. Very similar to both Paratrichobius and Megistopoda in the greatly elongated hind legs and in the structure of the underside of the thorax, which, though much narrower, is reminiscent of that of the Nycteribiidae. Neotrichobius resembles Megistopoda in possessing stenopterous wings, but in that genus the body is strongly, laterally compressed; the occipital lobes of the head are strongly elevated; the profemora lack spines on the inner face ; and the paired ventral setae of the male gonapophyses are situ- ated anterior to middle rather than subapically. Neotrichobius is more closely related to and possibly derived from Paratrichobius, which it re- sembles inter alia in the structure of the male gonapophyses (the paired ventral setae are subapical in both) ; in having a row of stout spines or setae on inner face of profemora ; and in possessing strong spine-like setae on the dorsal sclerites of the head as well as on the theca and along sides of oral cavity. It differs from Paratrichobius in being stenopterous ; in having subcylindrical profemora ; and in that the dorsal abdominal connexivum is entirely covered with setae. Neotrichobius differs further from both Megistopoda and Paratrichobius in having the median suture bifurcate posteriorly; the scutum small and not separated laterally from the pre- scutum ; a very small scutellum ; and the mesopleuron traversed by a longi- tudinal membranous suture just dorsal to mesocoxal cavity. DIAGNOSIS: Head. — Very similar to that of Paratrichobius. Laterovertices and oc- cipital plates differentiated but much reduced in size and not strongly sclerotized; with strong spine-like setae. Palpi irregularly oval, ventral surface with a mixture of spines and normal setae; theca with spines and setae; sides of oral cavity margined with spine- like setae. Thorax. — Mesonotum longer than head. Anterior margin of prescutum pro- duced at middle, emarginate on each side of projection, to accommodate occipital lobes of head. Median suture bifurcate posteriorly. Scutum not separated from prescutum laterally. Scutellum very small. Each mesopleuron divided into dorsal and ventral regions by a second longitudinal membranous fissure similar to that which separates the prescutum and episternum. Underside of thorax shield-like, the anterior and posterior margins rounded, the anterior sternopleural margin dorsally bent upwards like a ski. Pleurotrochantines not projecting as an angulate process on each side between meso- and metacoxae. Wings. — Stenopterous, venation greatly reduced, only three longi- tudinal veins basally, two apically. Legs. — Middle and hindlegs progressively much longer than the short forelegs; hindlegs greatly elongated, the hind femora extending beyond apex of abdomen; all femora with some macrosetae, in addition to short setae; middle and hind femora slightly bowed; profemora very stout, subcylindrical (much as in Paradyschiria) , inner face armed with very heavy spines. Hind trochanters with spines along inner margin; hind coxae armed with spines above and below; hind tibiae bowed. Protarsi strongly compressed antero-posteriorly; middle and hind tarsi elon- gated, the first tarsomere twice as long as wide, as long as second and third combined. Abdomen. — Lateral lobes of tergum I+II with strong spine-like setae on postero- lateral margins. Sternum I with spine-like setae at sides. Sternum II medially with both spine-like and normal discal setae ; posterior margin, with spine-like setae, forming a pseudo-ctenidium. Connexivum entirely covered with setae, without a bare dorsal 11 Thomas H. G. Aitken, Trinidad Regional Virus Laboratory, Port-of-Spain, Trinidad. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 537 Fig. 97. Neotrichobiusstenopterus,n.gen.,n.sp. Female (allotype), dorsal view. median longitudinal area; setae of upper and ventral sides similar in both sexes; lateral setae of connexivum shorter. Female: Tergum VII not evident. Supra-anal plate very short. Cerci united with ventral arc. Seventh sternites very small. Male : Hypopygium not strongly developed, but distinct. Sternum V present, transverse. Sternum VI ab- sent. Gonapophyses as in Paratrichobius, the paired ventral setae subapical in position, the macroseta not strongly developed, the accessory seta anterior to macroseta. 538 ECTOPARASITES OF PANAMA Fig. 98. Neotrichobiiis stenopterus, n. gen., n. sp. Female (allotype), underside. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATPLIES 539 Neotrichobius stenopterus Wenzel and Aitken, new species. Figures 97-99. DESCRIPTION : Head. — More or less horizontal, longer than broad. Genae with a few short spine-like setae. Laterovertices microsetose, each with one macroseta, another seta about two-thirds as long, an intermediate-size seta, and two short setae; occipital plates not microsetose, with a very stout long macroseta and five very short spine-like setae; mediovertex microsetose, membranous. Posterior margin slightly emarginate at Fig. 99. Neotrichobius stenopterus, n. gen., n. sp. A, apex of abdomen, male (holotype) , ventral view. B, same, female (allotype). C, wing. middle for reception of median prescutal projection. Eyes relatively conspicuous, with ±11 facets. Palpi irregularly oval, apical macroseta very long, ventral surface with six to eight setae, most of them spine-like. Theca with a pair of fine setae at apex and two spines on each side on apical half. Margin of oral cavity bordered with spine-like setae. Thorax. — Prescutum with two supraspiracular bristles, one or two short discal setae anterior to middle, and a pair near posterior margin. Scutum with eight to ten short bristles, and a long bristle at each lateral angle. Scutellum with a median pair of very long bristles and a shorter bristle on each side. Mesepisternum, above, with two or three long and five or six short bristles. Wings and Legs as illustrated. Abdomen. — Tergum I+II with a transverse basal cluster of short setae; inner and posterior margins poorly defined, side margins of lateral lobes with a row of strong spine-like setae. Each lateral lobe of sternum I with two spine-like setae. Sternum II with two pairs of heavy spine-like setae and four or five short spine-like setae between them, followed on apical half by three to five short stout setae ; apical margin with six to eight stout spine-like setae, and a pair of slender setae. Connexivum similar in both sexes, rather uniformly covered throughout with moderately heavy, fairly short setae, 540 ECTOPARASITES OF PANAMA these shorter along sides, especially apically. Female (figs. 97, 98, 99B) : Supra-anal plate as illustrated. Seventh sternites small, chaetotaxy as figured. Male (fig. 99 A) : Hypopygium without dorsal setae ; a long slender ssta on apical margin on each side, and lateral to it a short seta and a heavy macroseta ; a short seta also present each side near base; ventral chaetotaxy as illustrated. Gonapophyses as in Paratrichobius, paired ventral setae subapical; macroseta only slightly longer than accessory seta. Measurements : BL TL Male 1.70-1.92 0.55-0.56 Female 2.03-2.09 0.49-0.58 TYPE MATERIAL: Holotype male and allotype female (slides) from Uro- derma bilobatum (host no. 5492), Almirante (Bocas del Toro) 28 January 1960, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — From Artibeus cinereus: 3 (2 bats), Rio Changena Camp (Bocas del Toro) , elevation 2600 feet, 18 September 1961 ; 1, Cerro Hoya (Los Santos), 12 February 1962; 5 (4 bats), Armila (San Bias), 24 to 27 February 1963, C. 0. Handley, Jr. and F. M. Greenwell. From Artibeus j. jamaicensis: 1, Cerro Hoya (Los Santos), 9 February 1962. From Vampyressa pusilla: 1, Armila (San Bias), 18 March 1963, C. 0. Handley, Jr. and F. M. Greenwell; 1, La Laguna (Darien), 17 June 1963, elevation 2900 feet, GML. From Noctilio leporinus mexicanus: 1, Guanico (Los Santos), 13 February 1962. Non-Panamanian paratypes. — From Artibeus cinereus: 1, Track to Tree Station, Rio Grande Forest, Sangre Grande, TRINIDAD, 30 January 1957, T. H. G. Aitken. From Phyllostomus hastatus: 1, Kaiserberg Airstrip, east of Zuid River, SURINAM, 16 October 1960, ele- vation 900 feet, H. A. Beatty, CNHM Guianan Zoological Expedition 1960- 1962. Paratypes to be deposited in the collections of Chicago Natural History Museum; United States National Museum; Trinidad Regional Virus Laboratory; Gorgas Memorial Laboratory, Panama, Panama; and the Environmental Health Branch at Corozal, Canal Zone. REMARKS : We believe the primary host of this species to be Artibeus cinereus. Eight of the 14 lots known to us are from this host. About one-third of the specimens collected are females. Although the host records from Artibeus, Uroderma, and Vampyressa seem to be valid, we are re- luctant to accept those from Noctilio and Phyllostomus. There were a num- ber of dubious host associations in the material from Los Santos. Much of this material was collected from hosts caught in mist nets. The record from Phyllostomus is one of several dubious host associations in the material col- lected in Surinam. Genus Megistopoda Macquart Megistopoda Macquart, 1852, Ann. Soc. Ent. Fr., (2), 10: 332. Kolenati, 1863 (Megistopodia, sic!), Hor. Soc. Ent. Ross., 2: 89. Bigot, 1885, Ann. Soc. Ent. Fr., (6), 5: 235 (as Nycteribiidae). Speiser, 1900, Arch. Naturg., 66A, Bd. I. pp. 32, 57 (char.), 62 (cit.), 65 (keyed). Coquillet, 1900, Proc. U. S. Nat. Mus., 37, art. 1719, p. 566. Aldrich, 1905, Smiths. Misc. Coll., 46: 657 (as Nycteribiidae). Wil- liston, 1906, Man. N. Am. Dipt., 3rd ed., p. 385. Speiser, 1907, Ent. News, 18: 104 (cit.). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet, Nictheroy, 5: 26 (keyed), 30 (cit.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 28. Stiles WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 541 and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 356 (status). Bequaert, 1940, Revista Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418 (keyed) ; 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Parasitology, 39: 315. Maa, 1965, Jour. Med. Ent., 1: (4), p. 385 (synonymizes P ter ellipsis') . Pterellipsis Coquillet, 1899, Can. Ent., 31: 333; 1910, Proc. U. S. Nat. Mus., 37, no. 1719, p. 597. Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 25 (keyed), 29 (cit.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 25. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth. Wash., no. 155, p. 657. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 357 ff. (morph.). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418 (keyed) ; 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Parasitology, 39: 321; 1952, ibidem, 42: 134, fig. 2 A. Grandi, 1952, Introd. Stud. Ent., 2: 478. Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 177 (keyed), 187. [Type species: P. aranea Coquillet, 1899]. Type-species: Megistopoda pilatei Macquart, 1852. It is remarkable that the identity of Megistopoda pilatei has remained unresolved since 1852, when Macquart described and figured this remarkable streblid from Mexico (Teapa). The absence of wings seemed to preclude its being identifiable with any other streblid known. Although the very sketchy illustrations showed a close resemblance to Pterellipsis, it was dif- ficult to accept it as such, because of the absence of wings. Though very much reduced in Pterellipsis, the wings are nevertheless quite conspicuous and would hardly have been overlooked (see Speiser, 1900a) . Indeed, some authors, e.g., Bigot (1885) and Kolenati (1963) placed Megistopoda in the family Nycteribiidae, not only because of the absence of wings, but because Macquart's figure illustrated the head as thrown back over the mesonotum, much as it is in members of that family. In Pterellipsis, the head is more vertical than in most other Streblidae, and the anterior portion of the mesonotum is somewhat declivious, so that with due allowances for de- ficiencies of Macquart's figures, these characters should not be given any serious consideration. In fact, Speiser, in describing desiderata (=Pterel- lipsis aranea Coquillet) assigned it to the genus Megistopoda and separated it from M. pilatei because it possessed wings. There is always the possibility that a wingless species might yet be found that will answer Macquart's description. During the last 27 years, the senior author has identified approximately twenty or twenty-five thousand streblids from the New World tropics, including several thousand from Mexico, Guatemala and El Salvador, without finding any wingless species that would answer to the description of M. pilatei. However, in the Panama collection, there is a "wingless" male specimen of Pterellipsis aranea Coquil- let from Artibeus jamaicensis (GML host no. 300,084) , Tacarcuna (Darien) , elevation 2000 feet, 20 June 1963. The wings are reduced to such minute vestiges that they cannot be detected except under very high magnifications (ca. X150) . This specimen would correspond closely to the description and figure of Megistopoda pilatei, and it is suggested that Macquart's type was such a wingless specimen of Pterellipsis. It is not at all surprising that wingless individuals would appear within a species which normally has vestigial wings. I believe it is reasonable to assume that the species P. 542 ECTOPARASITES OF PANAMA aranea Coquillet and M. pilatei Macquart are synonymous. Megistopoda proximo, (Seguy) and M. theodori n.sp., though closely related to aranea, differ inter alia in having noticeably shorter legs. The figure of Megisto- poda pilatei shows very long hind legs, as in aranea. One can reasonably argue that the identity of pilatei is not settled, but we believe there is no longer any doubt concerning the identity of the genus Megistopoda. We agree with Maa (1965, p. 385) in considering Pterellipsis to be a synonym. KEY TO THE SPECIES OF MEGISTOPODA 1. Wings narrow (fig. 100D) . Legs very long, hind femora 1.54-1.81 mm. long, as long as or nearly as long as entire body. Prescutum with very weak setae along me- dian suture ; the lateral margins along longitudinal membranous suture, with one or two stronger setae. Hosts: Artibeus jamaicensis, A. lituratus palmarum .... aranea (Coquillet) Wings broader (fig. 101E). Legs shorter, hind femora 0.99-1.43 mm. long, only a little longer than abdomen. Prescutum with short but strong setae basally along median suture, the lateral margin along longitudinal membranous suture usually with four strong setae. Hosts : Sturnira spp 2 2. Male: Gonapophyses (figs. 100C, 101D) with ventral margin nearly straight in lateral view, with a row of translucent thorn-like setae between macroseta and apex. Macroseta of right gonapophysis inserted at about basal third, that of left anterior to it. Host: Sturnira lilium proxima (Seguy) Male: Gonapophyses (fig. 100B) with ventral margin distinctly curved in lateral view; thorn-like setae few in number, restricted to lateral face and one on both dorsal and ventral margins apically; macrosetae of right gonapophysis inserted only slightly posterior to that of left, which is inserted well beyond middle. Host : Sturnira ludovici theodori n.sp. Megistopoda aranea (Coquillet). Figure 100A, D. Pterellipsis aranea Coquillet, 1899, Can. Ent., 31: 334 (U. S. National Museum, type no. 4208); 1900, Proc. U. S. Nat. Mus., 22: 270 (records). Speiser, 1900, Zool. Anz., 23: 154 (synonymizes Megistopoda desiderata Speiser under P. aranea.). Aldrich, 1905, Smiths. Misc. Coll., 46: 658. Coquillet, 1910, Proc. U. S. Nat. Mus., 37, no. 1719, p. 597 (as type-species of Pterellipsis). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 23 (records), 30 (cit.), pi. 2, fig. 3 (wing). Wolcott, 1923, Jour. Dept. Agric. Porto Rico, 7: 235. Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 25, pi. 2, fig. 15. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 477 (fig.), 479 (fig., Streblidae II.-5). Jobling, 1936, Parasitology, 28: 359 (wing structure). Cooper, 1941, Yearb., Amer. Phil. Soc., 1941: 126 (chromosome no.). Bequaert, 1942, Bol. Ent. Venez., 1: 88 (records). Jobling, 1949, Parasitology, 39, pp. 316 ff., 327, fig. 2C (with P. proxima Seguy as syn., in error) ; 1952, ibidem, 42: 134, fig. 2A. Grandi, 1952, Introd. Stud. Ent., 2: 478. Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 183, 187 (records), pi. 3, fig. 1. Megistopoda aranea Maa, 1965, Jour. Med. Ent., 1: 385. Megistopoda desiderata Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 37, 42, 57-59 (descr.), 63, 65 (keyed), pi. 3, figs 6-8. PANAMANIAN MATERIAL EXAMINED : 213 specimens in 88 lots from more than 104 bats. From Artibeus j. jamaicensis: 188 (76 lots) as follows: 9 (2 bats), Fort Clayton (Canal Zone), 19 and 20 October 1960; 7 (4 bats), Fort Kobbe (Canal Zone), 24 and 31 July, 12 and 27 October 1960; 16 (5 bats), Natural Bridge, Madden Dam (Canal Zone), 31 August 1959; 3, WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 543 Summit (Canal Zone), 5 August 1930, L. H. Dunn No. 6 [AMNH] ; 23 (4 lots), Barro Colorado Island (Canal Zone), 22 November 1956 and 3 to 12 January 1957 [USNM] ; 18, Miraflores (Canal Zone), K. W. Cooper [MCZ] ; 3, Arraijan (Canal Zone), 26 March 1961; 19 (7 bats), Almirante (Bocas del Toro), 23 January to 5 February 1960; 6, Isla Colon (Bocas del Toro), 18 February 1960; 4 (3 bats), Rio Changena Camp (Bocas del Toro), 19 to 27 September 1961 ; 12 (18 bats), Cerro Hoya (Los Santos), 11 to 24 Febru- ary 1962; 1, Armila (San Bias), 1 April 1963; 38 (9 bats), Punta de Pina (Darien), 23 to 24 March 1960; 5 (4 bats), Rio Seteganti (Darien), 1 to 5 February 1961 ; 10 (4 bats) , La Laguna (Darien) , 9 to 17 June 1963, GML ; 17 (12 bats), Tacarcuna (Darien), 20 June to 15 July 1963, GML; 3 (2 bats), Rio Mono Camp (Darien), 24 and 25 July 1963. From Artibeus lituratus palmar um, 4(4 lots) as follows : 1, Fort Clayton (Canal Zone), 20 October 1960; 1, Barro Colorado Island (Canal Zone), 6 December 1956 ; 1, Sibube (Bocas del Toro) , 25 January 1963 ; 1, La Laguna (Darien), 6 June 1963, 2900 feet elevation, GML. From Carollia perspicil- lata azteca: 3 (3 bats), Almirante (Bocas del Toro), 23 to 31 January 1960 (mist nets). From Desmodus rotundus murinus: 1, Almirante (Bocas del Toro) , 25 January 1960 (mist net) . From Phyllostomus discolor: 1, Tacar- cuna (Darien), 5 September 1958. From mixed hosts (Artibeus cinereus, A. j. jamaicensis, and Carollia subrufa) : 1, Rio Changena Camp (Bocas del Toro). Without host: 3 (2 lots) Barro Colorado Island (Canal Zone), 4 November 1956, K15 and K16 [USNM] . OTHER MATERIAL EXAMINED : Numerous specimens, chiefly from Artibeus jamaicensis, from PUERTO Rico, MEXICO, GUATEMALA, EL SALVADOR, COSTA RICA, COLOMBIA, VENEZUELA, SURINAM, and BRAZIL. REMARKS : This species appears to be a parasite primarily of Artibeus jamaicensis, though it has occasionally been taken on Artibeus lituratus palmarum. Records from other hosts for the most part probably represent contaminations, disturbance transfers, and errors of association, though there certainly are some that represent valid associations. M. aranea was taken from approximately 54.4% of all Artibeus jamaicensis that were para- sitized by Streblidae, but on only 7.69% of A. lituratus palmarum. Its ana- logue on that host seems to be Paratrichobius longicrus. Megistopoda proxima (Seguy) . Figures 100C, 101. Pterelhpsis proxima Seguy, 1926, Ency. Ent., (B), II, Dipt. 3: 194-196, figs. 2-6 — Argentina: "Misiones: environs de San Ignacio: Villa Lutecia," from an unidenti- fied bat (Museum National d'Histoire Naturelle, Paris). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Jobling, 1949, Parasitology, 39: 327 (as syn. of P. aranea Coquillet, in error) ; 1952, ibidem, 42: 134, 135, fig. 4B (re- stored to specific status). Megistopoda proximum Maa, 1965, Jour. Med. Ent., 1: 385. In the collection of Chicago Natural History Museum is a male specimen of Megistopoda, taken on Sturnira sp., 5 miles north of Therezopolis, Brazil. This fly appears to be identical with the species described as proxima by Seguy, as well as with specimens taken from Sturnira lilium parvidens in Panama. Although Seguy's illustration of the gonapophyses (fig. 101D) is 544 ECTOPARASITES OF PANAMA Fig. 100. A-C, left male gonapophysis of species of Megistopoda. A, M. aranea (Co- quillet), from Artibeus j. jamaicensis (no. 6110), Pina Point (Darien). B, M. theodori, new species, paratype from Sturnira ludovici (no. 6181), Casa Tilley (Chiriqui). C, M. proximo, (Seguy), from Sturnira lilium (no. 6351), Rio Mandinga (San Bias). D, wing, M. aranea (=desiderata Speiser), after Speiser (1900). made from quite a different view than ours (fig. 101 C) , it shows the macro- seta of the right gonapophysis inserted far posteriorly, as in our specimens, and also the characteristic row of short thorn-like setae along the ventral margin. Therefore, we assign our specimens to M. proximo,. Measurements: BL TL FL Male 1.84-1.98 0.44-0.49 1.07-1.18 Female 1.76-2.09 0.99-1.21 PANAMANIAN MATERIAL EXAMINED: 34 species in 18 lots from (18) Sturnira lilium parvidens as follows : 10 (5 bats) , Guanico (Los Santos) , 21 WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 545 January to 8 February 1962; 4 (2 bats) , Cerro Hoya (Los Santos), 24 and 26 February 1962; 1, Isla Bastimentos (Bocas del Toro), 3 February 1963. 2 (2 bats) , Rio Tuira, 25 February 1958, P. Galindo [GML] ; 1, Cerro Tacar- cuna (Darien), 3 March 1964, C. 0. Handley, Jr.; 3 (3 bats), Tacarcuna (Darien), 2000 feet elevation, 20 June and 7 July 1963, GML; 1, Armila (San Bias), 27 March 1963, C. 0. Handley, Jr. and F. M. Greenwell; 12 (4 bats), Rio Mandinga (San Bias), 26 to 30 May 1957, P. Galindo [GML]. REMARKS: This species appears to be restricted to Sturnira lilium, at least in Panama. We have seen a few specimens from this host, from Para- guay and Venezuela, which appear to be this species. M. proximo, oc- curred on approximately 86^0 of the Sturnira I. parvidens that were para- sitized by Streblidae (nearly 100%), and averaged two specimens per bat. Megistopoda theodori Wenzel, new species. Figure 100B. With the characters of M. proximo, (Seguy), and apparently differing from that species only in its somewhat larger size and in the structure of the male gonapophyses (fig. 100B) , which are very distinctive. In M. proxi- mo, (fig. 100C) both gonapophyses are very similar except that the paired ventral setae are inserted further basally on the right than on the left ; the ventral margin of the gonapophyses appear nearly straight in lateral view ; the paired ventral setae are inserted further posteriorly ; and a row of dis- tinct thorn-like setae (coarser than in theodori) is present along ventral margin. In theodori, the right gonapophysis is not sinuate along dorsal margin and thus appears heavier ; the ventral margins are distinctly curved in lateral view, the paired ventral setae are inserted at middle or beyond ; and the fine thorn-like setae are finer, fewer in number, and most are in- serted on the lateral face rather than the ventral margin. M. theodori also differs in host and altitudinal distribution. M. theodori was taken on Sturn- ira ludovici together with Trichobius brennani, at elevations of from 4800- 5600 feet, while M. proximo, occurred on Sturnira lilium, parvidens, to- gether with Aspidoptera delatorrei n.sp., at elevations ranging from sea level to 2000 feet. DESCRIPTION : Head. — Occipital lobes with eight to nine setae of varying lengths, two of them conspicuously long macrosetae. Laterovertices with about six setae. Eyes with ±7 facets. Prescutum on each side with two spiracular bristles and 15-23 others in- cluding about four strong setae along margin of longitudinal membranous cleft and posteriorly a patch of from five to seven shorter bristles along median suture. Scutum with 13-14 bristles, the anteromedian ones usually noticeably shorter than the others. Mesepimeron with 13-17 bristles in two irregular rows, the marginal ones mostly longer and coarser. Wings. — Broader than in aranea, and with venation as in proximo, (fig. 101E). Legs. — As in proximo,. Abdomen. — Lateral lobes of tergum I+II with from 13-20 bristles, the three or four along inner dorsal margin fine, the posterior ones long and coarse, shorter ventrally. Connexivum covered with relatively short, moderately heavy bristles, these much shorter along sides, a little longer apically. Sternum I with two or three spines on each side. Sternum II subquadrate, typically with 9-14 discal setae and on posterior margin five to eight spiniform setae on each side, separated by two pairs of finer setae, the outer pair long. Female: Tergum VII and supra-anal plate united at middle for part of their width ; VII transverse, with a conspicuous macrosetae at each side and one or two shorter ones between them ; supra-anal plate with four apical macrosetae and on each side a pair of fine short setae. Connexivum with a long subapical latero-ventral macroseta. Seventh 546 ECTOPARASITES OF PANAMA Fig. 101. Megistopoda proximo, (Seguy). A, male, lateral view. B, antenna. C, eye. D, male gonapophyses (spread out), ventral view. E, wing. From Seguy (1926). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 547 sternites with 11-15 setae, several of them macrosetae, one conspicuously longer than the others. Male: Hypopygium with two or three dorso-lateral macrosetae basally (on VII+VIII), about eight apical macrosetae and four or five latero- ventral setae on tergum IX. Sternum V with discal setae of about the same size as those of connexivum; apical margin with six to eight macrosetae, alternating with one or two shorter but strong setae, two to four shorter strong setae at middle. Gonapophyses distinctly curved in lateral view (ventral margin nearly straight in proximo,), dorsal margin of the left gonapophysis sinuate near apex ; paired ventral setae of left gonapophysis inserted distal to middle, those of right gonapophysis slightly posterior to them; right gonapophysis slightly heavier than left, without dorsal sinuation ; each gonapophysis, anterior to paired ventral setae, with several extremely fine translucent, thorn-like setae along lateral face and ventral margins (one also seems to be present on dorsal margin above paired setae, though only the theca is visible in specimens examined). Measurements: BL TL FL Male 1.98-2.16 0.49-0.52 1.13-1.43 Female 2.17-2.47 0.55 1.18-1.35 TYPE MATERIAL : Holotype male and allotype female from Sturnira ludo- vici (host no. 10469), Casa Tilley, Cerro Punta (Chiriqui), elevation 5300 feet, 14 March 1962, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — 37, same data as the holotype; 7 (29 bats), same locality but 6 to 11 March 1962 ; 79 (26 bats) , same locality, 23 April to 4 May 1960 ; 5 (2 bats), Lava Flow, El Hato (Chiriqui), 5 May 1960; 9 (5 bats), Casa Lewis, Cerro Punta (Chiriqui), 5600 feet elevation, 3 and 4 May 1960; 15, same locality, 1 and 2 February 1960; 9 (6 bats), Rancho Mojica (Bocas del Toro), elevation over 4800 feet, 8 and 9 September 1961; 2 (2 bats), Rancho Caballero (Bocas del Toro) , elevation 6000 feet, 11 September 1961 ; 1, Cerro Mali (Darien), 9 February 1964, C. 0. Handley, Jr.; 3 (2 bats), same locality, 4800 feet elevation, 31 May and 3 June 1963, GML ; 19 (4 bats) , Cerro Tacarcuna (Darien), 29 February to 7 March 1964, C. O. Handley, Jr. Paratypes to be deposited in the collections listed on p. 410. REMARKS: We have studied M. theodori n. sp. and M. proximo, (Seguy) in considerable detail, without finding any external characters that could be used for routine identification. In theodori, the number of mesonotal setae (excluding the spiraculars) was distinctly higher in the females (median 44) than in the males (median 38) ; in proximo,, the number was only slightly higher in the females (median 33) than in the males (32) . In theodori, the number of setae on the lateral lobes of tergum I +11 ranges from 13-17 (median 14.5) in the females and 15-20 (median 16.5) in the males; in proximo, it is nearly identical (14-19, median 17) in the two sexes. The number of setae on each of the female seventh sternites ranges from 11-16 (median 13.5) in theodori and from 8-17 (median 12) in proximo,. The sample measured included about thirty of each species. There is such a broad overlap in the above ranges that, while bimodality is evident, the setal counts cannot be used for identification of individual speci- mens, nor for small series. This species is named for Prof. Oskar Theodor, of the Hadassah Medical School, Hebrew University, Jerusalem, Israel, in recognition of his outstand- ing contributions, especially to our knowledge of the pupiparous flies and of the Phlebotominae (Psychodidae). 548 ECTOPARASITES OF PANAMA Fig. 102. Speiseria ambigua Kessel. A, female, dorsal view. B, posterior margin of head and anterior portion of sternopleura, ventral view. C, left male gonapophysis, specimen from Carollia perspicillata azteca (no. 3940), Fort Sherman (Canal Zone). A and B from Jobling (1939a). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 549 Genus Speiseria Kessel Speiseria Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 19 (diagn.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 656. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, pp. 6 (keyed), 7. Jobling, 1936, Parasitology, 28: 359 ff. (morphology). Pessoa and Guimaraes, 1937, Ann. Fac. Med. Sao Paulo, 12: 265. Jobling, 1939, loc. cit., 31: 487-488 (as syn. of Synthesiostrebla, in error). Bequaert, 1940, Rev. Ent. Venez., 1: 86 (keyed). Jobling, 1947, Proc. Roy. Ent. Soc. Lond., (B), 16: 39; 1949, Para- sitology, 39: 321 (keyed). Hoffmann, 1953, Mem. Cong. Cient. Mex., 7: 177 (keyed) , 188. Synthesiostrebla (not Townsend, 1913), Jobling, 1939, Parasitology, 31: 487. Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed). Schuurmans-Stekhoven Jr., 1951, Beitr. Fauna Perus 3: 96 (part.). Type-species : Speiseria ambigua Kessel, 1925. The genus that was keyed out under the name Speisseria [sic/] by Cur- ran (1934b, p. 522) was probably Pseudostrebla (see S. ambigua, below). Like the preceding three genera, Speiseria appears to be related to the spe- cies of the Trichobius phyllostomae group and is similar in the female genital characters (See Paratrichobius) . However, the males differ from all of these in having the accessory gonapophyseal setae inserted posterior rather than anterior to the macrosetae. On the anterior face of the profemora (fig. 102) is a row of strong setae, which appear to be homologous with the spines of Paratrichobius and Neotrichobius. Speiseria ambigua Kessel. Figure 102. Speiseria ambigua Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 20, pi. 1 (figs. 1, 2). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 656. Curran, 1935, Amer. Mus. Novit., no. 765, p. 7. Pessoa and Guimaraes, 1937, Ann. Fac. Med. Sao Paulo, 12: 265, fig. 17 (in legend as fig. 16, in error). Cooper, 1941, Yearb. Amer. Phil. Soc., 1941: 126 (chromosome no.). Bequaert, 1942, Bol. Ent. Venez., 1: 88 (syn. notes). Jobling, 1947, Proc. Roy. Ent. Soc. Lond., (B), 16: p. 39; 1947, Parasitology, 39: 315 ff., 327, fig. 2B. Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 183, 188. Synthesiostrebla amorphochili (not Townsend, 1913), Jobling, 1939, Parasitology, 31: 488-489. Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418. Schuurmans-Stekhoven, Jr., 1951, Beitr. Fauna Perus, 3: 96 (part.). Paratrichobius anduzei Matheson, 1945, Jour. Parasit., 31: 191, fig. la-e. Jobling, 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 144 (synonymizes P. anduzei). The record of Curran (1934b, p. 523) was probably of a species of Pseudostrebla rather than of Speiseria ambigua. His specimen was from Tonatia bidens, host of a species of Pseudostrebla. To our knowledge Speiseria ambigua has never been taken on bats of the genus Tonatia. PANAMANIAN MATERIAL EXAMINED: From Carollia p. azteca: 395 speci- mens in 139 lots, from more than 391 bats, as follows: 7 (3 bats), Camp Chagres, Madden Dam (Canal Zone), 20 to 25 June 1963, GML: 4 (2 bats), Camp Pifia (Canal Zone), 11 and 26 October 1960; 2 (2 bats), bridge, Coco Solo (Canal Zone) ; 5 (4 bats), culvert, Gatun Tank Farm (Canal Zone), 19 November 1959; 1, culvert, Brazoa Brook, Navy Tank Farm (Canal Zone), 6 October 1959; 50 (28 bats), mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 September 1959 and 24 (18 bats), same locality, 25 November 1959; 1, Natural Bridge, Madden Dam (Canal Zone), 31 August 550 ECTOPARASITES OF PANAMA 1959; 20 (12 bats), air-raid shelter, Fort Davis (Canal Zone), 6 October 1959, and 5(6 bats) , same locality and site, 29 October 1959 ; 1, Fort Gulick, 15 October 1959; 2 (2 lots), Battery Pratt, Fort Sherman (Canal Zone), 16 March 1960 ; 6, same data but 30 July 1959 ; 3, same data but 23 Novem- ber 1959; 1, Battery Kilpatrick, same locality, 22 October 1959; and 34 (30 bats), from bats in cativo trees same locality, 2 and 4 December 1959; 6, tunnel, runway, France Field, 25 August 1959 ; 1, culvert, same locality, 22 September 1959 ; 1, magazine, same locality, 8 October 1959 ; 3 (3 bats) , Gold Hill, same locality, 20 October 1959; 4 (34 bats), culvert, same locality, 23 November 1959, and 1 (3 bats), same locality, 30 November 1959; 27 (34 bats), Sardanillo Caves, Summit (Canal Zone), 12 August 1961, C. Yunker [RML] ; 4 (2 lots) , Summit, 17 April 1957, C. B. Koford [USNM] ; 1, Gamboa (Canal Zone), 1 May 1957, C. B. Koford [USNM] ; 1, RR culvert, Paraiso (Canal Zone) , 16 September 1959 ; 17 (19 bats) , RR culvert, east of Summit Golf Club (Canal Zone), 26 October 1959; 5, hollow tree, 1 mile from Gatuncillo (?Canal Zone), 28 July 1960; 1, culvert, Chepo Road (Panama), 12 October 1959; 2, air-raid shelter, La Chorrera (Panama), 18 September 1959; 9 (29 bats), Chilibrillo Caves (Panama), 18 August 1959; 33, same locality, K. W. Cooper [MCZ] ; 4, same locality, 27 August 1944 [USNM] ; 5 (22 bats) , Buena Vista Caves (Colon) , 3 September 1959 ; 5 (4 bats) , same locality, 15 September 1959 ; 32 (20 bats) , same locality, 24 November 1959 ; 1, Porto Bello (Colon), 7 August 1923 [USNM] ; 11 (10 bats), Guanico (Los Santos), 12 January to 13 February 1962; 1, Cerro Hoya (Los Santos), 25 February 1962; 9 (5 bats), Cocos Point, Isla del Rey (Darien), 22 and 24 March 1960; 2, Rio Chucunaque (Darien), 19 February 1958, P. Galindo, [GML] ; 5 (3 lots), Rio Tuira (Darien), 25 February to 8 March 1958, P. Galindo [GML] ; 1, Tacarcuna Village (Darien), 2 September 1958, P. Galindo [GML] ; 32 (13 bats), Almirante (Bocas del Toro), 24 January to 1 February 1960; 2, "Panama," L. H. Dunn [MCZ]. From Carollia castanea: 3 (4 bats) , Sibube (Bocas del Toro) , 21 to 26 January 1963, C. 0. Handley, Jr.; 3, Rio Chucunaque (Darien), 17 February 1958, P. Galindo [GML]. From Carollia subrufa: 1, Rio Changena Camp (Bocas del Toro), elevation 2300 feet, 21 September 1961; 1, Sibube (Bocas del Toro), 23 January 1963, C. 0. Handley, Jr.; 1, Isla Bastimentos (Bocas del Toro), 3 February 1963, C. 0. Handley, Jr., and F. M. Greenwell; 1, Armila (San Bias), 19 March 1963, same collectors. From Carollia sp. : 2, Cerro Azul (Panama), elevation 2000 feet, 27 January 1959, E. Mendez [GML] ; 1, Fort Clayton (Canal Zone), 9 January 1953, F. Blanton. From mixed collections of Carollia p. azteca and Glossophaga s. leachii: 3, Huile (Panama), 24 October 1960; 2, San Lorenzo Cave, Fort Sherman (Canal Zone), 15 March 1961. From Desmodus rotundus murinus: 5 (2 bats), Guanico (Los Santos), 26 January 1962. From Glossophaga s. leachii: 1, Armila (San Bias), 5 March 1963, C. O. Handley, Jr. and F. M. Greenwell ; 1, Almirante (Bocas del Toro) , 28 January 1960 ; 1, cativo trees, Fort Sherman (Canal Zone), 4 December 1959. From Lonchophylla robusta: 1, Buena Vista Cave (Colon), 16 June 1960. From Lonchorhina aurita: 1, mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 551 September 1959, and 3 (2 lots, 10 bats), same locality, 24 and 25 November 1959; 4, two miles north of Summit (Canal Zone), 17 April 1957 [USNM]. From Natalus stramineus mexicanus: 1, Fort San Lorenzo, Fort Sherman (Canal Zone) , 17 March 1960 ; 1, San Lorenzo Cave (Canal Zone) , 15 March 1961. From Phyllostomus hastatus panamensis: 1, Guanico (Los Santos), 26 January 1962; 4 (20 bats), Chilibrillo Caves (Panama), 17 July 1959. From Pteronotus parnellii fuscus: 1 (3 bats), Buena Vista Cave (Colon), 3 September 1959; 2, Almirante (Bocas del Toro), 29 January 1960; 1 (4 bats) , mine shaft, Coco Plantation, Gamboa Road (Canal Zone) , 25 Novem- ber 1959. From Trachops cirrhosus: 1, culvert, Chepo Road (Panama), 12 October 1959; 1, Battery Pratt, Fort Sherman, 23 November 1959; 1 (2 bats), Cerro Hoya (Los Santos), 15 February 1962. From Vampyrops vittatus: 1, Rio Changena Camp (Bocas del Toro), elevation 2500 feet, 27 September 1961. Without host: 1, railroad culvert at tower 34-12H, Sum- mit (Canal Zone), L. H. Dunn 181A [MCZ] ; 8, Chilibrillo Caves (Panama), L. H. Dunn 277 [MCZ] ; 2, same locality, no further data [cu] ; 1, culvert between Miraflores and Pedro Manuel, W side of Canal (Canal Zone) , L. H. Dunn [MCZ]; 2, Barro Colorado Island (Canal Zone), 2 December 1956, Koford no. 19 [USNM]. REMARKS: Speiseria ambigua is an unusually excitable species and leaves the host readily when disturbed. Genus Aspidoptera Coquillet Aspidoptera Coquillet, 1899, Can. Ent., 31: 334. Speiser, 1907, Ent. News, 18: 104 (synonymizes Lepopteryx Speiser). Williston, 1908, Man. N. Am. Dipt., 3rd ed., p. 385. Coquillet, 1910, Proc. U. S. Nat. Mus., 37, no. 1719, p. 511 (type-species). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 25 (keyed), 28 (cit.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 25, 26 (char., key to spp.) . Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 55, p. 658. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 357 ff. (morph., syst. pos.). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418 (keyed); 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 135-144 (rev.) ; 1949, Parasitology, 39: 322 (keyed) ; 1951, Trans. Roy. Ent. Soc. Lond., 102: 216 (morph.). Hoffmann, 1953, Mem. Cong. Cient. Mex., 7: 177 (keyed). Lepopteryx Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 42, 53 (diagn.), 62, 65 (keyed). [Type-species: Lipoptena phyllostomatis Perty]. Lipoptena (not Nitsch, 1818), Kolenati, 1863, Hor. Soc. Ent. Ross., 2: 103 (keyed). Bigot, 1885, Ann. Soc. Ent. Fr., (6), 5: 226 (part). Type-species: Aspidoptera busckii Coquillet, 1899. Jobling (1949a) recognized four species of Aspidoptera, namely, megastigma Speiser, minuta Costa Lima, clovisi Pessoa and Guimaraes, and phyllostomatis (Perty) . It has long been apparent to the senior author that these were convergent types representing the independent evolution of small size and reduced wings. We have re-assigned the first three to new genera, Mastoptera, Exastinion, and Noctiliostrebla, respectively. Only one of these (Exastinion) appears to be monotypic. As restricted by us, Aspidoptera now includes those minute brachyp- terous species with the following characters : 552 ECTOPARASITES OF PANAMA DIAGNOSIS: Head. — Laterovertices and occipital plates well differentiated. Latero- vertices without a longitudinal or oblique dark suture; occipital plates rounded poste- riorly, without a posterior lobe or flap. Thorax. — With a complete median suture which is united with the transverse mesonotal suture. Longitudinal and vertical membranous clefts open. Sternopleura not strongly produced anteriorly, sides oblique. Pleurotro- chantines with a blunt reflexed median lobe, this not united with the metepimeron. Legs. — Subequal. Wings. — Brachypterous but with essentially complete venation. Abdomen. — Tergum I+II and sternum II without any distinctive characters. Female: Tergum Fig. 103. Aspidoptera phyllostomatis (Perty). A, male, dorsal view. B, labium. C, wing. From Jobling (1949a). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 553 VII very weakly sclerotized, with a transverse row of two to four short setae. Supra- anal plate short. Seventh sternites small. Cerci free, not united with ventral arc. Male : Sterna V and VI absent. Gonapophyses curved laterally and downwardly at apex; ac- cessory seta inserted posterior to the macroseta. We tentatively segregate four species of Aspidoptera: busckii Coquil- let, phyllostomatis (Perty) (sensu Jobling 1949a), delatorrei n. sp., and an undescribed species from Brazil. Two of these are known to us from Panama. Aspidoptera phyllostomatis (Perty). Figures 103, 104E. Lipoptena phyllostomatis Perty, 1833 (?), Delect. Anim. Artie., fasic. 3, p. 190, pi. 37, fig. 16 — "in Phyllostomatis specie brasiliensis, indeterminata" (type ap- parently lost). Westwood, 1840, Introd. Mod. Class. Ins., 2: 585. Kolenati, 1856, Parasiten d. Chiropteren, p. 48, pi. 4, fig. 48a, b; 1863, Hor. Soc. Ent. Ross, 2: 98 (notes), 103 (keyed). Leptotena phyllostomatis Macquart, 1835, Hist. Nat. Ins. Dipt., 2: 645. Lepopteryx phyllostomatis Speiser, 1900 (part., female), Arch. Naturg., 66A, Bd. I, pp. 32, 53-54 (descr.), 62, 65 (keyed), pi. 3, figs. 3, 4. Aspidoptera phyllostomatis Speiser, 1900 (part.), Zool. Anz., 23: 153 (synonymizes A. busckii Coquillet). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 20 (discuss.), 22 (keyed), 28 (cit.), pi. 1, fig. 4 (wing). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Jobling, 1936, Para- sitology, 28: 359 (wing struct.); 1949, ibidem, 39: 315 ff., fig. 1G; 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 136 (keyed), 137 (redescr.), fig. 1 (with busckii Coq. as synonym) . The type of Lipoptena phyllostomatis Perty, 1833, is apparently lost. Presumably, it was deposited in the Munich Museum (Zoologische Samm- lung des Bayerischen Staates) along with the other insects Perty described from the material collected in Brazil by Spix and Martius. Speiser (1900a, p. 53) inquired about the type and was told that it was lost. Dr. T. C. Maa (pers. comm.) has informed us that he saw nothing in European collections that could be regarded as the type, though he had not studied either the collection at Munich or the Perty collection at the Natural History Museum in Bern. At our request, Dr. W. Forster of the Munich Museum has kindly searched for the type again without success. He wrote (pers. comm.) : "Leider wurde in friiheren Zeiten das nicht als solches bezeichnete Typen-Material teilweise in der Schausammlung gezeigt, sodass auf diesem Wege manche Verluste eingetreten sind. Auch ein erheblicher Teil der Perty'schen Typen ging damals zugrunde. Ich glaube also, Sie kb'nnen, falls notig, es ohne weiteres verantworten einer Neotypus aufzustellen und zu publizieren." Dr. H. D. Volkart, Curator of Invertebrates at the Natural History Mu- seum at Bern searched for the type in the collection of that institution with- out success. He wrote : "Der Typus von Lipoptena phyllostomatis Perty befindet sich leider nicht in unseren Sammlungen. Wohl ist die Sammlung Perty hier im Museum, aber sie kamseinerseit leider in schlecht erhaltenem Zustand hierher und ist heute nicht mehr vollstandig. Es ist daher nicht mehr moglich, festzustellen, ob sich der Typus von phyllostomatis in Perty's personlicher Sammlung befand; wir besitzen heute kein Exemplar dieser Art." We agree with Dr. Forster that a neotype should be established. Un- fortunately, there is little basis for interpreting Perty's phyllostomatis. 554 ECTOPARASITES OF PANAMA The type locality is Brazil, the type host "in phyllostomatis specie brasil- iensis indeterminata." The name Phyllostomus was applied to many bats of the family Phyllostomidae even until the early Twentieth Century. Bats of the genus Phyllostomus are parasitized by Mastoptera minuta (Costa Fig. 104. A, B, terminal cone, abdomen, female. A, Aspidoptera busckii Coquillet, from Artibeus 1. palmarum (no. 9959) Cerro Hoya (Los Santos). B, A. delatorrei, new species, allotype. C-E, left male gonapophysis. C, A. busckii Coquillet, from Artibeus j. jamaicensis (5512), Almirante (Bocas del Toro). D, A. delatorrei, new species, holotype. E, A. phyllostomatis (Perty), from "Phyllostoma" sp., Humboldt, (Santa Catharina), BRAZIL. Lima) in Brazil (see "Remarks," p. 516). However, Perty's illustration, poor as it is, could hardly be of that species, but rather of a fly more nearly like that interpreted as phyllostomatis by both Speiser (1900) and Jobling (1949a). We follow Jobling's interpretation of phyllostomatis. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 555 Jobling's description and figure are based on a slide-mounted specimen collected from Phyllostomus sp. at Humboldt, Sta. Catharina, Brazil, by Ehrhardt. The host was obviously not a Phyllostomus. Jobling's specimen was from one of several lots, apparently collected by Ehrhardt from the same host species, which contained Trichobius phyllostomae Kessel (includ- ing the holotype) and a species of Megistopoda, as well as the Aspidoptera. This assemblage indicates that the host was either an Artibeus or a species of Sturnira. A specimen of T. phyllostomae in the collection of Chicago Natural History Museum was taken from "Artibeus literatus jamaicensis" at Joinville (Santa Catharina). A new species of the phyllostomae group but more closely related to brennani n.sp. (from Sturnira ludovici) than to phyllostomae was taken from Sturnira sp. at Therezopolis, BRAZIL, with a species of Aspidoptera that is related to delatorrei n.sp. (from Sturnira lilium) , rather than to phyllostomatis, sensu Jobling. It seems likely, then, that the host of Jobling's phyllostomatis was an Artibeus. We designate as the neotype of phyllostomatis the specimen (noted above) upon which Jobling based his description and illustration, in the collection of the British Museum (Natural History). We have seen an additional specimen, from the same host and locality, which was kindly sent to us by Dr. Harold Oldroyd of that museum. Jobling's figure (fig. 103) indicates that the dorsal surface of the mesepisterna have uniform, rather short setae, but in the Humboldt speci- men before us, the outer (lower) setae are short and the inner (upper) setae are long. When specimens are studied and illustrated from slide prepara- tions, the apparent length of the setae is often deceptive because of fore- shortening. As noted by Maa (1963), Westwood (1840, vol. 1, p. 585) refers Lipoptena phyllostomatis to Nitzsch ("Voyage Prince Maximilian of Bavaria") as well as to Perty. We have been unable to find the Nitzsch reference in any bibliographic sources, and thus cannot determine whether or not his use of the name antedates that of Perty. Aspidoptera busckii Coquillet. Figure 104A, C. Aspidoptera buskii Coquillet, 1899, Can. Ent., 31: 335 — "Bayamon, Porto Rico," from Artibeus sp. (U. S. National Museum, no. 4210) ; 1900, Proc. U. S. Nat. Mus., 22: 270 (record). Aldrich, 1905, Smiths. Misc. Coll., 46: 657. Coquillet, 1910, Proc. U. S. Nat. Mus., 37, no. 1719, p. 511 (as type-species of Aspidoptera). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 22 (status), 28 (cit.). Wolcott, 1923, Jour. Dept. Agric. Porto Rico, 7: 235. Kessel, 1925, Jour. N. Y. Ent. Soc., 33:26. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Jobling, 1936, Parasitology, 28: 359 (wing characters); 1949, Para- sitology, 39: 315 (syn. note). Cooper, 1941, Yearb. Amer. Phil. Soc., 1941: 126 (chromosome no.). Grandi, 1952, Introduz. Stud. Ent., 2: fig. 439. Lepopteryx phyllostomatis (not Perty, 1833), Speiser, 1900 (part.,), Arch. Naturg. 66A, Bd. I, pp. 32, 53 (keyed), pi. 3, fig. 3. Aspidoptera phyllostomatis Speiser, 1900 (part.), Zool. Anz., 23: 153 (with busckii as synonym) ; 1907, Ent. News, 18: 104. Jobling, 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 138 (part., Antillean records only). The name busckii is revived for the species of Aspidoptera that para- sitizes Artibeus jamaicensis and A. lituratus as well as bats of some closely 556 ECTOPARASITES OF PANAMA related genera, in the West Indies, Central America, and northwestern South America. We are unable at this time to give characters that will separate females of phyllostomatis and busckii. The males appear to be distinct in the structure of the gonapophyses (fig. 104C, E). The speci- mens recorded below from Vampyressa and Chiroderma appear to be the same species as from Artibeus jamaicensis and A. lituratus. Specimens from Panama seem to be identical with those on Artibeus jamaicensis in the West Indies. This is interesting, because Artibeus jamaicensis parvipes (Greater Antilles) and A. j. yucatanicus (in Mexico, Guatemala) are para- sitized by an undescribed species of Trichobius, while Artibeus j. jamaicen- sis in Panama is not. A few additional characters that apply to both busckii and phyllostomatis, not included in Jobling's description, follow. A heavily pigmented species. Head. — Ventral face of palpi with sparse setae on a little more than basal half. Postgenae with a row of rather heavy setae dorsal and par- allel to ventral margin of oral cavity, dorsal to these, a row of very short setae. Thorax. — Spiracles large. Sutures heavy, transverse suture nearly straight. Mesepisterna dorsally with several rows of setae, the outer ones conspicuously shorter. Abdomen. — Lateral lobes of tergum I+II with two longitudinal rows of about six heavy long bristles each, the most posterior ones of the inner (marginal) row shorter; ventral to these a group of seven to nine shorter bristles. Female: Supra-anal plate with six apical or subapical macrosetae and on each side near mid-length a short sub- marginal bristle. Seventh sternites small, roundly oval, with nine to ten setae. Male: Gonapophyses as shown in fig. 104C. Measurements : BL TL Male 1.85-2.03 0.56-0.60 Female 2.03-2.25 0.60-0.63 PANAMANIAN MATERIAL EXAMINED : From Artibeus j. jamaicensis: 34 (32 bats), Almirante (Bocas del Toro), 22 January, 18 February 1960; 17, Rio del Puente, Natural Bridge, Madden Dam (Canal Zone) , 31 August 1959 ; 1, same locality, 5 April 1933, L. H. Dunn no. 25 [AMNH] ; 1, cave, west side of Taboga Island (Panama), 24 September 1959; 20 (8 bats), Pifia Point (Darien) , 24 March 1960 ; 6 (3 bats) , Rio Seteganti (Darien) , 4-5 February 1961, P. Galindo [GML] ; 3 (1 bat), Rio Tuira (Darien), 10 March 1958, P. Galindo [GML]; 13 (11 bats), La Laguna (Darien), 2900 feet elevation, 2-15 June 1963; 21 (12 bats), Tacarcuna (Darien), 2000 feet elevation, 20 June to 12 July 1963, GML; 14 (9 bats), Cerro Hoya (Los Santos), 8 to 16 February 1962 ; 1, Guanico (Los Santos) , 12 January 1962. From Artibeus lituratus palmarum: 1, Summit (Canal Zone), K. W. Cooper [MCZ]. 3 (3 lots), Barro Colorado Island (Canal Zone), 1 lot each, 4 November 1956, 6 December 1956, and 3 January 1957 [USNM] ; 1, Cerro Hoya (Los Santos), 21 February 1962; 1, Rio Mandinga (San Bias), 15 May 1957, P. Galindo [GML]. From Chiroderma villosum -jesupi: 1, Almirante (Bocas del Toro) , 24 January 1960. From Vampyressa nymphaea: 4(1 bat) , Rio Mono Camp (Darien) 25 July 1963, GML. From Carollia p. azteca (mist net) : 2, Almirante (Bocas del Toro) , 26 and 31 January 1960. From Phyllostomus discolor (mist net) : 1, Almirante (Bocas del Toro), 30 January 1960. NON-PANAMANIAN MATERIAL EXAMINED: From Artibeus jamaicensis parvipes: 1, CUBA, Dethier [MCZ]. From Artibeus jamaicensis: about a dozen lots from various localities in MEXICO, GUATEMALA and COLOMBIA. WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 557 REMARKS : We have seen numerous specimens that are either busckii or phyllostomatis from Venezuela and Peru. We have not studied them suf- ficiently to identify them at this time. Aspidoptera delatorrei Wenzel, new species. Figure 104B, D. Similar to A. phyllostomatis (Perty) , and busckii Coquillet but differing from both in that the bristles of all rows on the dorsal surface of the mesepisterna are long (setae of outer rows short in busckii and phyllosto- matis) ; the supra-anal plate (female) has a transverse row of four to six shorter setae across middle (fig. 104B), rather than only one on each side, in addition to the apical macrosetae ; and the short dorsal bristle of the male gonapophyses is inserted near apex rather than near middle. DESCRIPTION : Smaller, more lightly pigmented, and on the whole more densely setose than busckii Coquillet and phyllostomatis (Perty). Head. — Ventral surface of palpi with numerous short setae distributed from base nearly to apex. Genae densely setose, bristles along dorsal margin long, gradually shorter ventrad; postgenae with bristles along margin of oral cavity; parallel and dorsal to these are two or three rows of sub- equal bristles. Thorax. — Spiracles small. Transverse suture distinctly sinuate on each side. Dorsal surface of mesepisterna with three longitudinal rows of long subequal bristles. Abdomen. — Tergum I+II with 23-24 bristles arranged in three or four irregular longitudinal rows, most of the bristles long, but the most anterior bristle of upper row short, as are two or three ventral ones. Female: Tergum VII very small, feebly sclerotized and indistinct, typically with four, sometimes three, bristles. Supra-anal plate usually with six (rarely, four or five) apical or subapical macrosetae and anterior to these, a transverse row of five or six shorter setae; the latter often divided into two lateral groups by the foreward displacement of the median pair of subapical macrosetae. Measurements : BL TL Male 1.65-1.90 0.48-0.49 Female 1.70-2.36 0.49 TYPE MATERIAL (from Sturnira lilium parvidens) : Holotype male and allotype female (slides, host no. 9406), Guanico (Los Santos), 13 February 1962, C.M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — 1, same data as holotypes; 3, same locality but 8 February 1962; 3 (2 bats), Rio Tuira (Darien), 25 February and 10 March 1958, P. Galindo [GML] ; 5 (2 bats), Rio Mandinga (San Bias), 26 and 27 May 1957, P. Galindo [GML]. Non-Panamanian paratypes. — 3, Finca El Zapote, Zapote (Escuintla), GUATEMALA, 2400 feet elevation, 14 July 1948, R. D. Mitchell and Luis de la Torre, CNHM Guatemala Zoological Expedition (1948) ; 2, La Catarina, 2 miles north of Ciudad Guzman (Ja- lisco) , MEXICO, 9 December 1954, L. de la Torre, CNHM Mexican Zoological Field Trip (1954) . Paratypes in the collections of Chicago Natural History Museum; the United States National Museum; Gorgas Memorial Labora- tory, Panama, Panama; Environmental Health Branch (U. S. Army) at Corozal (Canal Zone) ; Departmento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil ; Faculdad de Ciencias, Universidad Central de Venezuela ; and Bernice P. Bishop Museum, Honolulu. OTHER MATERIAL EXAMINED: From Corollia p. azteca: 1, Rio Tuira (Darien), 5 March 1958, P. Galindo [GML]. 558 ECTOPARASITES OF PANAMA REMARKS: This species is named for Dr. Luis de la Torre, College of Pharmacy, University of Illinois, Chicago. Dr. de la Torre has added much to our knowledge of ectoparasites of mammals, especially bats, through his collecting in Mexico and Guatemala. KEY TO THE PANAMANIAN SPECIES OF ASPIDOPTERA All dorsal setae of mesepisterna long. Tergum I+II with three or four rows of conspicuous, long bristles, in addition to the short bristles present near inner anterior margin and ventrally. Female: Supra-anal plate (fig. 104B) with five or six very long apical or subapical setae and a transverse row of six shorter setae across middle, this row sometimes separated into two lateral groups of three setae each by an anterior displacement of the median pair of macrosetae. Male: Gonapophyses as in fig. 104D delatorrei n. sp. Outer setae of dorsal surface of mesepisterna short. Tergum I+II with only two rows of long bristles, in addition to a group of short ventral bristles. Female : Supra-anal plate (fig. 104A) with five or six apical macrosetae and, on each side near mid-length, a short submarginal seta. Male: Gonapophyses as in fig. 104C busckii Coquillet Exastinion Wenzel, new genus Type-species : Aspidoptera clovisi Pessoa and Guimaraes, 1936. Resembling Aspidoptera but differing conspicuously in several charac- ters: each occipital lobe is produced as an oblique flap (fig. 105C), with crenated inner margin (posterior margin of head rounded in Aspidoptera) ; the longitudinal and vertical membranous thoracic clefts are closed, the line of fusion marked by dark pigmented sutures ; the median mesonotal suture is bifurcate anteriorly ; the dorsal abdominal connexivum is bare except toward sides ; and sternum V is present in the male. DIAGNOSIS: Minute species, 1.2-1.8 mm. long. Head. — Flattened. Theca cordiform. Palpi roughly oval, slightly longer than wide, anterior margins obliquely truncated, with setae along margins, and on ventral surface with fine setae on slightly more than basal half, the upper surface microsetose. Laterovertices and occipital plates dif- ferentiated, both microsetose, setae on laterovertices restricted to antero-lateral por- tions, laterovertices each with a long, posteriorly directed process, oblique occipital plates with a feebly sclerotized, posteriorly directed flap on each side, their posterior margins pigmented, crenated, with long bristles, and overlapping the anterior portion of the prescutum. Mediovertex unsclerotized, projecting downward as a minute process between the occipital flaps. Genae with long setae along dorsal margin, but ventral to these are numerous, shorter, thorn-like setae. Postgenae, below, with a large oval, lightly sclerotized transverse area on each side of the oral cavity. Eyes minute, in- distinctly faceted, with four to six facets. Thorax. — Mesonotum flat; median suture of prescutum bifurcate anteriorly, united posteriorly with the sinuate transverse suture; scutum very short, less than a third as long as prescutum. Mesipisterna prominent and flattened above, longitudinal and vertical membranous clefts closed, represented by rigid, pigmented sutures. Sterno- pleura broadly projecting and emarginate between the procoxae. Posterior margin of pleurotrochantines with a median, rounded, slightly reflexed lobe. Wings. — Brachyp- terous, with venation essentially normal, except that second and third crossveins are absent. Legs. — Short, subequal, femora stout, femora and tibiae subequal in length, clothed with long setae on antero-dorsal surface of fore- and midlegs, and on dorsal surface of hindlegs, the setae elsewhere shorter to very short. Tarsi short, tarsomeres 1-4 progressively wider and more strongly anteroposteriorly compressed, the fourth nearly as wide as the fifth, the latter broad, posteriorly rounded ; claws large. WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 559 B Fig. 105. Exastinion clovisi (Pessoa and Guimara.es). A, female, with pupa extruding, dorsal view. B, left male gonapophysis, specimen from Anoura cultrata (no. 8593), Rio Changena Camp (Bocas del Toro). C, head, dorsal view (setae of laterovertices and occipital lobes omitted). D, wing. E, mouthparts. A, D, and E from Jobling (1949a), based on specimen from Anoura g. geoffroyi, Aripa Cave, TRINIDAD. 560 ECTOPARASITES OF PANAMA Abdomen. — Dorsal connexivum bare except toward sides; paired dorsal segmental setae absent; sides and venter with fine, short setae. Tergum I+II very short at middle, with conspicuous, posteriorly directed lateral lobes. Sternum II well developed, setose in a median triangular area and along posterior margin, the setae borne in minute thecae, these, in turn, on a translucent plaque. Female: Tergum VII a small, feebly sclerotized, translucent, transverse plate, bearing two short setae. Supra-anal plate very short, transverse. Seventh sternites short and transverse. Cerci free, not united with the ventral arc. Male: Sternum V very short and broad; VI not distinguishable as a sclerite, though it seems to be represented by a thread-like, translucent band. Hypopygium very short. Gonapophyses as in Trichobius. The name of the genus is derived from the Greek exastis (rough edge or fringe) 4- inion (back of head, occiput) . Exastinion clovisi (Pessoa and Guimaraes), new combination. Figure 105. Aspidoptera clovisi Pessoa and Guimaraes, 1936, Ann. Fac. Med. Sao Paulo, 12, fasc. 2, pp. 262-263, figs. 5-6 — Ipiranga, Sao Paulo, Brazil, from Anoura geoffroyi ( ?Laboratorio de Parasitologia, Faculdade de Medicina, Universidad de Sao Paulo); 1940, Arq. Inst. Biol., Sao Paulo, 11: 424, fig. 1. Guimaraes, 1944, Papeis Avulsos, Sao Paulo, 6: 187, figs. 10-14 (descr. of "larva"). Jobling, 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 138 (redescr.), 136 (keyed), fig. 2; 1949, Parasitology, 39: 316 ff. (hosts) , 327 (records) , fig. U ; 1951, Trans. Roy. Ent. Soc., Lond., 102: 216. Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 183, 188 (records). Aspidoptera phyllostomatis (not Perty, 1833), Bequaert, 1940, Rev. Acad. Colomb. Cienc. Ex., Fis. y Nat., 3: 418. Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 26 (part.). PANAMANIAN MATERIAL EXAMINED: 53 specimens (9 lots) from 11 bats at localities from 2500-5300 feet. From Anoura geoffroyi lasiopyga: 4 (2 bats) , Rancho Mojica (Bocas del Toro) , 12 September 1961 ; 7 (3 bats) , Casa Lewis, Cerro Punta (Chiriqui), 1 to 5 February 1960, and 1, same locality, 4 May 1960; 11 (2 bats), Cerro Hoya (Los Santos), 11 February 1962. From Anoura cultrata: 3 (1 bat), Cerro Punta (Chiriqui), 12 March 1962; 18 (2 bats), Rio Changena Camp (Bocas del Toro), 27 September 1961. OTHER MATERIAL EXAMINED : We have examined about 175 other speci- mens from Anoura aculeata (VENEZUELA) and from subspecies of Anoura geoffroyi from TRINIDAD, VENEZUELA, COLOMBIA, ECUADOR and GUATEMALA. REMARKS: Jobling (1949a, p. 138, fig. 2B) indicated that the eyes of A. clovisi were not faceted and so figured one. All of our specimens show five to six minute facets in slide preparations (fig. 105C). The palpus fig- ured by Jobling (fig. 105E) is lying in an oblique plane and appears to be transverse. Actually, the palpi are very slightly longer than broad. The setae on sternum II are quite uniform in size in the male, though one or two at each postero-lateral angle may be a little longer. In the female there is a group of conspicuously longer setae within these angles. This may be a composite species, Jbut to date we have not been able to distinguish between specimens from different species of Anoura or from different geographic areas. Noctiliostrebla Wenzel, new genus Type-species: Lipoptena dubia Rudow, 1871. Superficially resembling species of Aspidoptera, Exastinion, Mastoptera, and Paradyschiria but not closely related to any of these. Easily separated WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 561 from them by the characters given in the key and distinctive in the follow- ing : longitudinal and vertical membranous clefts closed, the mesepisternum and prescutum, and the mesepisternum and mesepimeron completely fused without evidence of a suture ( ! ) ; tergum VII of female represented by paired transverse sclerites; male gonapophyses "free" anteriorly, articulated ventrally by short processes to forks of the gonapophyseal apodeme ; sterna V and VI absent in male ; hind tibiae along inner apices with a mixture of normal setae and extremely fine, translucent, flattened setae, the latter so closely placed as to appear to be branched when examined under low mag- nifications. DIAGNOSIS: Small louse-like species. Head. — Nearly round, viewed from above; ele- vated posteriorly, viewed in profile; with well-defined laterovertices and occipital lobes, each laterovertex divided by an oblique, dark suture. Palpi irregularly rounded, with marginal setae only, ventral surface lacking setae. Theca subcordif orm. Thorax. — Anterior margin of prescutum feebly emarginate but slightly projecting at middle; median suture strong, complete, united with the transverse suture, to form an inverted "Y". Spiracular openings large, doughnut-shaped, very conspicuous. Prescutum without discal setae, the setae restricted to sides and anterior margin. Longitudinal and vertical membranous clefts absent, the prescutum and pleura com- pletely fused, without evidence of a suture. Dorsal and ventral surfaces of thorax often with conspicuous irregular lighter areas in the cuticle. Wings. — Very short, oval, pointed, with marginal bristles, veins indistinct, very much reduced in number, usually three, sometimes four. Legs. — Short, subequal. Tibial setae sparse, arranged in rows; dense setae along inner margin of metatibiae on about distal fifth, a mixture of normal setae, and extremely fine flattened setae, which in profile appear almost filamentous under oil immersion. Tarsi stout, about three-fourths as long as tibiae; tarsomeres 1-4 antero-posteriorly compressed, the last tarsomere with subparallel sides, about as long as 1-4 combined; claws large, conspicuous. Abdomen. — Spiracles much smaller than thoracic spiracles but conspicuous and "doughnut-shaped." Abdominal connexivum transversely wrinkled, the setae relatively short and borne on feebly sclerotized plaques. Tergum I+II very well developed, emarginate at middle. Female : Tergum VII represented by a pair of transverse sclerites. Supra-anal plate short. Ventral arc about the same size as supra-anal plate, the two forming an annulus. Cerci free, not united with ventral arc. Male: Hypopygium greatly developed, conspicuous, obconical. Sterna V and VI absent. Gonapophyses short and blade-like, free and separate, not arising from a gonapophyseal sheath but each articulated ventrally by a process to the forked gonapophyseal apodeme and posteriorly to the recurved base of the aedeagus; aedeagus strongly laterally com- pressed, blade-like in lateral profile, its posterior end curved, but not coiled. The species of Noctiliostrebla and Paradyschiria do not appear to be at all closely related, but the presence in both genera of the peculiar, heavy, transversely wrinkled abdominal connexivum (with setae borne on con- spicuous plaques) and the conspicuous spiracles, suggest that these charac- ters are of adaptive value to parasites of the fish-eating bats. One might speculate that they are concerned with water regulation. The bats not only scull the water surface for food, but it is known that, at least in the case of Noctilio leporinus, the bats can swim when necessary. Protective adapta- tions that would insure survival of these parasites would be of considerable importance, especially to the parasites of leporinus, a bat which frequently fishes in salt water. Another peculiarity of the parasites of these bats lies in the chaetotaxy. The bristles are strong, sparse, and semi-erect in all of the streblidae of Noctilio, including a species of an undescribed genus, which 562 ECTOPARASITES OF PANAMA superficially resembles a large Trichobius. The fur of Noctilionidae has un- usually short hairs (Benedict, 1957) . Perhaps the strong semi-erect bristles give the flies additional purchase in the short pelage. When we first studied our material of this genus, it appeared that there was only one species, megastigma (Speiser) , that it occurred on both species of Noctilio, and that N. dubia (Rudow) was a synonym. As the picture of host-parasite specificity in Panamanian Streblidae developed, it no longer seemed reasonable that two bats that are as isolated ecologically as Noctilio labialis and N. leporinus should have the same species of para- site. We restudied both our Panamanian and other material and found that there were two species in Panama, one on each species of Noctilio. Our non- Panamanian material, too, seemed to segregate into these two forms. How- ever, we were puzzled by the fact that the species which occurred on N. labialis in Panama seemed to replace the other species on N. leporinus in Venezuela, Trinidad, and Amazonian South America. We could understand that it might be possible for the parasite of N. labialis to occur on N. leporinus in areas to which the parasite of leporinus had not been able to disperse. However, if the parasite of labialis could live on both labialis and leporinus in some areas, it did not seem reasonable that it should be re- stricted to labialis in Central America, Cuba, and Peru, where the parasite of leporinus also occurred. That is, it did not seem reasonable unless a factor such as competitive exclusion were operating. This we were not willing to accept until we had eliminated another alternative, namely that we were dealing with a complex of closely related species rather than just two. Study of nearly all available additional material confirmed the latter suspicion. We now know that there are about seven to nine species of Noctiliostrebla, five or six of them on N. leporinus and two or three on N. labialis. They may be separated into two groups. In the species of Group A, the aedeagus lacks a recurved dorsal subapical spine, while in Group B, the aedeagus possesses such a spine. Species of Group A apparently occur only on N. leporinus; one species (N. aitkeni n.sp.) of Group B occurs on leporinus, the others on N. labialis. It was the apparently anomalous occurrence of aitkeni on N. leporinus leporinus which caused us to re-examine our material. We are not yet in a position to revise the species in detail, but restrict our treat- ment to such species as necessary to enable us to apply names to the Panama- nian species. There are few records of flies of this genus occurring on bats other than Noctilionidae. Some of these undoubtedly represent contaminations and errors of association ; however, some probably represent temporary trans- fers to other hosts, such as Molossus, with which the Noctilionidae sometimes roost. The distributions of the species that occur on Noctilio leporinus seem to roughly correspond to those of subspecies of the host. This raises a num- ber of interesting problems. Mr. Philip Hershkovitz tells us that these are "nominal" subspecies that do not bear close scrutiny. If these are inter- breeding, intergrading populations of bats, it is quite possible that some of the species of Noctiliostrebla, too, will be shown to intergrade, as additional material becomes available. On the other hand, if such is not the case, one WENZEL, TIPTON, AND KIEWLIC2 : STREBLID BATPLIES 563 may raise the question as to whether or not the "subspecies" of Noctilio leporinus may actually be distinct species. While we recognize that speciation of host and parasite need not proceed at the same rate, nonethe- less, the data suggest that a careful restudy of the Noctilionidae and their parasites, based on larger and more extensive geographic samples than are presently available, is in order. The two species groups of Noctiliostrebla may be characterized as fol- lows: Female: Supra-anal plate with one or two long strong bristles on each side at mid- length (fig. 107D) ; sternum II very large, long, emarginate at middle of posterior margin (fig. 106B). Male: Aedeagus lacking a dorsal subapical spine (fig. 107B). Posterior margin of sternum II outwardly rounded or angulately projecting, the marginal setae at middle generally forming a false ctenidium (fig. 106A) Group A Female: Supra-anal plate with one or two short bristles on each side at mid-length (fig. 107C). Sternum II shorter, strongly transverse, not emarginate at middle of posterior margin (fig. 109B). Male: Aedeagus with a dorsal subapical spine (fig. 107A). Posterior margin of sternum II straight or feebly arcuate, the bristles along middle of posterior margin not forming a false ctenidium (fig. 109 A) Group B Group A In addition to the characters given above, it may be noted that all the species of this group, excepting one, lack setae near middle of the median wing vein ; the species of Group B typically have one or two, but an unde- scribed species from Brazil has from two to five. Noctiliostrebla dubia (Rudow), new combination. Lipoptena dubia Rudow, 1871, Zeitschr. gesammte Naturwiss., 37, (n. s., 3), p. 122; 1872, Ann. Mag. Nat. Hist., (4), 9: 407 — Venezuela, from Noctilio dorsatus [-N. leporinus] — (Zoologische Museum, Hamburg). Speiser, 1902, Syst. Hymen. Dipt., 2: 159-160 (synonymizes Paradyschiria fusca Speiser 1900 with dubia, in error). Aspidoptera megastigma (not Speiser, 1900) Bequaert, 1942, Bol. Ent. Venez., 1, (4), p. 88. Speiser (1902) synonymized his Paradyschiria fusca (Speiser, 1900) under dubia Rudow, 1871. He did so on the assumption that Rudow based his description of dubia on a mixed series containing three specimens of what Speiser regarded as his own Lepopteryx megastigma and a single specimen of his Paradyschiria fusca. This may have been true. Accord- ing to Speiser (op. cit.) Rudow's series contained two specimens of megastigma in a Canada balsam mount and one each of megastigma and Paradyschiria fusca in alcohol. However, Speiser further attempted to demonstrate that Rudow based his description of dubia primarily on the specimen of Paradyschiria. While it is apparently true that Rudow did have a mixed series, he cer- tainly did not base his description on the single specimen of Paradyschiria, as Speiser maintained. This is clear from both the German (1871) and English (1872) versions of his paper. For example, in referring to the median and transverse mesonotal sutures, Rudow says "mit einigen bogenformigen, quer und winkligen Langsfurchen von rother Farbe verse- 564 ECTOPARASITES OF PANAMA hen." This obviously refers to the condition in Noctiliostrebla, since these sutures are largely membranous in Paradyschiria. In 1962, Dr. T. C. Maa examined the Rudow types (slides only) of Lipoptena dubia and informed the senior author (pers. comm.) that they were the same as Lepopteryx megastigma Speiser. Because our studies of Noctiliostrebla indicated that a number of species had been confused under the name megastigma, it was important that the types both of Rudow's dubia and Speiser's megastigma be re-examined. Prof. Herbert Weidner of the Hamburg Museum kindly sent the slide of the male and female Lipoptena dubia Rudow with permission to remount them. These were not in Canada balsam as stated by Speiser (1902) but mounted dry and pressed flat between slide and coverslip. The specimens were remounted after long immersion in a solution of trisodium phosphate followed by treatment with KOH and the usual dehydration, clearing and mounting in Canada balsam. The specimen had been preserved dry so long that it was not possible to restore the telescoped abdomen of the female to normal condition. The male proved to be unusually brittle, and in dissecting the male terminalia, the abdomen was damaged. However, it is now possible, we believe, to determine the specific status of this species, and it will be re- described in a later paper. It differs conspicuously from traubi n.sp. (q.v.) in the characters of sternum II of the male. In dubia, the bristles of the posterior margin of sternum II do not form a ctenidial-like structure ; they are somewhat thorn-like and shorter than those toward the sides ; the discal setae are sparse, about eight in number. The species probably does not occur in Panama. Unfortunately the exact type locality is unknown, but we believe that dubia occurs on an Amazonian race of Noctilio leporinus. Noctiliostrebla megastigma (Speiser) , new combination. Lepopteryx megastigma Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 54-55, pi. 3, fig. 2 — Type locality unknown, off Noctilio dorsatus [=N. leporinus} — (Zoolo- gisches Museum der Humboldt Universitat, Berlin). Aspidoptera megastigma Speiser, 1900, Zool. Anz., 23: 154; 1902, Zeitschr. Hymen. Dipt., 2: 159. We have not examined the types of this species. Speiser had five dry specimens at the time he described it. According to Dr. T. C. Maa (pers. comm.) two are males and three are females. The country of origin is un- known. The host was given as "(N)octilio (d)orsatus" (=N. leporinus). Early in our studies, we sent copies of figs. 106 and 109 to Dr. H. Schu- mann of the Zoological Museum at Humboldt University, Berlin, with the request that he compare these with the type of megastigma. His reply indicated that sternum II of the female was of the type found in traubi n.sp. and dubia Rudow, i.e., long and with the posterior margin emarginate at middle. On the other hand the male apparently has a simple type sternum II, i.e., with posterior margin relatively straight and without a false ctenidium. We have seen two species with this combination of male and female characters of sternum II. However, until we are able to examine and dissect a male of the type series, it is not possible for us to settle the identity WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 565 of megastigma. It is clear that it is distinct from traubi, but it remains to be seen whether it is the same as dubia or is another species known to us from Amazonian Brazil. Noctiliostrebla traubi Wenzel, new species. Figures 106 ; 107B, D. Closely related to N. dubia (Rudow), 1871, and differing from it chiefly in that the posterior margin of sternum II is strongly produced at middle in the male, the marginal setae here more closely placed than they are laterally, subspiniform, distinctly longer than the discal setae, and forming a false ... , i > * , t ' , V-C\ \ Vi J% t >.., v vrS , \> •< r Fig. 106. Noctiliostrebla traubi, new species, sternum II. A, male holotype and B, female allotype from Noctilio leporimis mexlcanus (no. 5113), Fort Sherman. ctenidium (fig. 106A). In dubia, the posterior margin of sternum II is ap- parently straight at middle in the male and the median marginal setae are scarcely if any longer than the discal setae and do not form a false ctenidium. In the female of both species the first sternum is emarginate at middle, un- like that of N. maai n.sp. and N. aitkeni n.sp. The head and thorax are nearly identical throughout the genus and are not described below. DESCRIPTION : Wings. — Tips subacuminate ; apices with a coarse and a fine distal seta ; outer margin with about four stronger submarginal and seven or eight marginal setae, inner margin with three or four marginals and four or five submarginals; middle vein without bristles near mid-length ; vein along posterior margin usually divided into two veins which anastomose anteriorly and posteriorly. Legs. — Similar to those of maai and aitkeni. Abdomen. — Female: Lateral lobes of tergum I+II each with about 17 coarse setae, four to six of these (anteriorly and laterally) relatively short, being less than or only slightly more than half as long as the longest setae; inner margins on each side of mid-line, with four to eight additional small setae in a double row. Dorsal and lateral connexivum on each side posterior to tergum I+II with a cluster of about 10 setae that are coarser than the others but less than half as long as longest macrosetae of tergum I+II, and no more than twice as long as other dorsal connexival setae ; a transverse row of setae between sixth spiracles slightly longer than other connexival setae near them. Tergum VII a thin transverse strip with four minute setae. Supra-anal plate as on other members of group A (fig. 107D), with four apical macrosetae and a shorter but strong seta on each lateral margin. Setae of lateral connexivum minute; setae of ventral con- nexivum of about the same size as dorsals, except for a few long setae of four segmental 566 ECTOPARASITES OF PANAMA series. Sternum II (fig. 106B) very long, emarginate at middle with very few discal setae, marginal setae longer toward sides, but short along lateral margins. Seventh sternites very small, irregularly oval, usually with three or four shorter but conspicuous setae along apical margin, three long coarse setae just anterior to them and two to four very short (often thorn-like) setae anteriorly. Male: Tergum I+II generally similar to that of female, but macrosetae not as long ; short setae along inner margin relatively Fig. 107. Terminalia of species of Noctiliostrebla. A and B, male genital apparatus, lateral view: A, Noctiliostrebla maai, new species, paratype from Noctilio I. labialis (no. 6376), Rio Tuira (Darien) ; B, N. traubi, holotype. C and D, terminal cone, female abdomen: C, Noctiliostrebla aitkeni, new species, female paratype from Noctilio I. leporinus, Manzanilla, TRINIDAD ; D, N. traubi, new species, female from Noctilio leporinus, (CNHM 81166), Huasimo (Tumbes), PERU. longer than in the female. Sternum II (fig. 106 A) posteriorly strongly produced or rounded at middle, with few discal setae, the median marginal setae longer than the median discal setae and more closely placed Jthan the marginals towards the sides so as to form a false ctenidium. Gonapophyses (fig. 107B) shorter, more strongly curved than in maai and aitkeni, the aedeagus strongly laterally compressed as in those species but without a dorsal recurved subapical spine. Measurements : Male Female BL 1.59-1.65 2.06-2.28 TL 0.49-0.55 0.60-0.63 TYPE MATERIAL (from Noctilio leporinus mexicanus) : Holotype male and allotype female (slides) from host no. 5113, cativo trees, Fort Sherman WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 567 (Canal Zone), 2 December 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — 1, same data as the holotype ; 4, same data but 4 December ; 14 (2 lots), Rio Tuira (Darien), 5 March 1958, P. Galindo [GML] ; 4, from preserved museum specimens collected at Rio Santo, near Pacora (Panama) , 9 February 1935, by H. C. Clark [CNHM]. Non-Panamanian paratypes. — 7, Fortuna, 2 miles west of Pandora, Estrella Valley (Limon), COSTA RICA, H. W. Setzer [CNHM]. Paratypes to be deposited in the collections of Chi- cago Natural History Museum; the United States National Museum; the Gorgas Memorial Laboratory, Panama; Environmental Health Branch at Corozal (Canal Zone) ; the Departamento de Zoologia, Secretaria da Agri- cultura, Sao Paulo, Brazil ; and the Bernice P. Bishop Museum, Honolulu. OTHER MATERIAL EXAMINED. — From Noctilio leporinus : 3, Huasimo (Tumbes) , PERU, 750 meters elevation, 25 June 1954, C. Kalinowski, CNHM Peru Zoological Expedition (1953-54) ; 3, Rio Guimaral, Valledupar (Mag- delena), COLOMBIA, 18 September, no. 682, lot no. 69 [USNM]. We have also seen specimens taken in CUBA from Noctilio leporinus mastivus at Los Amacegos, 1 mile west of Santa Fe, Isle of Pines, and from Ariguanabo (Havana), H. Jaume [MCZ]. These differ slightly from our Panamanian specimens and could represent a separate entity. REMARKS: This species is named for Colonel Robert Traub, USA- MSC (Retired) , now at the School of Medicine, University of Maryland, not only in recognition of his outstanding contributions to the systematics of fleas, our knowledge of ectoparasites generally, and many other aspects of medical entomology, but also in appreciation of his invaluable assistance and cooperation through the years. Group B Noctiliostrebla aitkeni Wenzel, new species. Figures 107C, 108. Aspidoptera megastigma (not Speiser, 1900), Jobling, 1949, Proc. Roy. Ent. Soc. Lond., (B), 18: 140-142, fig. 3 (part.: description, figure, and Trinidad record) ; Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122: 220. This species is closely related to N. maai n.sp. and to an undescribed species from Bolivia. It is the only species of Group B which occurs on Noctilio leporinus (subsp. leporinus). The males are very similar to those of N. maai excepting as follows : there is a single rather than a double row of setae along inner margin of tergum I +11, the gonapophyses are slightly more slender and are subacuminate at apex. The females are easily dis- tinguished from maai by the very conspicuous cluster of long coarse connexival setae on each side posterior to tergum I+II and by the strong segmental groups of setae usually present posteriorly, medial to the spiracles on each side. Dr. Harold Oldroyd of the British Museum (Natural History) kindly sent us the female (slide) upon which Jobling based his illustration, from Pointe Gourde Caves, Trinidad. The description and illustrations of A. megastigma Jobling (loc. cit.) , apply to this species as does his record from Trinidad. In his figure, the coarse setae of the dorsal abdominal connexivum 568 ECTOPARASITES OF PANAMA Fig. 108. Noctiliostrebla aitkeni, new species, female. A, dorsal view. B, labium. C, wing. From Jobling (1949a, as Aspidoptera megastigma). Specimen from Noctilio I. leporinus. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 569 do not appear as long as they actually are. To Jobling's description, the fol- lowing may be added : Abdomen. — Posterior margin of sternum II nearly straight in both sexes, and nearly identical to that illustrated for maai n.sp. (fig. 109). Female: Lateral lobes of tergum I+II with about 18 coarse setae; dorsal inner margin with three to five short setae, usually in a single row. Connexivum on each side, posterior to tergum I+II, with a cluster of 12 very long setae, most of them as long as the shorter macrosetae of the lateral lobes of tergum I+II, some of them more than half as long as the longest macro- setae; medial to each of spiracles IV and V on each side of mid-line, are one or more coarser setae. Seventh tergites each with six to eight macrosetae, four of them in a transverse row across middle. Male: Inner margins of lateral lobes of tergum I+II with three or four, rarely five setae in a single row on each side. Genital apparatus very similar to that of maai n.sp. (fig. 107A). Measurements : BL TL Male 1.65-1.76 0.55-0.58 Female 1.95-2.20 0.60-0.66 TYPE MATERIAL: Holotype male and allotype female (slides) from Noc- tilio I leporinus, Manzanilla, TRINIDAD, 13 March 1957, T. H. G. Aitken. In the collection of Chicago Natural History Museum. Paratypes. — 55, same data as the holotype [TVL] ; 3 Scotland Bay, TRINI- DAD, September 1954, C. C. Sanborn, CNHM Trinidad Zoological Field Trip (1954) ; 2, Monos Island, B. W. I. [FCUCV] ; 16, Kaiserberg Airstrip, east of Zuid River, SURINAM, 12 October 1960, H. A. Beatty, CNHM Guianan Zoological Expedition (1960-62), 2, same expedition, Saramacca River, Loksie Hattie (Brokopondo), SURINAM, 5 January 1962, P. Hershkovitz; 8, Quebrada Esperanza, Rio Yauari-Mirim, Maynas (Loreto), PERU, 23 Sep- tember 1957, C. Kalinowski, CNHM Peru Zoological Expedition (1956-57) . Paratypes in the collections of Chicago Natural History Museum ; the United States National Museum ; the Bernice P. Bishop Museum, Honolulu ; Facul- dad de Ciencias, Universidad Central de Venezuela, Caracas ; Departamento de Zoologia, Secretaria da Agricultura Sao Paulo, Brazil ; and the Trinidad Regional Virus Laboratory, Port-of-Spain. REMARKS : This species is named for Dr. Thomas G. Aitken, of the Trini- dad Virus Laboratory (Rockefeller Foundation) , in recognition of his valu- able contributions to medical entomology. Noctiliostrebla maai Wenzel, new species. Figures 107A, 109. Closely related to N. aitkeni n.sp., but differing as indicated under that species. DESCRIPTION: Abdomen. — Female: Lateral lobes of tergum I+II with 10-14 coarse setae, and on dorsal inner margin with seven to nine short setae in a double row. Con- nexivum on each side posterior to targum I+II with a cluster of 14—16 setae that are coarser than other dorsal and lateral connexival setae, but none more than twice as long as other connexival setae and less than half as long as macrosetae of tergum I+II; a longer seta sometimes present medial to spiracle V; between sixth spiracles is a row of four long setae ; seventh tergites with four or five setae, four of them macrosetae. Supra-anal plate as in N. aitkeni n.sp. (fig. 107C). Male: Inner margin of each lateral lobe of tergum I+II with six to nine, rarely 10 setae in a double row. Measurements : BL TL Male 1.54-1.63 0.47-0.51 Female 1.87-1.98 0.49-0.55 570 ECTOPARASITES OF PANAMA TYPE MATERIAL: Holotype male and allotype female (slides) from Noctilio labialis labialis (host no. 6376), Rio Tuira (Darien), 3 March 1958, Pedro Galindo. In the collection of Chicago Natural History Museum. Paratypes. — From Noctilio labialis labialis: 19, same data as holotype; 2, Rio Chucanaque (Darien), L. H. Dunn [MCZ] ; 10, Juan Mina (Canal Zone) , 16 August 1945, H. Trapido [CNHM] ; 7, Chilibrillo Caves (Panama) , L. H. Dunn [MCZ]. From Noctilio labialis: 7, Playo del Medio (Bolivar), VENEZUELA, 19 April 1961, T. Ojasti [FCUCV] . Paratypes in the collection of Chicago Natural History Museum ; the United States National Museum ; the Museum of Comparative Zoology at Harvard; the Gorgas Memorial Laboratory ; the Environmental Health Branch, USAFSC at Corozal (Canal Fig. 109. Noctiliostrebla maai, new species, sternum II. A, male holotype. B, female allotype. Zone) ; Faculdad de Ciencias, Universidad Central de Venezuela ; Departa- mento da Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil; and the Bernice P. Bishop Museum at Honolulu. REMARKS: This species is named for Dr. Tsing C. Maa, in appreciation of the valuable assistance given by him, and in recognition of his important contributions to our knowledge of the batflies. KEY TO PANAMANIAN SPECIES OF NOCTILIOSTREBLA Median wing vein usually with one or two (rarely none) setae near middle. Posterior margin of sternum II (fig. 109) straight or nearly so, in both sexes, the marginal setae rather uniformly distributed. Female: Supra-anal plate much as in aitkeni (fig. 107C) with a pair of short setae on each side, one inserted at middle of lateral margin, the other on dorsal surface of 'plate at about lateral fourth. Male: Gonapophyses (fig. 107 A) similar to those of aitkeni; aedeagus with a dorsal subapical spine. Host: Noctilio labialis labialis maai n.sp. Median wing vein without setae near middle. Posterior margin of sternum II broadly emarginate (fig. 106B) in female; outwardly arcuate or angulate (fig. 106A) in the male, the median marginal setae subspiniform and more closely set, so as to form a false ctenidium. Female: Supra-anal plate (fig. 107D) typically with a strong seta on each side along lateral margin and a very short one ventral to it. Male: Gonapophyses as in fig. 107B; aedeagus without a dorsal subapical spine Host: Noctilio leporinus mexicanus traubi n.sp. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 571 Genus Paradyschiria Speiser Paradyschiria Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 38, 55-56 (diagn.), 62, 65 (keyed). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 25 (keyed), 29 (cit.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 26-27 (char., key to spp.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 357 ff. (morph., syst. pos.) . Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418 (keyed) ; 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Parasitology, 39: 321 (keyed) ; 1951, Trans. Roy. Ent. Soc. Lond., 102: 216 (morph. abdomen). Grandi, 1952, Introd. Studio Ent., 2: 478. Type-species: Paradyschiria fusca Speiser, 1900. The species of this genus are the only apterous Streblidae known and superficially resemble small Nycteribiidae. This resemblance is enhanced by the extensive membranous areas of the mesonotum with accompanying reduction of the sclerites. The females of P. fusca, Speiser and lineata Kessel are unique in possessing a vertical sclerite in the lateral connexivum between the posterior margins of the lateral lobes of tergum I +11 and spiracle III. This is feebly sclerotized excepting a narrow strip along an- terior margin which is more strongly sclerotized and easily visible in slide preparations as well as on specimens in alcohol. The female cerci are free and not united with the ventral arc. We tentatively recognize four species of this previously monotypic genus. Two of them, P. lineata Kessel and P. parvuloides n.sp., occur in Panama. PROVISIONAL KEY TO SPECIES OF PARADYSCHIRIA 1. Mesonotum (fig. 114A) with a short seta (rarely absent) anterior and lateral to the long macroseta of postero-lateral angle. Female: Lateral abdominal con- nexivum without a vertical sclerite between tergum I+II and spiracle III. Terminal cone (fig. 112D) with the usual four apical macrosetae, and with a short seta along each lateral margin and a pair of short discal setae at mid- length. Male: Gonapophyses slightly but distinctly hooked at apex 2 Mesonotum without a short seta anterior and lateral to the long macroseta of postero-lateral angle. Female : Lateral abdominal connexivum with an elongate- oval, vertical sclerite between posterior margin of lateral lobe of tergum I+II and spiracle III. Terminal cone (fig. 112B) with or without short discal setae, and in addition a pair of macrosetae, one on each side near posterior angle. Male: Right gonapophysis (fig. 113A) straight at apex, the left very feebly hooked at apex 3 2. Female: Seventh sternites (fig. 112C) with from two to four spine-like setae on apical margin. Male: Gonapophyses rather abruptly tapered on apical half parvuloides n.sp. Female: Seventh sternites with normal setae along apical margin. Male: Gona- pophyses rather evenly tapered to apex parvula Falcoz 3. Posterior margin of tergum I+II usually with one (rarely two) macrosetae that are conspicuously longer than the others (figs. 111A). Female: Mid-dorsal setae of abdominal connexivum conspicuously shorter than those lateral to them, those near base and apex shortest, the apical ones less than half as long as the setae lateral to them (figs. 110A, B). Terminal cone (fig. 112B) usually with baso-lateral macrosetae more widely separated than the outer setae of apical margin lineata Kessel Posterior margin of teigum I+II usually with two or three long subequal macro- setae that are conspicuously longer than the others. Female: Most of mid- 572 ECTOPARASITES OF PANAMA ij 13 ?- X^^* *VJ? r ' * Fig. 110. Female abdomen, dorsal view, of species of Paradyschiria. A and B, P. lineata Kessel; A, from Noctilio leporinus mastivus, Central Mercedes, Matas, CUBA; B, from AT. 1. mexicanus (no. 5558), Changuinola (Bocas del Toro). C, P. fusca Speiser, from N. 1. leporinus, Manzanilla, TRINIDAD. D, P. parvuloides, new species, from N. I. labialis (no. 6376), Rio Tuira (Darien). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 573 dorsal setae of abdominal connexivum (fig. HOC) shorter than those lateral to them, but not markedly so, many more than half as long as the setae lateral to them. Baso-lateral macrosetae of terminal cone usually no more widely sepa- rated than outer macrosetae of apical margin fusca Speiser Paradyschiria fusca Speiser. Figures HOC, 111B. Paradyschiria fusca Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 56-57 (descr.), 62 (cat.), 65 (keyed), pi. 3, fig. 1 — Orocue, Colombia, from Noctilio leporinus (Type repository unknown). [?] Cole, 1927, Proc. Calif. Acad. Sci., 16: 453 (morph. male genit.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658. Paradyschiria dubia (not Rudow, 1871), Speiser, 1902, Zeitschr. Syst. Hymenop. Dipt., 2: 159 (synonymizes P. fusca Speiser, in error). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 27. Stiles and Nolan, 1931, op. cit., p. 658 (part.) . Costa Lima, 1921 Arch. Esc. Sup. Agric. Med., Nictheroy, 5: 29 (part.). Bequaert., 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418. Guimaraes, 1941, Papeis Avulsos, Dept. Zool. Secret. Agric., Sao Paulo, 1: 217-222, figs. 1-4 (redescr.) ; 1944, loc. cit., 6: 186, figs. 5-9 (larva). Jobling, 1949, Parasitology, 39: 316 ff., 328, fig. 3 A (part.; synonymizes P. lineata Kessel, 1925 and P. parvula Falcoz, 1931). Good- win and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., 122: 220. Dr. T. C. Maa has informed us (pers. comm.) that he was unable to lo- cate the type of P. fusca and that it is probably lost. Thus, any interpreta- tion must be based upon Speiser's description and figure and the type host and type locality. Unfortunately, these do not permit us to determine which (if any) of the four species that we have segregated is fusca. His figure only permits one to determine that it is a Paradyschiria. No lateral macrosetae are shown on the supra-anal plate. This suggests that the species is either parvuloides n. sp. or one which we tentatively refer to P. parvula Falcoz. However, the figure does not show a small seta anterior and lateral to the posterior macroseta of the mesonotum. This is present in both parvuloides and parvula, neither of which occur on Noctilio leporinus. We doubt that there is a species with the combination of characters shown by Speiser and conclude that his illustration is too inaccurate to permit recog- nition of P. fusca. His description is of no help. The principal clues to its identity, then, are the type locality and type host. According to Bequaert (1940, p. 418), the type locality — Orocue, Colombia — is in the Department of Boyaca. He was unaware, apparently, that there is also a locality of that name in the Department of Vaupes. Both are in Amazonian Colombia, and in all likelihood the same species of Para- dyschiria occurs on Noctilio leporinus in both localities. P. lineata occurs in coastal Colombia, but judging from the distribution patterns of other streblids, including those on Noctilionidae, we doubt that it occurs in Ama- zonian Colombia. We have studied specimens — from Trinidad, Venezuela and Surinam — of a closely related population which we provisionally inter- pret as fusca. It appears to be the species redescribed and figured by Guimaraes (1941) as P. dubia, from Brazil. The specimens from Trinidad and Surinam are easily distinguishable from our Panamanian specimens of lineata, but those from Venezuela are difficult to separate. Perhaps P. fusca (sensu Guimaraes, 1941) and lineata Kessel represent clinal extremes. 574 ECTOPARASITES OF PANAMA Designation of a neotype of fusca must be deferred until specimens from near the type locality are available. Paradyschiria lineata Kessel. Figures 110A, B; 111A; 112A, B; 113A. Paradyschiria lineata Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 27-28 — Cuba (British Museum of Natural History). Jobling 1949, Parasitology, 39: 328 (as synonym of P. dubia Speiser, 1902, not Rudow 1871, in error). Jobling (loc. cit.) synonymized P. lineata Kessel under P. dubia Speiser 1902 (= P. fusca Speiser), not Rudow, 1871. As indicated above, the spe- cific identity of P. fusca Speiser is uncertain, though it seems that it is not Fig. 111. Abdominal tergum I+II, male. A, Paradyschiria lineata Kessel, from Noc- tilio leporinus mastivus (CNHM no. 93668), Los Amacegas, Isle of Pines, CUBA. B, P. fusca Kessel, from N. I. leporinus, Manzanilla, TRINIDAD. the same as lineata Kessel, described from Cuba. We cannot separate our Panamanian specimens, taken from Noctilio leporinus mexicanus, from Cuban specimens taken from N. I. mastivus. We therefore apply Kessel's name to the specimens from Panama. Measurements: BL TL Male 1.37-1.87 0.41-0.47 Female 1.76-1.98 0.47-0.49 PANAMANIAN MATERIAL EXAMINED. — From NoctlllO leporinus canus: 22 specimens (4 lots) as follows : 12 (1 bat) , Changuinola (Bocas del Toro), 29 February 1960; 10 (3 bats), Fort Sherman (Canal Zone), 2 and 4 December 1959 ; 1, removed from preserved museum specimen collected at Rio Santo, near Pacora (Panama) , H. C. Clark, 9 February 1935 [CNHM] . OTHER MATERIAL EXAMINED: CUBA. — From Noctilio leporinus mastivus: 1, removed from preserved museum specimen from Los Amacegos, 1 mile west of Santa Fe, Isle of Pines, February 1961, Gilberto Silva [CNHM] ; 1, Ariguanabo (Havana), H. Jaume [MCZ]. Paradyschiria parvula Falcoz Paradyschiria parvula Falcoz, 1931, Parasitology, 23: 267, figs. 4, 5 — Brazil, from Dirias albiventer [= Noctilio labialis] (Museum National d'Histoire Naturelle, WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 575 Paris). Jobling, 1949, Parasitology, 39: 328 (as syn. of P. dubia, in error). Paradyschiria dubia (not Rudow, 1871), Costa Lima, 1921 (part.), Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 23 (host record from N. labialis), 29 (cat.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 658 (part.). Guimaraes, 1941, Papeis Avulsos, Secret. Agric., Sao Paulo, 1: 221 (part., record from Noctilio albiventer). We have not examined the type of P. parvula, but we have a series col- lected by Dr. Merl Runs from Noctilio labialis at San Joaquin (Beni), BO- LIVIA, which we provisionally identify as this species. It is closely related to parvuloides n.sp. but is easily separated by the characters of the female seventh sternites (see key) . In addition to the characters given in the key, parvula and parvuloides may be separated from both fusca and lineata by their smaller size and the relatively uniform, subequal macrosetae of the posterior margin of tergum I +11. A small series received from Dr. Machado-Allison and collected in Vene- zuela at Playo del Medio (Bolivar) and Calabozo (Guarico), VENEZUELA, may be this species. However, they represent a puzzling mixture of char- acters of parvula, parvuloides, and fusca and more extensive series will be needed to determine their identity. They could represent a mixture of species, or an undescribed species, but it is also possible, as may be the case with some other streblids, that parvula and parvuloides represent either clinal extremes or subspecies, and that eastern Venezuela is an area of hybridization or an intermediate point on a cline. Paradyschiria parvuloides Wenzel, new species. Figures HOD; 112C, D; Paradyschiria dubia (not Rudow, 1871), Cooper, 1941, Yearb. Amer. Phil. Soc., 1941: 126 (chromosome no.). Jobling, 1949, Parasitology, 39: 316 (table 1), 328 (part., records from Dirias albiventer minor, Panama). Differing from P. lineata Kessel and fusca Speiser in having a short seta (fig. 114 A) lateral and slightly anterior to the posterior mesonotal macro- seta (absent in fusca and lineata) , and in lacking lateral macrosetae on the supra-anal plate of the female (fig. 112D). In these respects it resembles P. parvula Falcoz (of our interpretation) but differs from it and from both fusca and lineata in having from two to four short stout spine-like setae (fig. 112C) rather than normal setae on apical margins of seventh sternites of female. The male gonapophyses are rather abruptly tapered on apical half, rather than evenly tapered as in parvula. DESCRIPTION: Head. — As in P. lineata and P. fusca. Thorax. — As in those species, except that a short seta is present on prescutum on each side, lateral and slightly anterior to the postero-lateral macroseta (fig. 114A). Abdomen. — Sternum I+II setose in a median, roughly triangular area, the marginal setae longer toward sides. Female : Tergum I+II with (as usual) a pair of isolated setae near base; on each lobe there are well developed setae near mid-line on apical half and laterally on lateral half; usually one or two conspicuously longer and stouter submarginal setae near apical margin and a row of eight or nine setae along apical margin of each lobe, three or four of them much longer than the others. Tergum Vll+supra-anal plate (terminal cone) well sclerotized and pigmented, with four apical macrosetae, a pair of short discal setae at mid-length, and a minute seta at each posterior angle, lateral to the outer macroseta. Seventh sternites scarcely longer than broad, with eight to ten short setae basally and 576 ECTOPARASITES OF PANAMA two much longer, slender subapical setae, the outer seta nearly twice as long as the inner; with an apical row of three or four short, spine-like setae and lateral to them a stout seta which is nearly as long as the subapical macroseta. Dorsal and lateral connexivum clothed with moderately long setae, these slightly shorter along mid-line, those at extreme base and beyond middle as long as the setae lateral to them; setae of ventral connexivum very short, except for pairs of segmental setae; all setae borne on Fig. 112. A, seventh sternites and B, terminal cone, female abdomen of Paradyschiria lineata Kessel, from Noctilio leporinus mexicanus (no. 5558), Changuinola (Bocas del Toro). C and D, same structures, P. parvuloides, new species, paratype from Noctilio I. labialis (no. 6376), Rio Tuira (Darien). sclerotized plaques. Male: Tergum I+II with setae of posterior margin shorter than in female, three or four usually longer than others but not more than twice as long as discal setae. Connexivum covered with dense, relatively short setae, these shortest along the sides; ventral setae only a little shorter than dorsal setae. Sternum V di- vided into two short transverse oval sclerites bearing about six bristles each, four along apical margin, two near outer margin. Hypopygium covered with setae similar to those of connexivum, except for two or three longer slender setae apically near inner margin. Gonapophyses and aedeagus as shown in fig. 113B, abruptly tapered apically, in lateral view, and slightly hooked at apex. Measurements : BL TL Male 1.30-1.48 0.38-0.42 Female 1.37-1.66 0.38-0.41 TYPE MATERIAL (734 specimens, 26 lots, from Noctilio labialis minor) : Holotype male and allotype female (slides), host no. 6374, Rio Tuira WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 577 (Darien) , 2 March 1958, Pedro Galindo [GML] . In the collection of Chicago Natural History Museum. Paratypes. — 5 (1 bat), Fort Davis (Canal Zone), 26 October 1960; 3, Galeta Island (Canal Zone), 19 November 1959; 1, Gamboa (Canal Zone), 19 September 1960; 1, Juan Mina (Canal Zone), 20 April 1954, P. Galindo [GML] ; 10, same locality, 16 August 1945, H. Trapido ; 2, from preserved bats collected at Los Cascadas (Canal Zone) , 1933, by L. H. Dunn [CNHM] ; Fig. 113. Male genital apparatus, left lateral view. A, Paradyschiria lineata Kessel, same host data as for fig. 112 A. B, P. parvuloides, new species, paratype from Noctilio L labialis (no. 6374), Rio Tuira (Darien). 2, Barro Colorado Island (Canal Zone), 5 December 1956, C. B. Koford [USNM] ; 5 (2 lots), Summit (Canal Zone), 30 September 1932, L. H. Dunn [cu] ; 4 (slides), same data [BMNH — Jobling Collection] ; 121, same locality and collector, 5 August 1930 [AMNH] ; 3, same locality, K. W. Cooper [MCZ] ; 8, Salamanca, Rio Pequeni (Canal Zone) , 1 April, Griswold [MCZ] ; 1, Pacora (Panama), 6 June 1961; 1 from preserved bat collected at Panama (Pan- ama) [CNHM]; 202 (3 lots), Chilibrillo Caves (Panama), L. H. Dunn [MCZ] ; 15, same locality [USNM] ; 17, same locality, L. H. Dunn [cu] ; 2, Rio Mono Camp (Darien), 25 July 1963, Wilbur Lowe; 137, same locality, 3 March 1958, P. Galindo [GML] ; 5, Rio Chucanaque (Darien), L. H. Dunn [MCZ] ; 39, same data as holotype, and 133, same data but 2 March 1958 ; 4 (alcohol), "Darien," 4 February 1933, L. H. Dunn [BMNH — Jobling collec- tion, det. as P. lineata Kessel, by Jobling] ; 1, near Rio Guanico, Guanico (Los Santos), 8 February 1962. From Sturnira lilium parvidens: 1, same locality, 8 February 1962. 578 ECTOPARASITES OF PANAMA Non-Panamanian paratypes. — 75 (6 lots) from Noctilio labialis as fol- lows : 26, 6 miles west of Rama, Bluefields, NICARAGUA, elevation 50 feet, 29 April 1963, D. C. Carter; 26, 9 miles NE of Puerto Gulfito (Puntarenas) COSTA RICA, 29 April 1963, D. C. Carter; 18, Unguia, Golfo de Uraba (Choco) COLOMBIA, 8-11 March 1950, P. Hershkovitz, CNHM Colombian Zoological Expedition (1948-52) ; 7, Rio Docompado (Cauca), COLOMBIA, 350 meters elevation, 26 September 1958, Kjell von Sneidern, CNHM Co- lombian Zoological Expedition (1958-59) ;8 ( from type specimen of Noctilio Fig. 114. Paradyschiria parvuloides, new species, male. A, thorax, dorsal view (s = seta antero-lateral to posterior mesonotal macroseta). B, base of abdomen and thorax, showing posterior chaetotaxy of tergum I+II. minor Osgood), Encontrados (Zulia), VENEZUELA, 15 February 1908, Ned Dearborn, Field Museum Expedition (1908). Paratypes to be deposited in the collections listed on p. 410. REMARKS : This species appears to be restricted to Noctilio labialis and possibly to the subspecies minor Osgood. The single specimen from Sturnira lilium parvidens (no. 9351) is probably a contaminant from a specimen of N. I. minor (no. 9352) collected at the same time. Parastrebla Wenzel, new genus Type-species : Parastrebla handleyi Wenzel, new species. Similar to Pseudostrebla, Speiseria, and Trichobius, but differing as follows: from Pseudostrebla in that the laterovertices lack a pigmented suture, the postvertex is membranous and the head is deep and not flattened ; from Trichobius and Speiseria in having conspicuous tibial macrosetae; from Speiseria in not having greatly elongated hind legs ; and from all other known New World Streblidae by the presence in the female (male unknown) of a ventral, feebly sclerotized, subapical abdominal plate (fig. 116B) bear- ing blunt spines, anterior to the seventh sternites. DIAGNOSIS: Head. — Distinctly narrower than thorax; width greater than length from base of occipital lobes to apex of f rontoclypeus ; about as high as wide. Eyes WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 579 faceted, not projecting beyond postgenae. Laterovertices and occipital lobes well de- nned, the latter somewhat inflated; lateroverticss lacking a diagonal suture. Palpi oval, approximately as broad as long, with both ventral and marginal setae. Thorax. — Widest at level of transverse suture, narrowing apically; median suture present; humeral calli slightly produced but not prominent. Wings. — Venation as in Trichobius; anal vein with several strong setae near basal angle. Legs. — As in Pseudostrebla, the middle and hing legs progressively longer than the forelegs, the hindlegs not greatly elongated. All tibiae with conspicuously long macrosetae in addition to short setae. Abdomen. — As in Trichobius, but female with a very feebly sclerotized plate (sixth sternum?), anterior to seventh sternites, whose apical margin has a row of nine or ten blunt spines. Ventral arc well developed, translucent, with conspicuous flanges. Sterno- pleura and pleurotrochantines as in Speiseria, Pseudostrebla and Trichobius phyllos- tomae, the anterior margin projecting between coxae and angulately emarginate. REMARKS : The female genital structure of Parastrebla is very similar to that of the Trichobius major group, especially of T. truncatus Kessel. How- ever, we cannot determine with certainty, from the slide preparation, whether or not the cerci are fused or articulated with the ventral arc. The structure of the head and abdomen suggest that the genus is in some ways intermediate between Trichobius on the one hand, and Pseudostrebla on the other. Parastrebla handleyi Wenzel, new species. Figure 116. DESCRIPTION (Female) : Head. — Eyes small, transverse, with seven or eight in- distinct facets. Genae and postgenae with numerous, short, discal setae and con- spicuously longer marginal setae. Laterovertices subquadrate, declivous anteriorly, with about six conspicuous and one short setae. Occipital lobes each with three very long and about six other strong, shorter setae. Thorax. — Anterior margin biconcave for reception of occipital lobes of head, bi- tuberculate at middle; calli feebly produced. Median suture extending about two- thirds the distance to transverse suture ; disks of prescutum and scutum with numerous, rather uniformly distributed, recumbent, short setae, which become longer (though still short) anteriorly; longer, strong setae present laterally. Scutellum with four macro- setae. Underside rather uniformly, densely setose, the setae short, with longer, strong setae along sides and posterior margin, the latter fully emarginate at middle. Wings. — Costa with ten to twelve strong dorsal setae, these gradually shorter apically on basal half, the setae on apical half short; marginal setae very strong on about basal half and much more numerous than dorsal setae, becoming progressively shorter on apical half. Ri with one very long basal macroseta (as in Pseudostrebla) followed by about four shorter but strong setae, followed by short setae; Rs bare; third, fourth and fifth longitudinal veins each with three or four strong, basal setae, the other setae short. Legs. — Densely clothed with short setae. Profemora with about nine dorsal macro- setae arranged in two rows each side and two others laterally near apex. Mesofemora with about four macrosetae on apical half of dorsal surface, two ventro-laterally on each side near apex, and a row of about eight shorter erect setae on lateral surface on basal half. Metafemora with about 19 macrosetae on dorsal, lateral, and ventral surfaces. Protibiae with about four erect macrosetae along dorsal edge, mesotibiae with five, metatibiae with six. Tarsomeres 1-4 together a little longer than the last segment (excluding the claws), which is laterally slightly compressed; tarsomere 1 slightly longer than 2-4, its ventral surface with several pairs of short but rather strong setae. Abdomen. — Lateral lobes of tergum I+II with about 16 discal macrosetae and about a dozen short ones along the lateral margin. Dorsal connexivum bare except for four pairs of conspicuous segmentally arranged setae; lateral and ventral connexivum densely clothed with short setae of about same size; with two transverse rows, each of about four much longer setae anterior and lateral to sternum VI (?); tergum VII narrow, 580 ECTOPARASITES OF PANAMA parallel-sided, strap-like, continuous with the supra-anal plate and bearing four minute setae arranged in two pairs, one anterior to the other, evenly spaced. Supra-anal plate very short, with a short seta on each side and four apical macrosetae. Seventh sternites oval, with about 14 setae of which three on apical margin are macrosetae. Measurements: BL TL WL ww Holotype female 2.12 0.80 1.76 0.82 TYPE MATERIAL: A unique holotype female (slide) from Micronycteris nicefori (host no. 11565), Armila (San Bias), 13 March 1963, C. 0. Hand- A A Fig. 115. Synthesiostrebla amorphochili Townsend, female. A, dorsal view. B, ventral view, head and anterior part of thorax. From Jobling (1947). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 581 ley, Jr. and F. M. Greenwell. In the collection of Chicago Natural History Museum. REMARKS: This species is named after Dr. Charles 0. Handley, Jr., in recognition of his invaluable contributions to this volume. Fig. 116. Parastrebla handleyi, n. gen., n. sp., holotype female. A, thorax, dorsal view. B, apex of abdomen, ventral view. 582 ECTOPARASITES OF PANAMA Genus Pseudostrebla Costa Lima Pseudostrebla Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 23 (diagn.), 25 (keyed, as Psaudostrebla, sic!), 30 (cat.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 12 (keyed), 19. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 654. Curran, 1935, Amer. Mus. Novit., no. 765, p. 6 (keyed). Jobling, 1936, Parasitology, 28: 359, 367 ff., 370 (keyed); 1939, Arb. Morph. Tax. Ent., 6, (3), pp. 269-270 (diagn., keyed). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed) ; 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Parasitology, 39: 321 (keyed). Type-species: Pseudostrebla ribeiroi Costa Lima, 1921. Only one species of Pseudostrebla (ribeiroi Costa Lima) has been de- scribed. In our material from Panama we recognize two, one of them new. In the collections of Chicago Natural History Museum are two additional undescribed species. All are from bats of the genus Tonatia. It appears that each species of Tonatia is parasitized by a different species of Pseudo- strebla. The Panamanian species may be separated as follows : KEY TO PANAMANIAN SPECIES OF PSEUDOSTREBLA Head: Posterior margin of occiput on each side with a feebly developed, unisetose tubercle; oral cavity about as long as or slightly longer than broad. Legs: hind tibiae scarcely longer than the tarsi, with five macrosetae, of which the hind two are nearly approximate and on opposite sides of tibia, middle macroseta longest, slightly more than half as long as tibia. Host: Tonatia minuta greenwelli n.sp. Head: Posterior margin of occiput on each side with a well-developed bisetose tubercle; oral cavity strongly transverse, more than twice as wide as long. Legs: Hind tibiae about a third again as long as the tarsi, with six macrosetae, three on basal third, two very long ones just beyond middle, the sixth near apex. Host: Tonatia silvicola ribeiroi Costa Lima Pseudostrebla ribeiroi Costa Lima. Figures 117; 118, B, D. Pseudostrebla ribeiroi Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 23-25, pi. 2, fig. 4 — Brazil: Matto Grosso, from "Tonatia amblyotes" [sic!] (?Museu Nacional do Rio de Janeiro). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 19. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 656. Jobling, 1939, Arb. Morph. Tax. Ent., 6, (3), pp. 268 (notes), 271-273 (redescr.), fig. 1A-C; 1949, Parasitology, 39: 316 ff., fig. 2F. A single male specimen that we have identified as this species was col- lected from Tonatia silvicola (=amblyotis) at Armila (San Bias) , 12 March 1963, by C. 0. Handley, Jr. and F. M. Greenwell. We have seen 13 ad- ditional specimens (9 lots) from the same host, from COLOMBIA, PERU, and ECUADOR. Measurements are given below to supplement Jobling's ex- cellent figure and description. They are based on the single Panamanian specimen and a series of 11 Colombian specimens. Measurements: BL it, WL ww Male 2.23-2.30 0.85-0.88 1.63-1.65 0.77 Female 2.45-2.90 0.84-0.97 1.63-1.93 0.77-0.95 Pseudostrebla greenwelli Wenzel, new species. Figure 118A, C. Similar to P. ribeiroi Costa Lima, but differing from that species by the characters given in the key and those noted below in the description, as well as by its much smaller size and the different male gonapophyses. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 583 DESCRIPTION: Smaller, and less strongly pigmented and less strongly setose than ribeiroi. Head. — Transverse, but appearing less oblong than that of ribeiroi; latero- vertices with about 10 setae, including minute setae along anterior margin (12—13 in ribeiroi) ; occipital lobes with about nine coarse setae (about 12 in ribeiroi) in addition to the laterally directed setae along side margins, the posterior margin of each lobe with a feeble, unisetose tubercle (in ribeiroi a prominent bisetose tubercle) ; mediovertex with four setae, two each side of middle; palpi transverse but only about a third again as Fig. 117. Pseudostrebla ribeiroi Costa Lima. A, female, dorsal view. B, mouthparts. C, venter of thorax. From Jobling (1949b). 584 ECTOPARASITES OF PANAMA wide as long (nearly twice as wide as long in ribeiroi) ; oral cavity about as long as or slightly longer than broad (in ribeiroi strongly transverse, more than twice as wide as long, the theca retracted, in repose, into cavity above postgenae). Thorax. — Mesonotum about as wide as long; prescutum and scutum microsetose throughout (in ribeiroi, distinctly longer than broad, scutum and baso-lateral angles of prescutum microsetose). Anterior margin of prescutum only moderately projecting at middle (strongly in ribeiroi) ; median suture extending posteriorly to transverse suture, the latter slightly sinuous and poorly denned at middle; chaetotaxy in general much as Fig. 118. A, C, Pseudostrebla greenwelli, new species; A, left male gonapophysis, holotype; C, hind tibia, paratype male. B, D, same structures, P. ribeiroi Costa Lima, from Tonatia "amblyotis" (= silvicola), Rio Guapaya, La Macarena (Meta), COLOMBIA. ribeiroi, but less dense and setae paler; scutum at middle with two irregular rows of setae between long antescutellar scutal setae and transverse mesonotal sutures (three in ribeiroi) ; mesepisternum (viewed from above) with coarse setae along inner and posterior margins, with very short setae laterally. Wings. — Very similar to those of ribeiroi but the coarse setae a little shorter. Legs. — Fore-, mid-, and hindlegs progres- sively longer. Inner face of pro- and metafemora covered with short setae, setae of outer face a little longer; profemora with a row of longer setae along ventro-lateral margin, a few long setae dorsally and apico-laterally; mesofemora with about two pairs of conspicuously longer setae dorso-apically, a subapical seta on each side, and five or six along ventro-lateral margin; hind femora with a row of about six longer setae on basal half of dorsal margin and a double row of six macrosetae, and another pair of macrosetae distally on each side on lateral and ventro-lateral margins. Protibiae with about five setae along dorsal margin, that are distinctly longer than the others; meso- tibiae with five conspicuously longer setae along dorsal margin. Hind tibiae (fig. 118C) WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 585 Fig. 119. Stizostrebla longirostris Jobling. A, female, dorsal view. B, left male gonapophysis. C, labium. D, venter of head and thorax. A, C, and D from Jobling (1949b). B, from specimen off Tonatia sp. (CNHM no. 87942), Los Micos, San Juan de Arama (Meta), COLOMBIA. 586 ECTOPARASITES OF PANAMA a little shorter than hind femora, with five macrosetae. Hind tarsi slender, about three- fourths as long as tibiae; last segment strongly, laterally compressed. Abdomen. — Very similar to that of ribeiroi. Lateral lobes of tergum I+II with eight to ten long setae and eight to ten short ones (about 19 coarse macrosetae and eight to ten shorter setae in ribeiroi) . Sternum II as in ribeiroi but less strongly setose. Female: Tergum VII and supra-anal plates apparently fused, but basal portion longer, triangular, with a pair of short setae near apex of triangle, which points anteriorly; triangular portion continuous distally with a very small crescent-like sclerite ( ? supra- anal plate) which bears a pair of long setae and a pair of short setae. Ventral arc flat, crescentic, non-setose. Seventh sternites triangular, inner edges parallel; with 10-12 short setae anterior to long apical setae, four of which are macrosetae (about 15 discals in ribeiroi). Male: Gonapophyses (fig. 118A) rather evenly tapered apically, with a row of microsetae on each side (abruptly tapered apically in ribeiroi, and without row of microsetae). Measurements: BL TL WL ww Male 1.70-1.76 0.58-0.59 1.24-1.26 0.60-0.63 Female 1.81 0.62 1.29 0.66 TYPE MATERIAL : Holotype male and allotype female (slides) from Tonatia minuta (host no. 11764) , Armila (San Bias) , 20 March 1963, C. 0. Handley, Jr. and F. M. Greenwell. In the collection of Chicago Natural History Mu- seum. Paratype, a male, same data but 25 March ; in the collection of the United States National Museum. Genus Stizostrebla Jobling Stizostrebla Jobling, 1939, Arb. Morph. Tax. Ent., 6: 269 (discussion), 270 (keyed), 273 (diagn.). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed) ; 1942, Bol. Ent. Venez., 1: 86 (keyed). Jobling, 1949, Parasitology, 39:321 (keyed). Type-species: Stizostrebla longirostris Jobling, 1939. This genus contains the single described species, S. longirostris Jobling. Stizostrebla longirostris Jobling. Figure 119. Stizostrebla longirostris Jobling, 1939, Arb. Morph. Tax. Ent., 6: 273, fig. 2 — Brazil, "from a bat" (Deutsches Entomologisches Institut). In the collection of Chicago Natural History Museum is a series of eight specimens of this species, collected from Tonatia sp., apparently T. silvicola, at San Juan de Arama, Los Micos (Meta) , COLOMBIA by Kjell von Sneidern. This host occurs in Panama, and it is possible that the parasite does too. Genus Eldunnia Curran Eldunnia Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, pp. 5 (keyed), 6 (diagn.). Jobling, 1936, Parasitology, 28: 357 ff. (morph.), 370 (keyed) ; 1939, Arb. Morph. Tax. Ent., 6, (3), pp. 269 (discussion), 270 (keyed) ; 1949, Parasitology, 39: 322 (keyed). Bequaert, 1940, Revista Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed) ; 1942, Bol. Ent. Venez., 1: 84 (keyed). This monotypic genus is unique in having a postgenal ctenidium of only about 18 spines, limited to the underside of the head. The male genital ap- paratus is also distinctive. In addition to characters given by Curran, the following may be added : WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 587 Head. — Laterovertices transverse, without dark lines or sutures. Eyes small, with six facets. Occipital lobes and postgenae forming prominent lateral lobes which fit into deep concavities on anterior margin of thorax. Postvertex a sclerotized transverse strip which bears setae. Underside of head on each side of oral cavity with a pale oval area like that of Pseudostrebla and Parastrebla. Legs. — Hind tibiae with a Fig. 120. Eldunnia breviceps Curran. Head and thorax, dorsal view, male from Lonchophylla robusta (no. 12213), Puerto Obaldia (San Bias). subapical seta that is distinctly stronger than the others. Abdomen. — Female: Tergum VII feebly sclerotized, transverse. Seventh sternites triangular. Cerci free, ventral arc feebly sclerotized. Male: Cerci free, without apodemes that extend to surstyli. Gona- pophyses (fig. 121) short, strongly narrowed distally, with several pairs of ventral setae; aedeagus blade-like, but strongly narrowed apically, not coiled at base. Its very distinct male genital apparatus notwithstanding, Eldunnia ap- pears to share more characters, especially of head structure, with Pseudo- strebla and especially Stizostrebla, than with any other genera. It does not appear to be closely related to the genera which we regard as Streblinae. 588 ECTOPARASITES OF PANAMA Eldunnia breviceps Curran. Figures 39C, 120, 121. Eldunnia breviceps Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479; 1935, Amer. Mus. Novit., no. 765, p. 6 (descr.), figs. 7, 9 — Chilibrillo Caves, Panama, from Loncho- phylla robusta (American Museum of Natural History). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418. Jobling, 1949, Parasitology, 39: 316 ff., fig. 2B. Excepting the scutellum, the mesonotum of this remarkable species is covered with microsetae (fig. 120). Fig. 121. Eldunnia breviceps Curran. Male genital apparatus, left lateral view. Same data as for specimen in fig. 120. Measurements : Male Female BL 1.32-1.47 1.26-1.70 TL 0.49-0.51 0.49-0.52 WL 1.10-1.18 1.25-1.37 ww 0.49-0.60 0.60-0.66 PANAMANIAN MATERIAL EXAMINED : In addition to the type and para- type, we have examined 18 specimens in 11 lots (12 bats) , from Lonchophylla robusta as follows: holotype female, Chilibrillo Caves, Chilibre (Panama), 30 January 1931, L. H. Dunn [AMNH], and the paratype, same data but 9 March 1933 [MCZ] ; 1, same locality, 27 March 1958, GML; 5 (3 bats), Almirante (Bocas del Toro), 23 arid 31 January 1960; 4 (2 bats), Isla Bastimentos (Bocas del Toro) , 30 January and 1 February 1963 ; 4 (3 bats) , Buena Vista caves (Colon), 15 September and 19 November 1959; 2 (2 bats), Armila (San Bias), 14 March 1963, C. O. Handley, Jr. and F. M. Green well ; 2 (1 bat) , Puerto Obaldia (San Bias) , 4 April 1963, Handley and Greenwell. REMARKS : This species is known only from Panama. It appears to be restricted to Lonchophylla robusta, though it is possible that it may occur WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 589 on other species of the genus. None were taken from the few specimens of Lonchophylla mordax and L. thomasi that were collected in Panama. Of ap- proximately 55 specimens of L. robusta that were examined, only 11 (20% ) were positive for E. breviceps. In only one instance were more than two flies collected from a single host. Although Curran (1935) refers to page 479 of the Fam. Gen. N. Am. Dipt, as the original citation for E. breviceps, there is a question in our minds as to whether or not this (1934) usage was a nomen nudum. The genus Eldunnia was keyed out, and E. breviceps was given in parentheses as the included species. Because the paratype lacks the abdomen, its sex is unknown. Subfamily Streblinae As defined by Jobling (1936), the subfamily Streblinae would include the following genera : Anastrebla (= Strebla auct.) , Strebla (= Euctenodes Waterhouse), Metelasmus, Paraeuctenodes, Pseudostrebla, Parastrebla, n. gen., Stizostrebla, and Eldunnia. Jobling characterized the subfamily on the basis of shape and width of the head, shape and position of the palpi, and shape and relative size of the mesonotum. The palpi of Pseudostrebla, Parastrebla, and of Stizostrebla and Eldunnia exhibit a wide range of shape and width, including that (e.g., Pseudostrebla greenwelli n.sp.) typical of Trichobiinae. In none of these are they modified to form a shield-like front as in the other genera assigned to the Streblinae by Jobling. The other characters, too, vary. They seem to represent spe- cializations toward a polyctenoid body form, and probably have evolved in- dependently at least a couple of times. The ventral accessory setae of the male gonapophyses in Pseudostrebla and Stizostrebla are inserted posterior to the macrosetae, as in the uniformis, longipes, and dugesii groups of Trichobius and in the genera Speiseria, Noctiliostrebla, Aspidoptera, Mastoptera, Exastinion and Paradyschiria. The male of Parastrebla is unknown, but we predict that it will be of this type. In Strebla, Anastrebla, Paraeuctenodes, and Metelasmus, the ac- cessory setae are anterior to the macrosetae, as in the major-sparsus and caecus groups of Trichobius, and in the genera Megistopoda, Paratricho- bius, Neotrichobius, Joblingia, and Anatrichobius. These and some other characters, including female genital structure, appear to reflect natural groupings. If this is so, then the Streblinae of Jobling include convergent types. Until the relationships of the genera can be investigated more thoroughly, we restrict the Streblinae to those genera which have a complete postgenal ctenidium, which extends around the side of the head to the dorsal surface. The relationships of Eldunnia are puzzling. The male genital ap- paratus is strikingly different from that of any other New World Streblidae. However, the structure of the head appears to relate it to Stizostrebla and Pseudostrebla. In the following section, we have used some special terms which apply only to the Streblinae. The laterovertices are divided into two or more distinct areas by dark, internal lines or ridges. The area behind the eye is often set off separately by such an internal ridge and bears a single seta. 590 ECTOPARASITES OF PANAMA B Fig. 122. Dorsum of head. A, Strebla carolliae, new species (composite drawing from several specimens off Carollia p. azteca, from localities in Panama). B, Strebla hertigi, new species, from Phyllostomus h. panamensis (no. 4651), Chepo Road (Panama). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 591 We have called this area the "postocular sclerite." Behind the anterior mar- gin of the prescutum on each side, extending inwardly from the spiracle is a row of stout bristles of variable number, which we call the "epaulets" (fig. 134). Farther behind on each side of the prescutum is an arcuate row of setae, which are usually conspicuously longer than the other discal prescutal setae. We call these the "prescutal arc" of setae (fig. 134) . We have called the underside of the head, anterior to the ctenidium, the "ante-ctenidial area." Its length is measured along the mid-line between the anterior margin of the ctenidium and the anterior margin of the palpi ; its width is measured between the lateral margins of the postgenae, in front of the ctenidium. Measurements of body length were made from the ante- rior margin of the palpi, rather than of the frontoclypeus, as was done for the other Streblidae. Genus Strebla Wiedemann (=Euctenodes Waterhouse) , neiu synonymy. Strebla Wiedemann, 1824, Analecta Ent., p. 19; 1830, Aussereurop. Zweifl. Ins., 2: 612. Macquart, 1835, Hist. Nat. Ins., Dipt., 2: 637. Walker, 1849, List. Dipt. Ins. Brit. Mus., pt. 3, p. 1146 (part.). Kolenati, 1856 (& 1857), Parasiten d. Chiropteren, p. 46. Rondani, 1878, Ann. Mus. Civ. Stor. Nat. Geneva, 12: 167. Bigot, 1885, Ann. Soc. Ent. Fr., (6), 5: 226, 232 (keyed). Williston, 1896, Man. Fam. Gen. N. Am. Dipt., 2nd ed., p. 152 (keyed). Coquillet, 1910, Proc. U. S. Nat. Mus., 37: 609. Euctenodes Waterhouse, 1879, Trans. Roy. Ent. Soc. Lond., 1879: 310 (with E. mirabilis Waterhouse, as type-species). Bigot, 1885, Ann. Soc. Ent. Fr., (6), 5: 226 (discuss.), 233 (keyed). Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 41, 42, 63, 66 (keyed). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 26 (keyed), 32 (cit.). Kessel, 1924, Parasitology, 16: 406-409 (char.); 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 30 (char., key to spp.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 654. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1934, Bull. Amer. Mus. Nat. Hist., 66: 522 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 355 ff. (morph., syst. pos.), 370 (keyed). Pessoa and Gui- maraes, 1937, Ann. Fac. Med. Sao Paulo, 12: 255 ff. (comp. notes). Jobling, 1939, Arb. Morph. Tax. Ent., 6: 269 (discuss.), 270 (keyed). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed); 1942, Bol. Ent. Venez., 1: 86 (keyed). Schuurmans-Stekhoven, Jr., 1951 (& 1941?), Beitrage Fauna Perus, 3: 96. Jobling, 1949, Parasitology, 39: 322 (keyed). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 178 (keyed), 188 (records). Maa, 1965, Jour. Med. Ent., 1 : 386. Type-species : Hippobosca vespertilionis Fabricius, 1805. Wiedemann (1824, p. 19) based his genus Strebla on Hippobosca ves- pertilionis Fabricius. He also gave a short description and figure of the type specimen of vespertilionis, and indicated its location as "In Museo Regio Havniensi." His original figure (in color) did not show setae on the hind legs. The dark structures he figured at the basal angles of the head (and later interpreted to be eyes) are the dorsally curved ends of the ctenidium. The presence of the ctenidium and the shape of the thorax clearly show the insect to be a member of the Streblinae. The relatively short hindlegs indicate, however, that the insect is a species of Euctenodes Waterhouse, not of Strebla, Speiser (1900), Kessel (1924) and later authors. 592 ECTOPARASITES OF PANAMA H Fig. 123. Postvertex and occipital plates (of head) of species of Strebla. A, S. mirabilis (Waterhouse) from Phyllostomus h. panamensis (no. 4138), Natural Bridge, Madden Dam (Canal Zone). B, S. vespertilionis (Fabricius), from Desmodus r. murinus (no. 5236), Red Tank (Canal Zone). C, S. kohlsi, new species, from Tonatia s. silvicola (CNHM nos. 69897-99), Cauca Valley above Caceres, Puri (Antioquia), COLOMBIA. D, S. galindoi, new species, paratype. E, S. alvarezi, new species, from Saccopteryx bilineata (no. 4659). F, same, from Saccopteryx bilineata, Finca Santa Cristina, (Escuintlas), GUATEMALA. G, S. altmani, new species, paratype from Lonchorhina aurita. H, S. hoog- straali, new species, from Tonatia minuta (no. 9162). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 593 In 1830, Wiedemann added his interpretation and description of the "eyes" and slightly altered the illustration (now black and white) to change the wing somewhat and to add conspicuous setae on the hind tibiae. Mac- quart's (1835) figure is copied from this. Wiedemann again referred to the specimen as being in the Copenhagen Museum and it is assumed that the Fig. 124. A-E, postvertex and occipital plates (of head) of species of Strebla. A, S. diaemi, new species, paratype from Diaemus youngii, Rio Raposo, COLOMBIA. B, S. consocius, new species, paratype from Phyllostomus h. hastatus (TVL nos. 1406-21) Guanapo Heights, TRINIDAD. C, S. diphyllae, new species, paratype from Diphylla ecaudata centralis, San Lorenzo, N.E. of Volcan de Jumay (Jalapa), GUATEMALA. D, S. machadoi, new species, holotype. E, S. christinae, paratype from Phylloderma stenops (no. 11986). F, antennae and frontoclypeus, S. galindoi, new species. revised figure, too, was based on the type. Poor though the figure is, the illustration of the hind tibiae clearly shows the fly to be a species of Euctenodes, since the species regarded as Strebla vespertilionis by Speiser and Kessel (loc. cit.) has elongated hindlegs and lacks metatibial macrosetae. Thus, Euctenodes should be synonymized under Strebla, and a new name is needed for Strebla (not Wiedemann, 1824) of Speiser, Kessel et al. 594 ECTOPARASITES OF PANAMA The genus Strebla is taxonomically very difficult. In the past, a number of closely related species have been confused under mirabilis ( Waterhouse) . Only one other species was described (tonatiae Kessel). In our studies, we have segregated 15 species, 12 of them new. We considered it desirable to present a preliminary revision of the genus here, since 12 of the 15 species have been collected in Panama. After this study was completed, three ad- ditional new species from Colombia, Venezuela, and Surinam came to our attention, and it is likely that more will be discovered in the future. Further, some of the host data suggest that cryptic species exist which we presently are unable to separate on morphological grounds (see remarks under S. mirabilis) . Many of the species appear to be restricted to a single host. r I B D Fig. 125. Metatibiae, dorsal view. A, Strebla consocius, new species. B, S. vesper- tilionis (Fabricius). C, S. diaemi, new species. D, S. galindoi, new species. E, S. mirabilis (Waterhouse). F, S. hertigi, new species. KEY TO THE SPECIES OF STREBLA 1. Frontoclypeus entire (fig. 122B). The second, anterior pigmented mesonotal suture lacking or indistinct (fig. 127B) 2 Frontoclypeus with a pair of apical detached plates (fig. 122 A) or with an unsclerotized median suture (fig. 124F). Second anterior pigmented meso- notal suture usually distinct, as in fig. 134, though absent in christinae n.sp 6 2. Upper edge of posterior tibiae with two or three macrosetae (fig. 125D-F). An- terior margin of postvertex as in fig. 122B. Eyes consisting of a single row of about four facets. Hosts: Phyllostomus spp hertigi n.sp. Upper edge of posterior tibiae with six to eight macrosetae (fig. 125A-C) 3 3. Anterior margin of postvertex forming an obtuse angle (fig. 124A). Prescutum (fig. 127 A), anterior to the transverse mesonotal suture, broader than long; with a row of short bristles extending across entire width behind the anterior margin. Host: Diaemus youngii diaemi n.sp. Anterior margin of postvertex forming an angle less than or only slightly greater than 45° (figs. 123H, 124B). Prescutum, anterior to the transverse suture, longer than broad ; short, stout bristles near anterior margin limited to a group on each side, near the anterior angles (figs. 128, 129) 4 WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATPLIES 595 4. Bristle at apex of third longitudinal vein only slightly larger and stronger than the preceding "erect" setae. Angle formed by the anterior margin of the postvertex about equal to or slightly greater than a right angle (fig. 124B). Eyes small but distinctly faceted. Host: Phyllostomus hastatus . . .consocius n.sp. Bristle at apex of third longitudinal vein a strong and conspicuous macroseta (fig. 126) which is at least twice as long as the preceding erect setae. Eyes consisting of a single hyaline lens, facets indistinguishable. Angle formed by the anterior margin of the postvertex conspicuously less than a right angle (fig. 123H ) . Hosts : Tonatia spp 5 5. Mesonotum (fig. 129B) , more sparsely setose, the prescutum with only about 25-28 setae on each side behind the epaulets; scutum laterally with two irregular rows of setae between the antescutellar row and the transverse mesonotal suture ; setae of antescutellar row subequal. Host : Tonatia minuta hoogstraali n.sp. Mesonotum (fig. 129A), more densely setose, the prescutum with about 42-46 setae on each side behind the epaulets; scutum, toward the sides, with three irregular rows of setae between the antescutellar row and the transverse suture ; some of the lateral setae of antescutellar row distinctly longer than the median ones. Host: Tonatia brasiliensis tonatiae (Kessel) 6. Frontoclypeus with a median unsclerotized suture (fig. 124F). Anterior margin of postvertex as in fig. 124B ; setae of postvertex and occiput long and slender. Host : Tonatia bidens galindoi n.sp. Frontoclypeus with a pair of apical detached plates (fig. 122A) 7 7. Upper edge of posterior tibiae with two rows of setae which are conspicuously longer than the others, the apical ones longest 8 Upper edge of posterior tibiae either with a single row of longer setae of which the apical two or three are macrosetae or with only two or three subapical macrosetae that are distinctly longer than other tibial setae 10 8. Eyes consisting of a single elongate hyaline lens. Anterior margin of postvertex subacuminate (fig. 124D). Mesonotal chaetotaxy as in fig. 131B. Host: Micronycteris minuta machadoi n.sp. Eyes multi-faceted. Anterior margin of postvertex not thus 9 9. Second or anterior transverse mesonotal suture absent. Mesonotal chaetotaxy as in fig. 131A. Postvertex as in fig. 124E, the spinelets relatively short and stout, as are the inner three spinelets of occiput on each side. Female: Tergum VII with two pairs of setae. Anterior margin of ventral arc of terminal cone produced as a broad rounded lobe (fig. 44A). Male: Gonapophyses (in lateral view) strongly curved. Host: Phylloderma stenops christinae n.sp. Second or anterior transverse mesonotal suture present ; mesonotal chaetotaxy as in fig. 132. Postvertex as in fig. 123B, the spinelets more slender, as are the inner three occipital spinelets on each side. Female: Tergum VII typically with three pairs of setae. Anterior margin of ventral arc not produced. Male: Gonapophyses (lateral view) only very feebly curved, nearly straight. Host: Desmodus rotundus vespertilionis (Fabricius) 10. Anterior margin of postvertex broadly blunt (fig. 122A). Hosts: Carollia spp. carolliae n.sp. Anterior margin of postvertex much more strongly rounded, or angulate 11 11. Inner occipital setae fine, bristle-like, not spinelets (fig. 123G). Hosts: Lonchorhina aurita, Macrophyllum macrophyllum altmani n.sp. At least one or two, usually three or four of inner occipital setae on each side are spinelets 12 12. Anterior margin of postvertex as in fig. 123E, F. Prescutum (fig. 137B) with only two irregular transverse rows of setae laterally between the transverse mesonotal suture and the anterior suture alvarezi n.sp. Anterior margin of postvertex otherwise (figs. 123A, C; 124C). Prescutum (figs. 133, 134) with at least three irregular transverse rows of setae laterally be- tween the transverse mesonotal suture and the anterior suture . . . 13 596 ECTOPARASITES OF PANAMA 13. Prescutum (fig. 133) densely setose anteriorly; median discal setose area extend- ing anteriorly nearly to margin, the antero-lateral setose areas (anterior to arcs) extending along sides nearly to epaulets. Postvertex and occiput as in fig. 124C, their setae strong, but not stout spines or spinelets. Host: Diphylla ecaudata diphyllae n.sp. Prescutum (fig. 134) sparsely setose anteriorly; median discal setose area ex- tending only to about middle, distinctly separated from antero-median paired setae; area behind epaulets along lateral margins bare 14 14. Head elongate, the ventral ante-ctenidial area only slightly wider than long (24:22). Detached frontoclypeal plates distinctly longer than wide. Host: Tonatia silvicola kohlsi n.sp. Head short; ventral ante-ctenidial area nearly half again as wide as long (59 :41) Hosts: Phyllostomus hastatus panamensis, (?) Trachops cirrhosus mirabilis (Waterhouse) Fig. 126. Wing, Strebla hoogstraali, new species, male paratype from Tonatia silvicola (no. 9102), Las Palmitas (Los Santos). Strebla hertigi Wenzel, new species. Figures 122B, 125F, 127B. A very distinctive species not closely related to any other, that is easily recognized by the following combination of characters : frontoclypeus en- tire (without apical detached plates), eyes consisting of only four or five ocelli in a row; metatibiae with only two or three macrosetae; setae and macrosetae of occiput and spines of postvertex comparatively short ; dorsal plate of postgenae with a postero-lateral emargination ; postvertex long, subquadrate. DESCRIPTION: Head (fig. 122B). — Frontoclypeus without detached apical plates. Anterior plate of laterovertices with five to six setae; postocular sclerite distinct. Chaetotaxy of head as illustrated, except that long, outer setae of occiput are a little more slender than shown. Postero-lateral margin of dorsal sclerite of postgenae emarginate for reception of ctenidium. Eyes with four or five facets in a single row. Ventral ante-ctenidial area about as wide as long (24:23). Postvertex long, its anterior margin obtusely angulate. Thorax (fig. 127B). — Prescutum without a second pigmented suture; epaulets con- sisting of five or six setae which extend across anterior margin to median pair; prescutal WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 597 setae extending anteriorly to anterior margin ; arcuate row of long setae present but not conspicuously differentiated. Setose area posterior to arc consisting of about four ir- regular transverse rows at middle, three toward sides. Antescutellar scutal setae mostly subequal, not distinctly longer laterally, except for long macroseta at extreme lateral margin; two rows of scutal setae in addition to antescutellars, anterior half of scutum bare toward sides. Wings. — Rs approximately a third again as long as distance from fork to r-m (25:17.5). All veins setose to base, excepting the sixth longitudinal. Legs. — Protibiae with five exceptionally long macrosetae, mesotibiae with three. Hind tibiae (fig. 125F) apically with three macrosetae, the distal one less than half as long as the basal one. First segment of hind tarsi with a median row of heavy plantar bristles, one or two outwardly near base, one inwardly at apex and several fine ones anterior to this ; plantars of following segments minute. Abdomen. — Anterior face of tergum I+II with three short setae; inner dorsal margin with three to five setae, two very fine. Female: Tergum VII with two pairs of macro- setae ; the second, posterior pair more than half as long as anterior pair, sometimes with a supernumerary. Supra-anal plate with four long, apical macrosetae and a pair of short but well-developed basal setae. Seventh sternites with 13-16 setae consisting of four macrosetae (two very long), three to five intermediate setae, and four to nine short setae. Ventral arc triangularly produced anteriorly, with a single pair of setae. Male : Hypopygium dorso-laterally on mid-third with a macroseta and one or two shorter setae; dorso-apical margin with six to eight macrosetae; a long apical macroseta below these on each side. Gonapophyses nearly straight, slightly sinuate above, the macroseta inserted at about basal fourth. Sternum V at middle with two transverse rows of setae in addition to marginal setae. Measurements: BL TL WL ww Male 2.56-3.13 0.85-0.88 1.87-1.95 0.84-0.93 Female 2.56-3.08 0.88-0.93 1.87-2.14 0.91-0.96 TYPE MATERIAL: Holotype male and allotype female (slides) from Phyllostomus hastatus panamensis (host no. 4650) , hollow tree, Chepo Road (Panama) , 8 October 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes: PANAMA. — From P. h. panamensis: 38 (12 bats), same data as the holotype; 1 (20 bats) , Chilibrillo Caves (Panama) , 17 July 1959 and 1, 28 October 1959; 1, Tapia (Panama), 25 July 1923, J. P. Chapin [MCZ] ; 1, Fort Sherman (Canal Zone), 6 April 1960; 3 (2 bats), Natural Bridge, Madden Dam (Canal Zone) , 31 August 1959 ; 2, Almirante (Bocas del Toro) , 29 January 1960. From Phyllostomus d. discolor: 1, Rio Tuira (Darien), 10 March 1958, P. Galindo [GML] ; 3, Rio Mandinga (San Bias), 18 and 29 May 1957, P. Galindo [GML] ; 1, Armila (San Bias), 1 April 1963, C. 0. Handley Jr. and F. M. Greenwell. From Lasiurus egregius (mist net) : 1, Armila (San Bias), 23 February 1963, Handley and Greenwell. From Artibeus j. jamaicensis: 1, Natural Bridge, Madden Dam (Canal Zone), 31 August 1959. From Desmodus r. murinus: 1, Guanico (Los Santos), 26 January 1962. COSTA RICA. — From Phyllostomus h. panamensis: 1, Playa del Coco, Guancaste, 20 August 1962, F. Truxal, Los Angeles County Museum. NICA- RAGUA.— From Phyllostomus h. panamensis, 5 km. N and 14 km. E of Condega : 2 (1 bat) , 23 June 1964, J. Knox Jones [KU] ; 8 (1 bat) , 25 June 1964, J. D. Smith [KU] ; 19 (1 bat), 24 June, and 10 (1 bat), 25 June 1964, T. E. Lawlor [KU]. EL SALVADOR [H. Felten, SNI], — From Phyllostomus discolor verrucosus: 5, Hacienda San Diego (La Libertad), 15 November 1953 ; 1, same data but 8 February 1953 ; 5, San Rafael Cedros (Cuscatlan) , 598 ECTOPARASITES OF PANAMA 7 September 1953; 496 (1 lot), "Hohle Ceiba pentandra, an Strasse San Salvador-Sta. Ana bei San Andres" (La Libertad), 482 meters elevation, 27 September 1953; 11, "Hohle bei Suchitoto (Cuscatlan)," 7 June 1953; From Desmodus rotundus: 1, "Hohle Ceiba pentandra, bei km. 31, Strasse San Salvador, Sta. Ana," 4 August 1953. Without host: 2 lots, 27 speci- mens. GUATEMALA. — Without host: 5, Patulul, April 1906. COLOMBIA. — From Phyllostomus discolor: 1, Cucuta, May 1940. VENEZUELA. — From Phyllostomus d. discolor: 1, Biological Station, Rancho Grande (Aragua), B Fig. 127. Thorax, dorsal view. A, Strebla diaemi, new species, male paratype from Diaemus youngi, Rio Raposo, COLOMBIA. B, S. hertigi, new species, male paratype from Phyllostomus discolor, Port-of-Spain, TRINIDAD. 31 August 1962, C. Machado and R. Antequera [FCUCV]. TRINIDAD (TVL unless otherwise indicated). — From Phyllostomus discolor [discolor]: 2, Alta Gracia Trace, Siparia, 20 January 1958; 31, Long Circa, Port-of- Spain, 2 August 1954 ; 6, Masson Hospital, Port-of-Spain, 29 October 1956 ; 7, Emperor Valley Zoo, Port-of-Spain, 13 and 18 March 1958 ; 1, Santa Cruz, 28 June 1956 ; 2, Port-of-Spain, C. C. Sanborn, 8 November 1954, CNHM Trinidad Zoological Field Trip (1954). BRITISH GUIANA. — Without host: 2, Kartabo, Bartica District, 1921, A. E. Emerson. SURINAM. — From Phyllostomus discolor: 3, Kaiserberg Airstrip, east of Zuid River, 12 Octo- ber 1960, and 4, same locality, 12 December 1960, H. A. Beatty, CNHM Guianan Zoological Expedition, 1960-62. PERU. — From Phyllostomus dis- color: 3, Hacienda Bogetes, Salitral (Piura), 23 April 1959, 180 meters ele- vation, C. Kalinowsky, CNHM Peruvian Zoological Expedition. Paratypes to be deposited in the collections listed on p. 410. OTHER MATERIAL EXAMINED : We have at hand dozens of additional lots collected in MEXICO, COSTA RICA, and NICARAGUA from Phyllostomus discolor WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 599 verrucosus by J. Knox Jones [KU] and collaborators and by Fred Truxal and collaborators [LACM] . We have not yet tabulated these records. REMARKS : This species is named in honor of Dr. Marshall Hertig, dis- tinguished medical entomologist of the Gorgas Memorial Laboratory. S. hertigi may occur on Phyllostomus d. discolor throughout most, if not all of its range ; but we have seen specimens only from the tropical lowlands, from Mexico to Peru and northeastern South America. In Panama, it often occurs together with S. mirabilis on Phyllostomus h. panamensis, but per- haps only where ecological situations are favorable for its transfer to that host from P. discolor. Interestingly, S. mirabilis does not seem to occur on P. h. panamensis in Costa Rica and Nicaragua, but is entirely replaced by hertigi. For further discussion, see p. 645, and following paper, p. 683. Strebla diaemi Wenzel, new species. Figures 124A, 125C, 127A. Similar to S. hertigi n. sp., S. consocius n. sp., and S. tonatiae (Kessel) in having the frontoclypeus entire. Among these, it resembles hertigi in the shape of the postvertex, but differs conspicuously in having seven meta- tibial macrosetae (rather than two or three) and in having convex, multi- faceted eyes, whereas hertigi has a single row of four facets. It resembles consocius in the metatibial setae and eyes, but differs markedly from that species in the shape of the postvertex as well as in other characters given in the key. DESCRIPTION : Head. — Frontoclypeus entire, without apical detached plates. Anterior plate of laterovertices with about seven setae; postocular sclerite distinct. Dorsal sur- face of postgenae not emarginate laterally; posteriorly with about 15 short setae and, near inner posterior margin, a conspicuous macroseta of about same length as the long, outer occipital setae; lateral to the macroseta is a shorter seta of about same length as inner occipitals. Postvertex and occiput as illustrated (fig. 124A), but the setae usually more nearly like those shown on right side. Eyes multi-faceted and projecting, seven or eight facets visible from above. Ventral ante-ctenidial area more than half again as wide as long (29.5:18.5). Thorax (fig. 127A). — Prescutum with one pigmented suture; epaulets consisting of eight or nine setae on each side and extending inwardly on each side to paired median setae behind anterior margin; arc of longer setae not well-defined, though the lateral setae of arc are conspicuous. About six to eight median antescutellar setae scarcely longer than the scutal setae anterior to them; on each side of these are three or four longer setae, one conspicuously longer ; to each side of antescutellars are two prominent setae, one a very long macroseta. Wings. — All veins except sixth longitudinal, setose to base. Costa without a macroseta at apex. Rs about two and a half times as long (29 :11) as distance from fork to r-m. Legs. — Pro- and mesotibiae dorsally each with eight or nine conspicuous macrosetae in a double row; metatibiae (fig. 125C) with seven macro- setae, the apical ones about as long as the first three tarsomeres combined. First segment of hind tarsus elongate, slightly longer than next three segments combined; ventrally with a median longitudinal row of five very heavy, long plantar bristles, and lateral to these, two others on basal half of outer edge and three or four slightly weaker ones along inner edge. Plantar bristles of succeeding segments very small, relatively in- conspicuous. Abdomen. — Dorsal inner margin of lateral lobes of tergum I+II with three to four setae. Female : Tergum VII with a pair of very long macrosetae and, posterior to these, two pairs of setae in tandem, the anterior of these shorter, the posterior pair about two- thirds to three-fourths as long as the anterior macrosetae. Supra-anal plate with four apical macrosetae and on each side, near base, one or two pairs of shorter, conspicuous 600 ECTOPARASITES OF PANAMA setae. Seventh sternites with 12-15 setae, four to seven of these being macrosetae. Ventral arc produced anteriorly as a finger-like process, its posterior edge with eight fine setae, the two median setae much shorter than the others. Male: Hypopygium very long and prominent; sternum VII+VIII dorsally with two conspicuous macrosetae a little beyond middle, and medial and slightly posterior to these, two setae that are more than half as long as the macrosetae; tergum IX dorsally with a row of four macrosetae. Gonapophyses gradually tapered and curved from base to knobbed apices; sides with scattered microsetae, chiefly on apical third, a few anterior to macroseta ; ventral macro- seta inserted at about apical third ; accessory seta minute. Measurements: BL TL WL WW Male 2.53-2.80 0.88-0.93 1.76-1.92 0.82 Female 2.89-3.16 0.93-0.99 1.87-1.92 0.82-0.88 TYPE MATERIAL : PANAMA. — Holotype male and allotype female (slides) , from Diaemus youngii cypselinus (host no. 11422), Armila (San Bias), 4 March 1963, C. 0. Handley, Jr. and F. M. Greenwell. In the collection of Chicago Natural History Museum. Paratypes. — 6, same data as the holotype; 4, same host and collectors, Isla Bastimentos (Bocas del Toro). COLOMBIA. — 7, same host, Rio Raposo (Valle de Cauca), 10 August 1962, C. J. Marinkelle and W. Thornton; 1, same host, Rio Mecaya (Caqueta), 1851 meters elevation, 2 March 1952, P. Hershkovitz, CNHM Colombian Zoological Expedition (1948-52) . PERU.— 14, same host, Rio Calleria, Colonia Calleria, "15 km. from [?Rio] Ucayali" (Loreto), 8 October 1961, B. Malkin and J. S. Hurley. Paratypes in the collections of Chicago Natural History Museum ; the United States National Museum ; and C. J. Marinkelle, Universidad de los Andes, Bogota, Colombia. OTHER MATERIAL EXAMINED : 2, from "bats," Orinoco River, VENEZUELA, collected by Olalla, [MCZ, det. as mirabilis by Bequaert]. REMARKS : The only host known for this species is Diaemus youngii, a vampire bat which often attacks poultry (Goodwin and Greenhall, 1961) . Strebla consocius Wenzel, new species. Figures 124B, 125A, 128. Closely related to S. tonatiae (Kessel) but differing in the greater angle formed by the anterior margin of the postvertex, the convex, multi-faceted eyes (a single hyaline lens in tonatiae), and in not having a conspicuous macroseta at apex of costal wing vein. DESCRIPTION : Head. — Frontoclypeus without apical detached plates. Anterior plate of laterovertices with six setae; postocular sclerite distinct. Dorsal surface of post- genae not emarginate laterally, posteriorly with about 10 short setae, as well as a con- spicuous macroseta near inner posterior margin and lateral to it a shorter but conspicu- ous seta. Postvertex and occiput as illustrated (fig. 124B) but anterior margin generally a little more pointed and the festoon setae usually more like those shown on right side. Eyes relatively small, multi-faceted, about seven facets of varying sizes, visible from above. Ventral ante-ctenidial area wider than long (29:22). Thorax (fig. 128). — Prescutum without second or anterior transverse suture; epaulets usually with five to six setae ; discal setae relatively sparse, setose area not quite reaching epaulets along sides ; arc of longer setae not distinguishable, though long outer seta is present. Antescutellar setae conspicuously longer than other scutals. Wings. — All veins except sixth setose to base. Costa without a macroseta at apex. Rs only about a fourth longer (23:17) than distance from fork to crossvein r-m. Legs. — Protibiae with about eight conspicuously long macrosetae, mesotibiae with nine to ten. Metatibiae with six or seven conspicuously long macrosetae along dorsal edge, the apical ones WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 601 longest. First segment of hind tarsus about as long as next two segments combined, the ventral surface with two irregular rows of conspicuous plantar bristles and an inner row of much shorter ones; plantars of following segments minute, inconspicuous. Abdomen. — Inner dorsal margins of tergum I+II with five to six setae. Female: Tergum VII with two macrosetae and posterior and medial to them a pair of setae nearly half as long. Supra-anal plate with four apical macrosetae only. Seventh sternites Fig. 128. Thorax, dorsal view, Strebla consocius, new species, female paratype from Phyllostomus hastatus (CNHM no. 87115), Santa Rita, Iquitos (Loreto), PERU. with 11—12 setae, about five of these very long and coarse, one much longer than the others. Ventral arc very narrow, with two longer setae and eight minute ones along middle. Male: Hypopygium as in diphyllae. Gonapophyses heavy anteriorly, ventral margin nearly straight, the dorsal margin gradually curved to near apex where both are markedly curved, apices knobbed. Sides with scattered microsetae from apices nearly to base; ventral macroseta inserted slightly beyond middle, extending nearly to apex; accessory seta minute. Measurements: BL TL WL ww Male 2.86-2.97 0.96-0.99 1.76-1.90 0.82-0.93 Female 3.08-3.11 0.96-1.04 1.98-2.23 0.91-0.99 TYPE MATERIAL: Holotype male and allotype female (slides), from Phyllostomus In. hastatus (CNHM host nos. 61915-19), cave, Guanapo Heights, TRINIDAD, Frank Wonder, CNHM Trinidad Zoological Expedition (1947) . In the collection of Chicago Natural History Museum. 602 ECTOPARASITES OF PANAMA Paratypes: TRINIDAD. — From Phyllostomus h. hastatus: 9, same data as holotype; 4, same locality, 11 July 1954 and 1 same locality, 1 October 1957, T.H.G. Aitken [TVL] ; 1, Tamana Cave, Mount Tamana, 20 November 1957, T.H.G. Aitken [TVL]. From Carollia p. perspicillata: 1, same data as the preceding. Without host: 5, same locality and date as the preceding. VEN- EZUELA.— From Phyllostomus hastatus subsp. : 1, San Sebastian Cave (Aragua), 2 September 1961, Carlos Bordon [FCUCV]. SURINAM (CNHM Guianan Zoological Expedition, 1960-62) . — From Phyllostomus h. hastatus: 6, Lelydorplan (Surinam), 21 January 1962, Philip Hershkovitz; 12 (3 lots) , Kaiserberg Airstrip, east of Zuid River, elevation 900 feet, 15 October to 23 November, H. A. Beatty. From Phyllostomus sp. (probably hasta- tus) : 2, same locality and collector as the preceding, 12 October 1960. PERU (CNHM Peru Zoological Expeditions, Celestino Kalinowski). — From Phyllostomus sp. : 6, Mann, (Madre de Dios), 17 October 1954; 10, Santa Rita, Iquitos (Loreto), 16 October 1956. From Trachops sp. (!) : 9, Que- brada Esperanza, Rio Yauari-Mirim, Maynas (Loreto), 22 September 1957. Without host (removed from cloth wrappings containing numerous pre- served bats) : 1 each, presumably from Huajyumbe (Cuzco), 16 September 1950, and Marcapata (Cuzco) , 24 August 1950. REMARKS. — The specimen with doubtful host from Huajyumbe was rest- ing lightly on the fur of an alcohol-preserved specimen of Mesophylla mac- conelli. The specimen from Marcapata was doubtfully associated with Molossus rufus. No species of Strebla are known to us from these host genera. The association with Molossus seems especially dubious. The only streblids known to us from this genus are species of the genus Trichobius (see Trichobius dunni) . S. consocius seems to be restricted to Phyllostomus hastatus in north- eastern South America and Amazonian Peru. It probably occurs on this host elsewhere in the Amazonian region, but we are unable to verify this from the limited material collected in that region. Strebla tonatiae (Kessel). Figure 129A. Euctenodes tonatiae Kessel, 1924, Parasitology, 16: 411-412, figs. 7, 8 — Gualaguiza, Ecuador, from Tonatia brasiliensis (British Museum of Natural History) ; 1924, Jour. N. Y. Ent. Soc., 33: 30, pi. 4, fig. 26. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 654. Pessoa and Guimaraes, 1937, Ann. Fac. Med. Sao Paulo, 12: 255 ff., figs. 10-12. Jobling, 1949, Parasitology, 39: 316 ff. We have not seen the type of S. tonatiae (Kessel) , but our Peruvian specimens agree well with Kessel's figure and with the host of tonatiae. The identity of the host of our Peruvian specimens has been verified by Mr. Philip Hershkovitz of Chicago Natural History Museum. The host of the type of S. tonatiae has not been checked. S. tonatiae and S. hoogstraali n. sp. differ from all others known to us in possessing a long conspicuous macroseta at the apex of the costal wing vein. The eyes of both species consist of a single hyaline lens as in S. machadoi n. sp., but they differ markedly from this species in the shape of the postvertex and in having the frontoclypeus entire (without apical de- tached plates) . They appear to be most closely related to S. consocius n. sp., WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 603 from Phyllostomus h. hastatus. Though small and of nearly the same shape as in tonatiae, the eyes of consocius have distinguishable facets. Further, the postvertex is of a different shape. Measurements: BL TL WL ww Male 1.90 0.66 1.10 0.55 Female 2.67 0.82 1.21 0.55 MATERIAL EXAMINED: From Tonatia brasiliensis (host no. 75586) : a male and female, Huajyumbe (Cuzco), PERU, elevation 630 meters, 23 May 1953, Celestino Kalinowski, CNHM Peru Zoological Expedition (1953-54). A Fig. 129. Thorax, dorsal view. A, Strebla tonatiae, from Tonatia brasiliensis (CNHM no. 75586), Huajyumbe (Cuzco), PERU. B, S. hoogstraali, new species, same data as for fig. 126. Strebla hoogstraali Wenzel, new species. Figures 123H, 126, 129B. Very closely related to S. tonatiae (Kessel) but easily separated from that species by the distinctive mesonotal chaetotaxy, which is much sparser than it is in tonatiae (see fig. 129). In addition to the strikingly different chaetotaxy of the prescutum, the scutal setae of the antescutellar row are subequal in hoogstraali, while they are much longer at sides than at middle in tonatiae. DESCRIPTION : Head, — Almost identical to that of S. tonatiae. Frontoclypeus entire, without apical detached plates. Anterior plate of laterovertices with six setae. Post- ocular sclerite distinct. Postgenae posteriorly with eight short bristles and, in addition one long and one medium bristle. Postvertex and occiput as illustrated (only the two inner setae are spinelets, the others are thinner than shown) . Eyes consisting of a single elongate hyaline lens. Ventral ante-ctenidial area a little wider than long (21.5:18). Thorax. — Prescutum without second pigmented suture; epaulets with four setae; disk rather sparsely setose, setae gradually sparser and a little longer anteriorly, prescutal arc of longer setae present but not conspicuous. Antescutellar setae subequal, scutum with two irregular transverse rows of setae at sides anterior to antescutellars. 604 ECTOPARASITES OF PANAMA Wings. — Setae of wing veins relatively longer than usual in the genus; a conspicuous macroseta at apex of costa. Length of Rs approximately equal to distance from fork to r-m (12:11). Sixth longitudinal bare for only a short distance at base. Third cross- vein situated approximately midway between second crossvein and r-m. Legs. — Pro- and mesotibiae each with seven or eight macrosetae. Metatibiae with six macrosetae on dorsal margin; hind tarsus elongate, more than half as long as tibia, the first tarsomere slightly shorter than 2-4 combined, with a median row of six or seven long heavy plantar bristles and two on each side basally; remaining tarsomeres with very minute plantars. Abdomen. — Anterior face of tergum I+II with one or two short setae; dorsal inner margin with two or three fine short setae ; dorsal connexivum bare, except for four pairs of minute segmental setae; lateral connexival setae nearly twice as long as ventrals. Female: Tergum VII united with the supra-anal plate; with two pairs of macrosetae, the anterior pair longer and more widely separated than the posterior pair. Supra-anal plate with four, long, thin apical setae. Seventh sternites with about eight setae, three apical ones conspicuous macrosetae. Male: Sternum V with two transverse rows of discal setae at middle and laterally, sometimes with three. Setae of apical margin long, subequal, a little longer toward sides. Sternum VI large. Hypopygium as in alvarezi. Gonapophyses very slender, curved apically, apices knobbed; macrosetae rather short, not reaching apices of gonapophyses, inserted distal to mid-length. Measurements: BL TL WL ww Male 1.93-1.99 0.63-0.68 0.99-1.10 0.49-0.57 Female 2.06-2.26 0.66-0.71 1.04-1.18 0.51-0.62 TYPE MATERIAL (19 specimens [6 bats] from Tonatia minuta) : Holotype male and allotype female (host no. 12018) from Armila (San Bias), 29 March 1963, C. O. Handley, Jr. and F. M. Greenwell. In the collection of Chicago Natural History Museum. Paratypes. — 2, same data as the holotype ; 3, same data but 7 March ; 7, Las Palmitas, Guanico (Los Santos), 24 January 1962; 8 (3 bats), Sibube (Bocas del Toro) , 17 to 24 January 1963, C. O. Handley, Jr. Paratypes to be deposited in the collections of Chicago Natural History Museum; the United States National Museum ; the Gorgas Memorial Laboratory, Panama ; the Environmental Health Branch, USAFSC, at Corozal (Canal Zone) ; Faculdad de Ciencias, Universidad Central de Venezuela ; and the Departa- mento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil. REMARKS: This species is named for Dr. Harry Hoogstraal, Chief of Medical Zoology, United States Naval Medical Research Unit No. 3, Cairo, Egypt, in recognition of his many outstanding contributions to our knowl- edge of ectoparasites and many other aspects of medical entomology, and in appreciation of his cooperation through the years. Strebla galindoi Wenzel, new species. Figures 123D, 124F, 125D, 130. Distinct from all other species in that the f rontoclypeus is neither entire nor with apical detached plates but is intermediate in structure, having a median unsclerotized suture (fig. 124F). It is apparently related to S. mirabilis but is distinct from that species in that it has no detached fronto- clypeal plates and the anterior margin of the postvertex is broadly obtuse rather than strongly projecting. DESCRIPTION : Head. — Frontoclypeus without apical detached plates, but with a median unsclerotized suture anteriorly. Anterior plate of laterovertices with seven to eight setae ; postocular sclerites distinct. Posterior portion of dorsal sclerite of postgenae with eight short setae, and (near posterior margin) two long conspicuous ones; lateral margin WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 605 not emarginate. Postvertex and occiput as illustrated in fig. 123D (the outer seta slightly more slender than shown). Eyes multi-faceted and projecting, about eight facets visible from above. Ante-ctenidial area of underside half again as broad as long (30:20). Thorax. — Prescutum with a second anterior pigmented suture ; epaulets with five long setae. Three transverse rows of setae between transverse and second sutures. Scutum at middle with two rows of setae anterior to the antescutellars, the latter unusually long and becoming even longer laterally. Wings. — All veins, except sixth longitudinal, with setae basally. Apex of costa without a macroseta. Rs about half again as long as Fig. 130. Thorax, dorsal view, Strebla galindoi, new species, male paratype from Tonatia bidens (no. 5489), Almirante (Bocas del Toro). distance from fork to r-m (17 :11) . Legs. — Protibiae with four to five much longer setae (four stout) on outer edge, about three on inner. Mesotibiae with two oblique rows of heavy setae apically on inner face, the basal row with four setae, the apical one with two; another conspicuous macroseta along dorsal edge at middle, preceded by several long, weaker setae. Metatibiae with three macrosetae on apical two fifths, the two on dorsal margin approximately as long as first two tarsomeres combined, the third lateral in position and a little more than half as long as other two. Three rows of heavy plantar bristles on first tarsomere, minute plantars on following segments. Abdomen. — Inner dorsal margins of lateral lobes of tergum I+II with four fine setae. Connexivum with a group of three to five setae on each side of most anterior pair of seg- mental setae. Female : see Remarks. Male : Hypopygium with a dorso-lateral macro- seta and medial to it a short seta; tergum IX with two pairs of macrosetae in a trans- verse row. Gonapophyses strongly curved and rather abruptly tapered apically; sides 606 ECTOPARASITES OF PANAMA with numerous microsetae on a little more than apical third; ventral macroseta inserted at about basal third, not reaching apex, accessory seta well developed. Measurements: BL TL WL ww Male 2.47 0.82-0.93 1.65-1.76 0.82 TYPE MATERIAL : Holotype male and paratype male (slides) from Tonatia bidens (host no. 5489) , Almirante (Bocas del Toro) , 28 January 1960, C. M. Keenan and V. J. Tipton. Holotype in Chicago Natural History Museum ; paratype in the United States National Museum. OTHER MATERIAL EXAMINED: A female, from Tonatia sp., Coora T. P. D. Estate, Quiram Road, St. Patrick Co., TRINIDAD, 13 January 1959 [TVL]. REMARKS: The female from Trinidad is too badly damaged to permit study of the abdominal characters. It has about 12 setae on each of the seventh sternites. This species is named in honor of Mr. Pedro Galindo of the Gorgas Memo- rial Laboratory, in recognition of his noteworthy contributions to our knowl- edge of the ectoparasites of Panama, and for his invaluable assistance in fa- cilitating the field work in Panama. Strebla christinae Wenzel, new species. Figures 44A, 124E, 131A. S. christinae n. sp. is the only species of those with detached fronto- clypeal plates that lacks a second or anterior pigmented mesonotal suture. From related species that have numerous strong metatibial setae and multi- faceted eyes — diphyllae n.sp., vespertilionis (Fabricius), mirabilis (Water- house) — it may be recognized by the relatively short spine-like setae and shape of the postvertex (fig. 124E) and, in the female, by the conspicuous anteriorly produced lobe of the ventral arc. DESCRIPTION: Head. — Detached frontoclypeal plates irregular, about as wide as long. Anterior plate of laterovertices with seven setae; postocular sclerite distinct. Dorsal surface of postgenae not emarginate laterally, its posterior portion with 10-11 shorter but conspicuous setae, one very long posterior seta and lateral to it one that is about half as long. Postvertex and occiput as illustrated. Eyes multi-faceted, project- ing, about seven facets visible from above. Ventral ante-ctenidial area a little wider than long (21.5:19.5). Thorax (fig. 131 A). — Prescutum lacking a second pigmented suture; setae rather uniformly distributed and extending anteriorly nearly to anterior margin along median suture, and to epaulets along sides, the epaulets consisting of three to five setae which merge into the median prescutal setae; arc of longer setae poorly defined, only the two outer setae distinctive. Scutal setae of antescutellar row longer than discals and becom- ing longer toward sides; discal setae arranged in three transverse rows. Wings. — All veins except sixth longitudinal setose to base. A longer seta at apex of costa but not a macroseta. Rs nearly half again as long as distance from fork to r-m. Legs. — Dorsal edge of metatibiae with a double row of 10-12 macrosetae, of varying length. First segment of hind tarsi with a transverse row of three coarse plantar bristles a little be- yond middle, and five or six shorter, finer ones basally; plantars of segments 2-4 dis- tinctive but not as coarse as those of first segment. Abdomen. — Inner dorsal margin of lateral lobes of tergum I+II with from three to five setae. Female: Tergum VII with a pair of macrosetae and posterior and median to them a pair of setae about one-fourth as long. Supra-anal plate with four apical macro- setae and on each side four conspicuous setae, two dorso-lateral, one lateral, and one ventral. Seventh sternites with about nine setae, three of them macrosetae. Ventral arc produced anteriorly (fig. 44 A) as a conspicuous microsetose lobe; with four short setae, inner pair twice as long as outer. Male : Sternum V short, with no more than two WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 607 transverse rows of discal setae throughout width. Sterna VI and VII+VIII forming a wide ring. Hypopygial setae as in diaemi n.sp. Gonapophyses curved basally, nearly straight apically, apices mucronate below; ventral macroseta inserted at about basal third, short, extending only a little more than half the distance to apex; accessory seta minute ; with scattered microsetae along sides on apical third. Measurements: BL TL WL ww Male 2.75-2.87 0.93 1.65-1.81 0.95-0.99 Female 2.97-3.00 0.93-0.96 1.81-1.92 0.93-1.04 TYPE MATERIAL: Holotype male, allotype female (slides) and 7 paratype females from Phylloderma, stenops (host no. 11986) , Armila (San Bias) , 26 A B Fig. 131. Thorax, dorsal view. A, Strebla christinae, new species, male paratype. B, S. machadoi, new species, holotype. March 1963, C. 0. Handley, Jr. and F. M. Greenwell. 1 male paratype, same data, but 27 February 1963. Holotype and allotype in the collection of Chi- cago Natural History Museum. Paratypes in the collections of Chicago Natural History Museum; the United States National Museum; and the Environmental Health Branch, U. S. Army, at Corozal (Canal Zone). This species is named for Christina Johnson Fowler in grateful appre- ciation of her invaluable assistance in producing this volume. Strebla machadoi Wenzel, new species. Figures 124D, 131B. S. machadoi n. sp. is easily separated from all other species in that it is the only one with detached frontoclypeal plates that has non-faceted eyes. The subacuminate anterior margin of the frontoclypeus is also distinctive. DESCRIPTION (Female) : Head. — Detached frontoclypeal plates small, about as wide as long. Anterior plate of laterovertices with six setae; postocular sclerite distinct. Dorsal sclerite of postgenae posteriorly with five or six short bristles, a long, conspicuous one on postero-median angle and lateral to this a seta of intermediate length; lateral margin emarginate for reception of ctenidium. Postvertex and occiput as illustrated 608 ECTOPARASITES OF PANAMA (fig. 124D) , but with the inner three or four setae on each side more distinct spinelets than shown. Ventral ante-ctenidial area wider than long (21:17). Eyes consisting of a single elongate hyaline lens. Thorax. — Prescutum with a pigmented second or anterior transverse suture; prescutum sparsely setose, a bare area along median line on anterior division, except for two long setae and a short seta on each side of line; epaulets con- sisting of five to six stout setae ; arc of longer setae distinct ; two transverse rows of setae laterally between transverse and anterior suture; antescutellar bristles very long at sides, becoming gradually and distinctly longer medially, the median bristles distinctly longer than the discal scutal setae arranged in two or three irregular transverse rows. Fig. 132. Thorax, dorsal view, Strebla vesper tilionis (Fabricius), from Desmodus rotundus murinus (no. 5236), Red Tank (Canal Zone). Wings. — All veins except sixth longitudinal setose to base. Apex of costa without a macroseta. Legs. — Pro- and mesotibiae without distinctive characters. Metatibiae with two conspicuous macrosetae near apex, preceded by a double row of eight to ten setae that are shorter than the macrosetae but at least twice as long as the other tibial setae. First segment of hind tarsus with two rows of heavy and an inner row of weaker but still prominent plantar setae, four across apical margin; segments 3 and 4 with distinct plantar bristles, but none as heavy or as long as those of first segment, those of the fourth segment relatively weak. Abdomen. — Anterior face of tergum I+II with four short setae; inner dorsal margin with two fine setae. Female: Tergum VII with a pair of macrosetae and posterior to them a pair of more closely placed, shorter setae that are less than one-half as long as macrosetae. Supra-anal plate with four long apical setae and a pair of short setae on each side near base. Seventh sternites with eight setae, two of these macrosetae. Ventral arc produced anteriorly as a broad, short, rounded microsetose lobe about twice WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 609 as wide as long, with two long and two short setae on posterior edge. Male: Unknown. Measurements: BL TL WL ww Female 2.39 0.77 1.65 0.85 TYPE MATERIAL: Holotype female from Micronycteris minuta (host Machado- Allison no. 495), La Guanote, Caripe (Monagas), VENEZUELA, J. Ojasti. In the collection of the Faculdad de Ciencias, Universidad Central de Venezuela, Caracas. A paratype, sex undetermined (abdomen missing) , same data as the holotype, in the collection of Chicago Natural History Mu- seum. REMARKS : This species is named for Dr. Carlos Machado-Allison, of the Departamento de Parasitologia y Microbiologia, Faculdad de Ciencias, Uni- versidad Central de Venezuela, in recognition of his valuable contributions to the study of ectoparasites. Strebla vespertilionis (Fabricius). Figures 123B, 125B, 132. Hippobosca vespertilionis Fabricius, 1805, Syst. Antliat., p. 339.6 — "America meridionalis vespertilione" (Type: See discussion below.) Strebla vespertilionis Wiedemann, 1824, Analect. Ent., p. 19, fig. 7; 1830, Aus- sereurop. Zweifl. Ins., 2: 612, pi. xb, fig. 13a, b. Macquart, 1835, Hist. Nat. Ins. Dipt., 2: 637, pi. 24, fig. 16. Walker, 1849, List. Dipt. Spec. Brit. Mus., 4: 1146 (cit., part.). Osten-Sacken, 1878, Cat. N. Am. Dipt., Smiths. Misc. Coll., no. 270, p. 214 (cit., part.) Strebla wiedemannii (new name for vespertilionis Fabricius [not Nitsch, 1803], in error), Kolenati, 1856, Parasiten d. Chiropt., p. 46. Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 32, 43 (status); 1907, Ent. News, 18: 103. Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 31 (cit. only, not fig.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 654 (cit.). Euctenodes mirabilis (not Waterhouse, 1879), Kessel, 1924, Parasitology, 16: 411 (part., from Desmodus sp.), figs. 2, 3; 1925, Jour. N. Y. Ent. Soc., 33: 30 (part., from Desmodus sp.), pi. 3, fig. 21, pi. 4, figs. 22, 23. Stiles and Nolan, 1938, Bull. Nat. Inst. Hlth., Wash., no. 155, pp. 654, 732 (part., from Desmodus sp.). Jobling, 1949, Parasitology, 39: 318-19, 328 (part., from Desmodus rotundus), (? figs. 3 D, E.). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 188 (? part., from Desmodus r. murinus). Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., no. 122, p. 269. Kolenati (1856) erroneously regarded Strebla vespertilionis as a homo- nym, changed the name to Strebla iviedemannii, and noted : "Das Original exemplar sich in konigl. Museum zu Kopenhagen." In 1863 (p. 18) , however, Kolenati stated : "Das Original exemplar aus der zu Kopenhagen existierenden alten Toiider Liend-Schestedt'schen Sammlung, welches Wiedemann zur Beschreibung und Abbildung iiberlassen wurde, ist entweder bei die Zeichung verloren gegangen, oder nicht mehr zuruckgegeben warden, weil an dem Platze dieses Originalstiickes, welcher vorhalten blieb als die Sammlung vor 30 Jahren umgestellt wurde, kein nadelstich ist [italics ours]. Diese Mitthieilung machte mir Prof. Steenstrup, am 10 Nov. 1857." Dr. Sven G. Larson (pers. comm.) of the Copenhagen Museum recently confirmed the absence of the type of vespertilionis and added : "If there is a possibility that Wiedemann, for some reason or another, did not return the animals to the owners (Sehested & T0nder Lund) , the type may possibly be in Vienna, where most of Wiedemann's animals are found, as far as I know." We then asked Prof. Dr. Max Beier of the Vienna Museum to see if there were specimens in Wiedemann's material that could possibly be Fabricius' 610 ECTOPARASITES OF PANAMA type. To our surprise, Dr. Beier answered : ". . . iibersende ich Ihnen mit gleicher Post unser einziger Exemplar von Strebla vespertilionis F. Dieses Stuck stammt zweifellos aus der Sammlung Wiedemann, da es die griine Etikett am Klebeblattchen tragt. . . . Wahrscheinlich handelt es sich also um die verschollene Type von Fabricius." The specimen is of a species that parasitizes Desmodus rotundus. It carries four labels : one reading "Strebla vespertilionis, Pernambuco, Bgt. ;" a second, reading "Schiner 1869 ;" a third "vespertilionis det. Speiser" ( !) ; and the fourth, reading "Strebla vespertilionis F." The specimen is glued to a celluloid card, which bears a green, quadrangular marker at the base. The "Bgt." on the first label is puzzling. It is possible that it is an abbrevi- ation of Bigot. The word "Strebla" is hand-printed and resembles the sample of Bigot's hand-printing given by Horn and Kahle (Ent. Beihefte, 2, pi. 11, fig. 1). The specimen seems to have come to the Vienna Museum through the Schiner collection in 1869. This, too, is puzzling, since, accord- ing to Horn and Kahle (ibidem, 4, p. 302), the Wiedemann Diptera went to Vienna via W. von Winthem, and the same authorities (p. 305) state that the von Winthem Diptera went to that museum in 1852. We are not convinced that this specimen is the type of vespertilionis, but in view of Dr. Beier's experience, we accept it as an authentic Wiedemann specimen. If it is not the type, however, it is quite possible that it is the specimen upon which Wiedemann based his 1830 description and illustration (see p. 591). It is certainly congeneric with the specimen figured, and it matches in the setation of the hind tibiae, taking into account the poor quality of the figure. Since we cannot demonstrate that this specimen is Fabricius' type, we desig- nate it as a neotype, so as to fix the identity of the type-species of the type- genus of the family. As pointed out by Speiser, Strebla iviedemannii Kolenati, 1863, is not the same as wiedemannii Kolenati, 1856 (& 1857). Speiser considered it to be identical with Trichobius parasiticus and T. dugesii Townsend. Actually, it probably was a mixture of species and the specimen illustrated (Kolenati, 1863, pi. 16, fig. 36) may be an undescribed species of Trichobius that occurs in the West Indies, Mexico, and Guatemala. Strebla vespertilionis of von Roder (1896, p. 26) from Vampyrops lineatus was probably a species of Paratrichobius. Strebla avium Macquart (1854, p. 308) could have been a hippoboscid. We have not seen the original description. To our knowledge, no one has re-examined the type. It was supposed to be in Macquart's collection at Marseilles. We do not know if it exists or not. Dr. T. C. Maa has examined the type of Strebla mexicana Rondani ( 1878, p. 168) . According to him (pers. commr.) , it is heavily coated with dirt, but on the basis of a superficial examination, he believes it to be "a Euctenodes" 1= Strebla'}. Only a few species of Strebla are known to us from Mexico. They include S. vespertilionis, carolliae n. sp., diphyllae n.sp. and hertigi n. sp. We hope that in the future there will be an opportunity to clean and make a slide preparation of the type of mexicana so as to establish its iden- tity. It is in the museum at the University of Florence (Italy) . S. vespertilionis is similar to machadoi n. sp. and christinae n. sp. in WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 611 possessing two rows of six or seven conspicuously longer setae each, on the upper edge of the metatibiae (fig. 125B) . It differs conspicuously from both in mesonotal chaetotaxy (fig. 132) and from machadoi in having multi- faceted eyes rather than a single hyaline lens, and from christinae in having a second or anterior pigmented transverse suture, as well as the other char- acters given in the key. Actually, vespertilionis appears to be more closely related to diphyllae, but this species has only a single dorsal row of longer metatibial setae and it has distinctly curved gonapophyses. The only other species known to us that have nearly straight gonapophyses are S. hertigi n. sp., consocius n. sp., and diaemi n. sp. In the two last-named species, the ventral macroseta is inserted distal to middle on the gonapophysis, while in hertigi and vespertilionis, it is inserted basally. S. hertigi, however, be- longs to the group of species with entire f rontoclypeus ; its eyes consist of a row of four facets ; and its gonapophyses, apically, are upwardly sinuate rather than downwardly curved. DESCRIPTION: Head. — Frontoclypeus with apical detached plates that are distinctly longer than broad. Anterior plate of laterovertices with eight setae; postocular sclerite distinct. Dorsal sclerite of postgenae not emarginate laterally; posterior portion with 11-13 short setae and near posterior margin, two long setae. Postvertex and occiput as figured (fig. 132B), but with most of the long occipital setae and spinelets of postvertex more slender than shown. Eyes multi-faceted. Ventral ante-ctenidial area distinctly wider than long (30:22). Thorax (fig. 132). — Prescutum with a pigmented second or anterior transverse suture; epaulets consisting of five or six spinelets; arc of longer setae distinct; discal setae not reaching epaulets ; three irregular transverse rows of setae between transverse mesonotal suture and anterior suture. Antescutellar setae much longer than other scutal setae, those near sides longer than those at middle. Wings. — All veins except sixth setose to base. Seta at apex of costa not a macroseta. Rs nearly twice as long (22.5:13) as distance from fork to crossvein r-m. Legs. — Protibiae dorsally with three heavy setae along inner edge, near apex, preceded by four slender, shorter setae; outer edge with one or two strong setae near apex. Mesotibiae apically with about five strong dorsal setae on outer edge ; inner edge with two strong setae preceded by a row of weaker ones. Metatibiae (fig. 125B) with about 14 conspicuous setae in a double row, the apical two nearly twice as long as the others. Ventral surface of hind tarsus with a median, longi- tudinal row of five very heavy plantar bristles and lateral to these, basally, a row of five very short plantars along inner edge ; plantar bristles of succeeding segments minute. Abdomen. — Inner dorsal margins of lateral lobes of tergum I+II with three to four setae. Female : Tergum VII with variable chaetotaxy, usually with three pairs of setae, an anterior pair of macrosetae and posterior to these two pairs of shorter strong setae. Supra-anal plate with four apical macrosetae and on each side near base a pair of setae that are about one-third as long. Seventh sternites with about 17 setae, seven or eight of them very long, about five much longer than the others. Ventral arc with both anterior and posterior margins slightly indented, not produced; with four setae, two longer, two minute. Male: Hypopygium as in S. diphyllae n.sp. Gonapophyses gradually tapered, with both dorsal and ventral margins curved to the knobbed apices ; sides, especially on apical third, with numerous microsetae ; ventral macroseta inserted at about basal third, extending about three-fourths the distance to apex; accessory seta minute. Measurements: BL TL WL ww Male 2.97-3.03 0.93-1.00 1.70-1.91 0.82-0.89 Female 2.94-3.41 1.00-1.04 2.03-2.17 0.89-0.96 PANAMANIAN MATERIAL EXAMINED: 50 lots, 288 specimens, from more than 92 bats. From Desmodus r. murinus: 1, Paraiso (Canal Zone) , 17 No- vember 1960; 9, Red Tank (Canal Zone), 27 January 1960; 45 (7 bats), 612 ECTOPARASITES OF PANAMA Almirante (Bocas del Toro) , 23 to 31 January 1960 ; 1, Changuinola (Bocas del Toro), 27 February 1960; 72 (40 bats), Casa Tilley near Cerro Punta (Chiriqui), 23 January and 6 February 1960; 1, same locality, 11 March 1962; 5, Chilibrillo Caves (Panama), 2 May 1957 [USNM] ; 2, same locality, L. H. Dunn no. 878 [MCZ] ; 4, San Pedro Cave, Taboga Island (Panama), L. H. Dunn [MCZ] ; 12, near La Chorrera (Panama), 18 April 1959, H. Trapido; 112 (10 lots, 12 bats), Guanico (Los Santos), 26 January to 25 Fig. 133. Thorax, dorsal view, Strebla diphyllae, new species, male paratype, same data as holotype. February 1962; 22 (3 bats), Rio Tuira (Darien), 24 to 27 February 1958, P. Galindo [GML] ; 3, Rio Mono Camp (Darien), 24 July 1963, W. Lowe [GML]. From Artibeus jamaicensis: 3 (9 bats), Almirante (Bocas del Toro), 26 and 28 January 1960. From Vampyrodes major: 2, Almirante (Bocas del Toro), 22 January 1960. From Lasiurus egregius: 1, Armila (San Bias), 23 February 1963. OTHER MATERIAL EXAMINED: 1162 flies in 113 lots, from more than 150 bats, mostly from Desmodus rotundus, from 69 localities in MEXICO, GUATE- MALA, HONDURAS, EL SALVADOR, COLOMBIA, ECUADOR, PERU, VENEZUELA, TRINIDAD, and SURINAM. REMARKS : 96% of all the lots and 97% of all the specimens of S. vesper- tilionis examined were from Desmodus rotundus. Excluding one collection WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 613 from mixed hosts that included Desmodus, only 7 lots (including those from Panama) are from bats other than Desmodus. These bats include Lasiurus egregius (a genus not known to harbor Streblidae), Artibeus jamaicensis, Vampyrodes major, Saccopteryx bilineata, Centurio senex, Carollia per- spicillata and Phyllostomus discolor. From the history and nature of the collections, we are convinced that some of these represent contaminations or errors of association. However, several obviously represent temporary or transitory associations. Kessel (1924, p. 411) stated that her redescription of E. "mirabilis" was based on a specimen from Hemiderma [= Carollia] perspicillatum, but her figures are obviously of Strebla vespertilionis, which occurs on Desmodus rotundus. Jobling has informed us (pers. comm.) that his figures (1949, figs. 3D, E) of "mirabilis" were based on a specimen from Desmodus r. rotundus from Diego Martin Cave, Trinidad. One would assume, then, that this specimen was vespertilionis. However, the short, detached fronto- clypeal plates are much more like those of mirabilis than those of vesper- tilionis, and it will be necessary to examine his slide to determine the identity of his specimen. Strebla diphyllae Wenzel, new species. Figures 124C, 133. Euctenodes mirabilis (not Waterhouse, 1879), Bequaert, 1933, Publ. Carnegie Inst. Wash., no. 431, p. 571. ?Hoffmann, 1953, Mem. Congr. Cient. Hex., 7: 184, 188 (part., from Diphylla). Very closely related to S. vespertilionis (Fab.), but differing from it in having shorter frontoclypeal plates; more densely setose prescutum; tergum VII of female with four rather than six setae; and male gon- apophyses distinctly bent rather than nearly straight. DESCRIPTION : Head.- — Short. Frontoclypeus with apical detached plates, these slightly wider than long. Anterior plate of laterovertices with eight setae; postocular sclerite distinct. Postgenae with lateral margins not marginate, posteriorly with eight or nine short bristles and near margin two long conspicuous bristles, the inner one nearly as long as the setae of postvertex, the outer one a little shorter ; lateral to them a shorter bristle. Postvertex and occiput as illustrated (fig. 124C) . Eyes multi-faceted with seven or eight facets visible from above. Ventral ante-ctenidial area wider than long (32.5:21). Thorax. — Prescutum with anterior transverse suture; epaulets consisting of five to six stout spinelets; setose area of disk extending anteriorly along mid-line to the double pair of short setae behind the anterior margin, and along sides nearly to epaulets ; arc of seven longer setae distinct; three rows of setae laterally between transverse and anterior sutures. Scutum with about three transverse irregular rows of setae; antescutellar setae long. Wings. — All veins except sixth with setae to base; costa without an apical macroseta. Rs about half again as long (30:21) as distance from fork to r-m. Legs. — Protibiae with about three heavy conspicuous setae, preceded by several long fine setae on inner margin; outer margin with two or three heavier setae. Mesotibiae with eight or nine conspicuously coarser setae. Metatibiae with a dorsal row of nine or ten long setae, the apical one about as long as first segment of hind tarsus, the others progres- sively shorter anteriorly, the basal ones about twice as long as short setae on lateral face. Ventral surface of first segment of hind tarsus with three longitudinal rows of heavily pigmented conspicuous plantar bristles, those along inner edge smaller; plantar bristles of following segments very small but stout. Abdomen. — Inner dorsal margins of tergum I+II with about four setae. Female: Tergum VII with two long macrosetae and a pair of shorter setae, posterior and medial to them, only a little more than half as long. Supra-anal plate with a median pair of 614 ECTOPARASITES OF PANAMA strong setae along basal margin in addition to the four apical macrosetae. Seventh sternites with 20-21 setae. Ventral arc slightly thickened at middle, with two longer and two minute setae. Male : Hypopygium dorsally with a lateral macroseta and medial to it a very short seta on sternum VII+VIII, and on tergum IX a transverse row of six macrosetae, the medial pair longest. Gonapophyses strongly bent a little beyond Fig. 134. Thorax, dorsal view, Strebla mirabilis (Waterhouse), male from Phyllostomus h. panamensis (no. 4138), Natural Bridge, Madden Dam (Canal Zone), ep = epaulet; a = prescutal arc of setae. middle; macrosetae inserted near middle, not reaching knobbed apices; sides with scattered microsetae on a little more than apical third. Measurements: BL TL WL ww Male 2.11-3.20 1.02-1.04 1.81-1.92 0.91-0.93 Female 3.20-3.53 1.00-1.07 1.84-2.47 1.10-1.21 TYPE MATERIAL: GUATEMALA (CNHM Guatemala Zoological Expedi- tions, 1936, 1948, 1951-52). — Holotype male and allotype female (slides) from Diphylla ecaudata centralis (CNHM cat. nos. 73191-94, 73345-46), San Lorenzo, 4 miles northeast of Volcan de Jumay ( Jalapa) , 5700-5750 feet elevation, 15 February 1952, R. D. Mitchell and Luis de la Torre. In the collection of Chicago Natural History Museum. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 615 Paratypes. — From Diphylla e. centralis: 13 (7 bats), same data as the holotype ; 3 (5 bats) , Finca El Progresso (Santa Rosa) , 4500 feet elevation, 3 August 1951, Luis de la Torre; 4, Finca Recreo, Municipio Yepocapa (Chimaltenango), elevation 4400 feet, 11 May 1948, R. L. Wenzel, L. de la Torre, and R. D. Mitchell ; 15 (5 bats) , same locality, 25 October 1948, R. D. Mitchell and L. de la Torre. From Trachops cirrhosus coffini: 1, Finca Recreo (Chimaltenango), 2 May 1948, Wenzel, Mitchell and de la Torre. From Desmodus rotundus murinus: 5 (5 bats), Finca El Progreso (Santa Rosa), 3 August 1951, L. de la Torre; 2, Finca Olas de Moca (Solola), 2700 feet elevation, 3 March 1934, F. J. W. Schmidt. Non-Guatemalan Paratypes. MEXICO. — From Diphylla e. centralis: 2, Yocat (Yucatan), R. T. Hatt [MCZ, det. Bequaert, 1933, as E. mirabilis~] ; 18, Xilitla (San Luis Potosi), C. R. Shaw [MCZ] ; 4, 2 km. north of Felipe Carillo Puerto (Quintana Roo), 16 August 1962, J. Knox Jones [KU]. Paratypes to be deposited in the collections of Chicago Natural History Museum; the United States National Museum; Snow Entomological Mu- seum, the University of Kansas; the Museum of Comparative Zoology at Harvard University; and the Environmental Health Branch, USAFSC at Corozal (Canal Zone). REMARKS: The Guatemalan records from Trachops (Finca Recreo) and Desmodus (Finca El Progresso) almost certainly represent strays from other bats collected at the same time and placed in the same bags as specimens of Diphylla e. centralis. This does not seem to be true for the specimens from Desmodus collected at Finca Olas de Moca. Six specimens of S. vespertilionis n. sp. were collected together with three S. diphyllae, from the same host specimen. As nearly as can be determined from the field records, no specimens of Diphylla ecaudata were collected at that locality. Although Diphylla ecaudata was collected in Panama, no specimens of Strebla diphyllae were obtained. In Venezuela, a new species (not de- scribed here) related to mirabilis, was collected from this host by C. 0. Handley, Jr. In view of the fact that monotypic New World hosts do not, in any instance known to us, have different streblids in different parts of their range, the suggestion of Burt and Stirton (1961, p. 37) that Diphylla ecaudata is monotypic should be questioned. Strebla mirabilis (Waterhouse), new combination. Figures 123A, 125E, 134, 135A. Euctenodes mirabilis Waterhouse, 1879, Trans. Roy. Ent. Soc. Lond., 1879: 310, pi. 10, figs. 1, 2. Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Veter., Nictheroy, 5: 32 (cit.). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Ex., Fis. y Nat., 3: 418 (part., from Phyllostomus h. panamensis). Cooper, 1941 (part.), Yearb. Amer. Phil. Soc., 1941: 126 (chromosome no.). Closely related to S. kohlsi but distinct in having a shorter head and shorter frontoclypeal plates (see key) . It is primarily a parasite of Phyllo- stomus hastatus, though it may also occur on Trachops (see below). S. kohlsi is known to us from Tonatia silvicola. Phyllostomus, Trachops, and Tonatia are related genera of the subfamily Phyllostominae. 616 ECTOPARASITES OF PANAMA DESCRIPTION : Head. — Frontoclypeus with apical detached plates, these approximately as long as wide, though inner margin is appreciably longer than apical. Anterior di- vision of laterovertices with eight setae; postocular sclerite distinct. Dorsal sclerite of postgenae not emarginate laterally, its posterior portion with ten fine and mostly very short setae and, along posterior margin, two conspicuous long setae. Postvertex and occiput as illustrated (fig. 123A). Eyes convex, multi-faceted, about eight facets visible from above. Ventral ante-ctenidial area wider than long (59:41). Thorax (fig. 134). — Prescutum with second anterior pigmented suture; epaulets with four to five setae; discal setose area not extending to epaulets; arc of long setae very distinct. Scutal setae of antescutellar row longer toward sides; three transverse rows of setae present between them and the transverse mesonotal suture. Wings. — Rs less than twice as long (22:13) as distance from fork to crossvein r-m. Seta at apex of costa longer than others preceding it but not a conspicuous macroseta. Fifth longi- tudinal vein bare for a short distance before base. Legs. — Protibiae with a double row of longer setae along dorsal edge, the basal ones small, the apical three on inner edge and two on outer edge very stout; apices with two strong curved ventral setae. Setae of mesotibiae very similar to those of protibiae. Metatibiae with two conspicuous macro- setae on apical third, these preceded by a distinctly shorter and finer seta and then a row of five or six shorter setae, all of them longer than the other tibial setae. Ventral surface of first segment of hind tarsi with two rows of heavy, short plantar bristles along outer half and an irregular row or two of shorter bristles along inner edge; fol- lowing segments with minute, very fine plantar bristles that can hardly be detected ex- cept in slide preparations. Abdomen. — Inner dorsal margins of lateral lobes of tergum I+II with four to six setae on each side. Sternum II with setose median triangular area, the apical marginal setae conspicuously longer than the discals, a conspicuously long macroseta on each side of middle, separated by about four shorter setae. Dorsal connexival setae along sides longer than ventral setae, those along lateral margins shortest. Female: Tergum VII roughly tear-drop shaped; distally with a pair of very long macrosetae and medial and posterior to these, a pair of setae that are only half as long. Supra-anal plate with four apical macrosetae and a pair of shorter discal setae. Seventh sternites with 11-12 setae, two to four of them very long macrosetae. Ventral arc with two long and two minute setae; produced anteriorly as a small poorly sclerotized lobe. Male: Sternum V well developed ; apical margin with about eight conspicuous macrosetae alternating with shorter setae, all longer than discals, of which there are two transverse rows at middle and three toward sides. Hypopygium with a very long macrosetae on each side and medial to this a short bristle on sternum VII+VIII; dorsum of tergum IX with three pairs of macrosetae. Gonapophyses bent apically, with scattered microsetae along middle on apical half; apices knobbed; ventral macroseta not reaching apex. Measurements: BL TL WL ww Male 2.32-2.72 0.86-0.93 1.54-1.65 0.81-0.84 Female 2.58-3.08 0.92-0.97 1.73-1.94 0.91-0.95 PANAMANIAN MATERIAL EXAMINED. — From Phyllostomus Hcistatus pan- amensis: 140 (17 bats) , hollow tree, Chepo Road (Panama) , 8 October 1959 ; 112 (20 bats), Chilibrillo Caves, 17 July 1959; 4 (2 bats), same locality, 18 August 1959; 322 (13 bats), 28 October 1959; 3, same locality, 2 May 1957 [USNM] ; 11, same locality, K. W. Cooper [MCZ] ; 2, same locality from pre- served museum specimens collected 4 September 1934 by L. D. Lamm; 147, same locality, L. H. Dunn, no. 436 [MCZ] ; 1, Tapia (Panama), 25 July 1923, J. P. Chapin [MCZ]; 1, "Panama" [MCZ]; 12, Farfan (Canal Zone), 19 December, T. Hallinan [AMNH] ; 168 (20 bats), Natural Bridge, Madden Dam (Canal Zone), 31 August, 28 September 1959; 77 (5 bats), hollow Ficus tree, Fort Kobbe (Canal Zone), 9 October 1959; 4 (3 bats), Fort Sherman (Canal Zone), 30 July 1959; 149 (19 bats), 4 and 6 April 1960; 14, France WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 617 Air Force Base (Canal Zone), 8 October 1959; 7, hollow tree, 1 mile from Gatuncillo (Canal Zone), 28 July 1960; 3, Almirante (Bocas del Toro), 29 January 1960. From Phyllostomus sp. : 11, Camp Pina (Canal Zone), 26 October 1960. From Carollia p. azteca: 25 (3 bats), Natural Bridge, Mad- den Dam (Canal Zone) , 31 August 1959. Without host: 1, Chilibrillo Caves (Panama), 27 August 1959; 30, same locality [MCZ] ; 2, same locality, 1 January 1916, T. Hallinan [AMNH] ; 31, Barro Colorado Island (Canal Zone), 2 December 1956, K-18 [USNM]. OTHER MATERIAL EXAMINED: MEXICO. — Without host: 1, "either from Campeche, Mexico or Yucatan. Original vial had label no. 15 plus included." COLOMBIA. — From Phyllostomus hastatus: 6 (3 bats), "Las Campanas," Coloso (Bolivar) , 23 May 1949, P. Hershkovitz, CNHM Colombian Zoological Expedition (1949-51) ; 1, Pitalito (Huila) , 1350 meters elevation, 29 Novem- ber 1951. From Tonatia sp. : 1, Rio Guayapa, La Macarena (Meta) , 10 April 1957, Kjell von Sneidern, CNHM Colombian Zoological Expedition, 1957. PERU. — From Phyllostomus hastatus: 2, Vitoc Valley, Province Parma (Junin), 18 September 1940, F. Woytkowski. From Phyllostomus sp. : 1, Santa Rita, Iquitos (Loreto) , 4 September 1956, C. Kalinowski, CNHM Peru Zoological Expedition, 1956. BOLIVIA. — From "Fledermaus," 2, Mina Cris- talmaya (Cochabamba), October 1949, Luis Pena. REMARKS: Both Mr. B. Jobling and Prof. Oskar Theodor have kindly examined the type of mirabilis and compared it with specimens of several related species which we sent them. In addition, Prof. Theodor referred to our manuscript keys and figures. He supplied us with notes and sketches which leave no doubt as to the identity of mirabilis. Probably most records that have been published for mirabilis from Phyllostomus hastatus are of this species. However, one cannot be certain without examining the speci- mens, since at least two other species also occur on this host. In addition to the verified records given above, mirabilis has been reported from P. hastatus by Kessel (1924, 1925), Stiles and Nolan (1931), Pessoa and Guimaraes (1940), Schuurmans-Stekhoven Jr. (1941), Jobling (1949b), and Goodwin andGreenhall (1961). We have additional specimens from hosts such as Tonatia bidens in Brazil (Urucum de Curumba) and Vampyrops vittatus (same locality and expedi- tion) , but we are not certain of their identity. We have recently segregated specimens of a very closely related species, taken from Diphylla ecaudata in Venezuela, which we were unable to describe and illustrate in time to include in this paper. Strebla ( ?) mirabilis from Trachops cirrhosus. Figure 135B. We have specimens that appear to be mirabilis, taken from Trachops c. cirrhosus in Panama and from T. c. coffini in Guatemala. The gonapophyses of these appear to be slightly more angulately bent (fig. 135B) than in mirabilis (fig. 135A) from Phyllostomus. The number of setae on the seventh sternites (each) ranged from 13-16 in specimens from P. h. pana- mensis and 11-13 in specimens from T. c. coffini. It should be noted that to the north in Costa Rica and Nicaragua, Phyllostomus h. panamensis does not appear to carry Strebla mirabilis, but 618 ECTOPARASITES OF PANAMA only S. hertigi (see "Host-parasite Relationships"). One explanation for the absence of mirabilis there, is that its range, like that of many ectopara- sites, may not be as great as that of the host. However, it does appear to oc- cur on Trachops much farther north, in Guatemala. Further, all of the Panamanian specimens of mirabilis from Phyllostomus were taken below ±500 feet elevation, though P. h. panamensis was taken at elevations of up to 2000 feet. The specimens from Trachops c. cirrhosus in Panama were taken from between sea level and 1700-3200 feet, while those from T. c. coffini in Guatemala were taken at 4400 feet. Its altitudinal distribution plus the fact that mirabilis is absent from the usual host in Costa Rica and Nicaragua, and yet is present on Trachops to the north in Guatemala suggest that the population on Trachops is a separate, cryptic species. A Fig. 135. Strebla mirabilis (Waterhouse), left male gonapophysis. A, of specimen from Phyllostomus h. panamensis (no. 4138), Natural Bridge, Madden Dam (Canal Zone). B, of specimen from Trachops cirrhosus coffini (CNHM nos. 65181-90), Finca Recreo, Municipio Yepocapa (Chimaltenango), GUATEMALA. MATERIAL EXAMINED : PANAMA. — From Trachops c. cirrhosus: 2 (2 bats) , Fort Sherman, 22 October and 23 November 1959 ; 9 (5 bats) , culvert, Chepo Road (Panama), 12 October 1959 and 24 May 1960; 29 (24 bats), Cerro Hoya (Los Santos), 1700-3200 feet elevation, 10 to 25 February 1962; 1, Armila (San Bias), 16 March 1963, C. O. Handley, Jr. and F. M. Greenwell. GUATEMALA. — From Trachops cirrhosis coffini: 30 (5 bats), Finca Recreo, Municipio Yepocapa (Chimaltenango), 4400 feet elevation, 11 May 1948, R. L. Wenzel, R. D. Mitchell, and L. de la Torre, CNHM Zoological Expedition (1948). Strebla kohlsi Wenzel, new species. Figure 123C. Related to mirabilis (Waterhouse) and distinguished from it by the elongated head, with ventral ante-ctenidial area hardly wider than long (24 :22 as opposed to 59 :41 in mirabilis) , its slightly smaller size, and more elongated f rontoclypeal plates. DESCRIPTION : Head. — Elongated. Detached f rontoclypeal plates present, their inner edge longer than apical margin, though basal margin is almost as wide as inner edge is long. Anterior plate of laterovertices with six setae; postocular sclerite distinct. Dorsal surface of postgenae not emarginate laterally; posteriorly with about six short setae and a long conspicuous seta and a shorter one near posterior margin. Postvertex WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 619 and occiput as illustrated, but outer two setae somewhat finer than shown. Eyes multi- faceted and projecting, about seven or eight facets visible from above. Ante-ctenidial area of underside only slightly wider than long (24:22). Thorax. — Second pigmented suture of prescutum present but not very distinct; epaulets consisting of four setae; discal setose area not extending anteriorly beyond anterior end of longitudinal membranous fissures; arcuate row of long setae well de- fined; three transverse rows of setae laterally between sutures. Antescutellar setae more or less uniform in length. Wings. — Seta at apex of costa distinctly longer than those preceding it, but not a conspicuous macroseta. Rs half again as long (23:15) as distance between fork and crossvein r-m. Fifth longitudinal vein with a short, bare area at base, this less than half as long as bare area on sixth vein. Legs. — Protibiae dorsally with four heavy macrosetae on inner edge, and two or three on outer ; mesotibiae with one or two on inner edge and four or five on outer, preceded by more slender long bristles. Metatibiae with two conspicuous macrosetae on apical third. Ventral surface of first segment of hind tarsi with two longitudinal rows of shorter plantar bristles near inner edge and outwardly at base a longer more conspicuous one, and across apex three additional coarser ones; succeeding segments with distinct plantar bristles but without very coarse longer ones. Abdomen. — Anterior face of tergum I+II with two short setae; inner dorsal margin with about four fine setae on each side. Female: Tergum VII with a pair of macro- setae and posterior to these a pair of shorter ones that are less than half as long as anterior pair. Supra-anal plate with four apical macrosetae and, basally, a pair of shorter setae that are slightly shorter than the posterior pair on tergum VII. Seventh sternites each with about 14 setae, one a macroseta. Ventral arc slightly longer at middle, not produced as a lobe, with four setae. Male : Hypopygium with a dorso-lateral macroseta on each side; tergum IX with two pairs of macrosetae dorsally and another macroseta on each side. Measurements: BL TL WL ww Male 2.20-2.47 0.88-0.93 1.59-1.81 0.77-0.82 Female 2.28-2.58 0.80 1.59-1.76 0.74-0.77 TYPE MATERIAL: PANAMA. — Holotype male and allotype female (slides), from Tonatia silvicola (hosts no. 11352, 11547, resp.), Armila (San Bias), 1 and 12 March 1963, C. O. Handley, Jr. and F. M. Greenwell. In the col- lection of Chicago Natural History Museum. Paratypes. — 1, same data as holotype and 4, same data but 28 March 1963 ; 5, same host, Cerro Mali (Darien) , 8 February 1964, C. O. Handley, Jr. COLOMBIA, from Tonatia silvicola (=amblyotis) : 1, Puri, Cauca Valley above Caceres, 23 August 1952, P. Hershkovitz, CNHM Zoological Ex- pedition. REMARKS : This species is named in honor of Glen M. Kohls of the Rocky Mountain Laboratory in recognition of his outstanding contributions to our knowledge of ectoparasites, especially the systematics and biology of ticks. Strebla carolliae Wenzel, new species. Figures 122A, 136. Euctenodes mirabilis (not Waterhouse, 1879), Kessel, 1924, Parasitology, 16: 409 (descr.), 411 (part., from Hemiderma [=Carollia] perspicillatum ; 1925, Jour. N. Y. Ent. Soc., 33: 30 (part., from Hemiderma perspicillatum and Carollia p. azteca). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, pp. 654, 738 (part.). Pessoa and Guimaraes, 1940, Arq. Inst. Biol. Sao Paulo, 11: 426 (part., gives records of Kessel, 1925). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Ex., Fis. y Nat., 3: 418 (part., from Carollia p. azteca). Cooper, 1941 (part.), Yearb. Amer. Phil. Soc., 1941: 126 (chromosome no.). Jobling, 1949, Parasitology, 39: 318-19, 328 (part., from Carollia perspicillata) . Goodwin and Greenhall, 1961, Bull. Amer. Mus. Nat. Hist., no. 122, p. 250. 620 ECTOPARASITES OF PANAMA S. carolliae may be separated from all other species of the genus by the broadly blunt anterior margin of the postvertex. It is most closely related to alvarezi, a smaller species with a more narrowly rounded projection of the postvertex. DESCRIPTION: Head. — As illustrated (fig. 122A). Detached frontoclypeal plates wider than long. Postocular sclerite distinct. Anterior margin of postvertex very bluntly rounded. Lateral margins of postgena (above) not emarginate. Ante-ctenidial area of underside wider than long (25:17). Thorax (fig. 136). — Second or anterior pigmented suture of prescutum present, not well defined; epaulets with five setae; lateral discal setae extending to epaulets; arc of longer setae very distinct; three rows of setae at sides, between sutures. Antescutellar scutal setae subequal, except for long, outer macroseta on each side. Wings. — Fifth longitudinal vein basally with conspicuous bare area near base, this less than half as long as that of sixth vein. Costa without apical macroseta. Rs nearly twice as long (19.5:10) as distance from fork to crossvein r-m. Legs. — Fore- and midlegs very similar to those of altmani, metatibiae with two macrosetae, one subapical, the other at about apical fourth, preceded by a row of five or six setae that are about half as long as the macrosetae and twice as long as the other tibial setae. Ventral surface of first segment of hind tarsus with long heavy plantar bristles along outer edge, and shorter weaker ones along inner edge, these of about the same size as those on following segments. Abdomen. — Anterior face of tergum I+II with one short seta; inner dorsal margins of lateral lobes with four to five fine setae. Dorso-lateral setae of connexivum con- spicuously larger than ventrals. Female: Tergum VII with a pair of macrosetae and posterior to them, two or three setae that are less than half as long. Supra-anal plate with four apical setae only. Seventh sternites with 12-15 setae each. Ventral arc pro- duced anteriorly at middle as a narrowly triangular or spiniform process. Male: Sternum V with two transverse rows of discal setae at middle and three at sides in addition to apical marginal setae. Sternum VII+VIII (of hypopygium) on each side with a dorso-lateral macroseta and medial to it a short seta; tergum IX with a lateral macroseta on each side. Gonapophyses slender, angulately bent, apically knobbed, sides with scattered lateral microsetae apically; ventral macroseta inserted slightly posterior to basal third, extending almost to apex. Measurements: BL TL WL ww Male 1.99-2.31 0.74-0.81 1.42-1.46 0.71-0.75 Female 2.40-2.58 0.82-0.85 1.54-1.65 0.75-0.84 TYPE MATERIAL : Holotype male (host no. 4616) and allotype female (host no. 4618) , on slides, from Carollia perspicillata azteca, air-raid shelter, Fort Davis (Canal Zone) , 6 October 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes include only those specimens, listed below, that were collected from species of Carollia in Panama and in the countries to the north. Paratypes. PANAMA. — From Carollia p. azteca, 296 flies, from more than 287 bats, as follows : 15 (12 bats) , same data as the holotype and 6 (7 bats) , same locality, 17 November 1960, and 18 and 19 November 1959 ; 2, Camp Pina (Canal Zone) , 26 October 1960 ; 8 (4 bats) , Coco Solo (Canal Zone) , 13 January 1960 ; 1, Fort Clayton (Canal Zone) , 9 January 1953, F. Blanton ; 2, same locality, 8 December 1960; 5 (21 bats), Fort Gulick (Canal Zone), 15 September and 29 October 1959; 1, Old Battery, Fort Randolph (Canal Zone) , 1 October 1959 ; 10 (32 bats), Fort Sherman (Canal Zone), 23 Novem- ber and 4 December 1959 ; 3 (2 bats) , same locality, 16 March and 18 October 1960; 32 (66 bats), France Air Force Base (Canal Zone), 8 October to 30 November 1959; 11 (1 bat), Juan Mina (Canal Zone), 28 July 1960; 16 (23 WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 621 bats) , mine shaft, Coco Plantation, Gamboa Road (Canal Zone) , 9 September and 25 November 1959; 3 (3 bats), Brazoa Brook, Navy Tank Farm, 6 October 1959; 2 (2 bats), Rousseau Road, culvert, 13 October 1959; 18 (21 bats), railroad culvert, east of Summit Gold Club (Canal Zone), 26 October 1959; 9, "second culvert W. of bridge 5742 on the old RR" (Canal Zone), L. H. Dunn [MCZ] ; 4, "RR tower culvert at tower 34-12, Summit" (Canal Zone), L. H. Dunn [MCZ] ; 1, culvert, Gatun Tank Farm (Canal Zone), 19 November 1959 ; 9, Gatun Dam (Canal Zone), L.H.Dunn 185 [MCZ] ; 3, Camp Fig. 136. Thorax, dorsal view, Strebla carolliae, new species, male paratype from Carollia p. azteca (no. 3984), Rio Chilibrillo (Panama). Chagres, Madden Dam (Canal Zone), 21 and 25 June 1963, GML; 2, Summit (Canal Zone) [USNM] ; 7, same locality [cu] ; 2, same locality, K. W. Cooper [MCZ] ; 8, RR culvert, same locality, L. H. Dunn [MCZ] ; 5 (2 lots), same locality, 17 April 1957, C. B. Koford [USNM] ; 3, Gamboa (Canal Zone), 1 May 1957; 11, hollow tree, one mile from Gatuncillo (Canal Zone), 28 July 1960; 1, Taboga Island (Panama), L. H. Dunn 449-A [MCZ] ; 13 (10 bats), La Chorrera (Panama), 18 September 1959; 1, culvert, Chepo Road, 12 October 1959 ; 3, Chilibrillo Caves (Panama) , 28 October 1959 ; 7 (29 bats) , 18 August 1959 ; 37 (32 bats) , Buena Vista Cave (Colon) , 3 and 15 Septem- ber 1959, and 24 November 1959; 4, Porto Bello (Colon), 7 August 1923 [USNM det. as E. mirabilis by Kessel; see Kessel, 1924] ; 4, Penonome (Code), L. H. Dunn 704-A [MCZ] ; 1, Tacarcuna (Darien), 2 September 1958, P. Galindo [GML] ; 4 (2 bats), Rio Tuira (Darien), 26 February and 5 March 1958, P. Galindo [GML] ; 1, Rio Chucanaque, 19 February 1958, P. 622 ECTOPARASITES OF PANAMA Galindo [GML] ; 3 (3 bats), Punta Pina (Darien), 23 and 24 March 1960; 2 (2 bats), Guanico (Los Santos), 27 January 1962; 3 (3 bats), Cerro Hoya (Los Santos) , 18, 23, and 25 February 1962 ; 13 (11 bats) , Almirante (Bocas del Toro), 26 January to 4 February 1960. From Carollia castanea: 2 (2 bats) , Sibube (Bocas del Toro) , 19 and 23 January 1963, C. O. Handley, Jr. ; 2 (2 bats), Armila (San Bias), 17 February and 17 March 1963, C. 0. Handley, Jr. and F. M. Greenwell; 1, La Laguna (Darien), 2900 feet eleva- tion, 9 June 1963, GML; 1, Tacarcuna (Darien) , 2000 feet elevation, 19 July 1963, GML. From Carollia subrufa: 1(2 bats) , Isla Bastimentos (Bocas del Toro), 3 February 1963, C. 0. Handley, Jr. and F. M. Greenwell. From Carollia sp. : 1, Cerro Tigre (Canal Zone), 15 January 1953. From Glos- sophaga soricina leachii: 1, Bella Vista, L. H. Dunn, 397-B [MCZ]. From Desmodus r. murinus: 1, Chilibrillo Caves (Panama), L. H. Dunn [MCZ] ; 2, Almirante (Bocas del Toro) , 29 January 1960 ; 1, Fort Clayton, 13 October 1959. From Lonchorhina aurita: 1, two miles north of Summit, 17 April 1957 [USNM] ; 1, culvert E of Summit Golf Club (Canal Zone), 26 October 1959; 1, mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 25 November 1959. From Trachops cirrhosus: 1, Rio Mandinga (San Bias), 30 May 1957. From Macrophyllum macrophyllum: 4 (3 bats) , Fort Gulick (Canal Zone), 15 September 1959. From Phyllostomus h. panamensis: 1, Guanico (Los Santos), 26 January 1962; 1, hollow tree, Chepo Road (Pan- ama) , 8 October 1959. From Pteronotus parnellii fuscus: 3 (3 bats) , culvert east of Summit Golf Club (Canal Zone), 26 October 1959. From Natalus mexicanus: 1, Penonome (Code), L. H. Dunn [MCZ]; From Artibeus j. jamaicensis: 1, Almirante (Bocas del Toro) , 23 January 1960. From Loncho- phylla robusta: 1 (16 bats), Buena Vista Cave (Colon), 16 June 1960; 1, Chilibrillo Caves (Panama), 2 August 1960. From "Leaf nosed bat": 8, Chilibrillo Caves, 10 July 1945, [USNM]. Without host: 1, Barro Colorado Island (Canal Zone), 2 December 1956, K-19 [USNM]; 1, Farfan Cave (Canal Zone), L. H. Dunn 329-A [MCZ]. Non-Panamanian paratypes. GUATEMALA. — From Carollia p. azteca: 3 (3 bats), small cave, 0.2 mile northeast of Cueva de Lanquin, Lanquin (Alta Vera Paz) , 10 June 1948, R. D. Mitchell and Luis de la Torre, CNHM Guatemala Zoological Expedition, 1948. From Glossophaga s. leachii: 1, Sonarate (Progresso), 15 October 1948, Luis de la Torre. EL SALVADOR. — From "Glossophaga soricina or Carollia sp.": 1, near Santa Tecla (Liber- tad), Felten [SNl]. From Glossophaga soricina: 1, Mineral Encuentro, 19 June 1953, H. Felten [SKI]. BRITISH HONDURAS. — From Carollia p. azteca: 1, Belize, 7 October 1939, Ivan T. Sanderson. HONDURAS — Without host: 1, Coxen Hole, Ruatan Island, 9 January 1940, D. D. Davis, Field Museum- Leon Mandel Caribbean Expedition. OTHER MATERIAL EXAMINED: COLOMBIA. — From Carollia perspicillata: 1, Rio Raposo, 10 July 1962, C. J. Marinkelle. VENEZUELA. — From Carollia subrufa: 1, Rancho Grande, El Limon (Aragua), 21 March 1960, 3576 feet elevation, C. 0. Handley, Jr. From Carollia p. azteca: 1, same locality and collector, 30 March 1960; 4, Guacharo Cave ("Cerro de la Cueva"), C. Monagas, 16 August 1962 [FCUCV], From Artibeus lituratus: 1, 38 km. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 623 south of El Dorado (Bolivar), 2 August 1962, Ojasti [FCUCV]. TRINIDAD [TVL, T.H.G. Aitken, unless otherwise indicated]. — From Carollia p. per- spicillata: 1, Moko Estate, Maraval, 27 November 1956 ; 1, Masson Hospital, Port-of -Spain, 5 December 1956 ; 1, Arena Forest, Brazil, 8 January 1957 ; 4, Arena Trace (Los Lomas) , east of Cunupia, 5 September 1957 ; 1, Marocas Valley, 7 November 1954 ; 1, Belmont Valley Road, Port-of -Spain, 26 Febru- ary 1958 ; 4, Coroney Cave no. 1, Blue Range, Diego Martin, 5 May 1958 ; 1, Las Lomas, 27 June 1958 ; 6, La Fontaine Cave, Petit Valley, Diego Martin, 13 and 20 March 1959; 1, Waller Field, 5 July 1956; 1, near San Rafael, 17 February 1947, F. Wonder, CNHM Trinidad Zoological Expedition (1947). Without host: 3, Melapo Forest, 9 March 1956, W. G. Downs. TOBAGO. — From Carollia p. perspicillata: 2, Fort George, Scarborough, 2 September 1956. BRITISH GUIANA. — Without host: 7, Georgetown, 18 August 1924, A. E. Emerson. SURINAM. — From Carollia sp. : 1, Krakka-Pdehra Road (Surinam) , 25 October 1962, B. Malkin. BRAZIL. — From Carollia perspicil- lata: 1, Rocha (Sao Paulo) , 9 September 1961, A. M. Olalla. REMARKS : As may be judged from the above records, the normal hosts of this fly are species of Carollia. The fairly numerous but scattered records from other hosts reflect the ubiquity of the principal host, C. perspicillata (see "Host-parasite Relationships," below). Strebla altmani Wenzel, new species. Figure 123G, 137A. Distinct from all other species that have short inner occipital setae in that these are all fine setae, none are spinelets ; the male gonapophyses, too, are distinct in that they are rather abruptly narrowed and tapered from mid-length to apex. DESCRIPTION : Head. — Detached f rontoclypeal plates very small and weakly sclero- tized, difficult to see even in alcohol specimens. Anterior plates of laterovertices with about six setae ; postocular sclerite distinct. Dorsal surface of postgenae not emarginate laterally; with about ten short bristles, a long, conspicuous bristle near inner posterior margin and lateral to this a medium bristle. Postvertex and occiput as illustrated (fig. 123G) except that all of occipital setae are fine (not spinelets as shown). Eyes multi- faceted and projecting, about six large facets visible from above. Ventral ante- ctenidial area approximately as wide as long (17:17 or 17.5). Thorax (fig. 137A). — Second pigmented suture of prescutum present; epaulets with four or five setae; prescutal setae relatively sparse and more or less uniformly dis- tributed, setose area extending to epaulets along sides ; arc of long setae present but not conspicuous ; two or three irregular transverse rows of setae between transverse sutures. Scutum with two transverse rows of discal setae anterior to the row of subequal ante- scutellar setae. Wings. — Fifth longitudinal vein bare for a short distance before base, the bare area less than half as long as that of sixth vein. Costa without apical macro- seta. Rs more than twice as long (17.5:7.5) as distance from fork to crossvein r-m. Legs. — Pro- and mesotibiae dorsally with a double row of longer setae, only one or two of these on each side stronger than the others (three on inner edge of mesotibiae). Metatibiae with two subapical macrosetae preceded by a double row of setae that are much shorter and finer than the macrosetae but much longer than other tibial setae. Ventral surface of first segment of hind tarsi with two or three irregular rows of plantar setae, these not heavily pigmented and scarcely heavier than those of following segments. Abdomen. — Anterior face of tergum I+II with one short seta; inner dorsal margin of lateral lobes with three or four fine setae. Female : Tergum VII with two macrosetae 624 ECTOPARASITES OF PANAMA and posterior to these a pair of setae that are half as long. Supra-anal plate small, with four long apical setae only. Seventh sternites with nine or ten setae including three or four macrosetae. A small crescent-shaped postgenital sclerite present anterior to ventral arc; arc with two short and two minute setae. Male: Sternum VI very short, with only one transverse row of discal setae at middle, two laterally in addition to apical marginal setae. Hypopygium as in carolliae n. sp. Gonapophyses very slender, curved, apices not knobbed, with microsetae scattered along sides; macrosetae extending nearly to apex. Measurements: BL TL WL ww Male 1.40-1.49 0.49-0.52 1.10-1.13 0.51-0.55 Female 1.59-1.95 0.56-0.59 1.24-1.31 0.63-0.65 TYPE MATERIAL: Holotype male (slide) and allotype female (slide) (host no. 4191) from Lonchorhina aurita aurita, mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 September 1959, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — From Lonchorhina a. aurita, 208 flies (37 lots, 47 bats) as follows: 163 (29 lots, 31 bats), same data as the holotype; 26 (10 bats), same data but 25 November 1959; 13 (5 bats), railroad culvert east of Summit Golf Club (Canal Zone), 26 October 1959; 6 (1 lot), two miles north of Summit, 17 April 1957 [USNM]. From plastic bag containing Carollia p. azteca and Lonchorhina a. aurita: 25, mine shaft, Coco Planta- tion, Gamboa Road (Canal Zone), 25 November 1959. From Macrophyl- lum macrophyllum, 108 flies (19 lots, 46 bats) as follows: 83 (18 lots, 21 bats) , Fort Davis (Canal Zone) , 13 October 1959 ; 25 (25 bats) , same locality, 29 July 1960; 21 (9 bats), Natural Bridge, Madden Dam (Canal Zone), 31 August 1959; 2, Rio Tuira (Darien), 10 March 1958, P. Galindo [GML] ; 2 (2 bats), culvert, Chepo Road (Panama), 12 October 1959. From Carollia p. azteca: 1, air raid shelter, Fort Davis (Canal Zone), 6 October 1959; 20 (13 bats), mine shaft, Coco Plantation, Gamboa Road (Canal Zone), 9 September 1959; 8 (18 bats), same locality, 25 November 1959; 1, railroad culvert east of Summit Golf Club (Canal Zone) , 26 October 1959 ; 2 (2 bats) , Buena Vista Cave (Colon) , 3 September 1959. From Trachops c. cirrhosus: 1 (15 bats) , culvert, Chepo Road (Panama) , 24 May 1960. From Pteronotus parnellii fuscus : 1, mine shaft, Coco Plantation, Gamboa Road (Canal Zone) , 25 November 1959. From unidentified host (probably Macrophyllum macrophyllum) : 1, Natural Bridge, Madden Dam (Canal Zone), 31 August 1959. Non-Panamanian paratypes. — From Lonchorhina aurita: 12 (4 lots), Rancho Grande (Aragua), VENEZUELA, 18 July, 7 and 12 August 1949, Dr. J. Racenis [FCUCV] . Paratypes to be deposited in the collections listed on p. 410. REMARKS : S. altmani appears to be a normal parasite of both Lonchor- hina aurita and Macrophyllum macrophyllum, closely related genera of phyllostomine bats. We are unable to distinguish between specimens from these two hosts. However, it appears to occur in larger numbers per indi- vidual host bat on Lonchorhina than on Macrophyllum (See "Host-parasite Relationships"). It also occurred consistently on Carollia p. azteca, in larger numbers than one would expect if these were merely strays or transi- tory transfers, but only where Carollia was roosting with one of the normal WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 625 hosts. It was never taken on Carollia in the absence of one of these hosts. This species is named in honor of Lt. Col. Robert M. Altman, in recogni- tion of his important contributions to our knowledge of the ectoparasites of Panama, especially his initiation of a systematic survey. Strebla alvarezi Wenzel, new species. Figures 123E, F ; 137B. Closely related to carolliae n.sp., but easily distinguished from that species by its smaller size, the shape of the anterior margin of the post- vertex (fig. 123E, F) which is not broadly blunt as in carolliae (fig. 122A), the much smaller detached f rontoclypeal plates, and the less strongly curved male gonapophyses. A Fig. 137. Thorax, dorsal view. A, Strebla altmani, new species, male paratype from Lonchorhina aurita (no. 4220), mine shaft, Coco Plantation, Gamboa Road (Canal Zone). B, Strebla alvarezi, new species, paratype male from Micronycteris megalotis (no. 7955), near Huile (Canal Zone). DESCRIPTION: Head. — Frontoclypeus with apical detached plates small, wider than long. Anterior plate of laterovertices with six setae. Postocular sclerite not dif- ferentiated. Postgenae not emarginate laterally, posteriorly with eight or nine short setae, a conspicuously longer seta near inner posterior margin, and lateral to this, a seta of intermediate length. Eyes multi-faceted and projecting, about seven facets visible from above, the tipper ones not very distinct. Ventral ante-ctenidial area a little wider than long (17.5:14.5). Postvertex and occipital lobes as in fig. 123E, F. Thorax (fig. 137B). — Prescutum with well-defined second pigmented suture; epaulets consisting of four setae; prescutal setae sparse, setose area not reaching epaulets; arc of longer setae present; two transverse rows of setae laterally between transverse mesonotal and second pigmented suture. Scutum with about eight subequal ante- scutellar setae and two short median ones, in addition to a long macroseta each side, in addition to the discal setae. Wings. — Seta at apex of costa not conspicuously long. Rs about three times as long (22:7) as distance from fork to crossvein r-m; Rs and fifth longitudinal vein bare basally but not as extensively as sixth. Legs. — Protibiae dorsally with three macrosetae on inner edge and two on outer; mesotibiae similar. Metatibiae with two macrosetae on apical third, preceded by a row of gradually smaller setae, the 626 ECTOPARASITES OF PANAMA basal one scarcely longer than the numerous short tibial setae. Ventral surface of first segment of hind tarsus with three irregular, longitudinal rows of plantar bristles, these not heavily sclerotized, distinctly longer and heavier than those of following seg- ments, though these are not minute. Abdomen. — Dorsal inner margin of lateral lobes of tergum I+II with three to five setae. Dorso-lateral connexival setae scarcely longer than those of venter. Female: Tergum VII with a pair of macrosetae and, posterior to these, a pair of shorter setae. Supra-anal plate with four apical macrosetae only. Seventh sternites with seven to nine setae, three of them much longer than the others. Ventral arc triangularly but not strongly produced anteriorly, with four setae, two larger, two minute. Male: Sternum V with two transverse rows of discal setae in addition to long marginals. Sternum VII+VIII dorsally with a macroseta on each side; tergum IX dorsally with a single macroseta on each side. Gonapophyses long, slender, straight on basal two-thirds, curved on apical third; sides apically with scattered microsetae, apices rounded; ventral macroseta inserted at about basal third, extending about two-thirds the distance to apex; accessory seta minute. Measurements: BL TL WL ww Male 1.59-1.76 0.58-0.59 1.10-1.21 0.55-0.60 Female 1.92-2.14 0.63-0.67 1.35-1.37 0.63-0.66 TYPE MATERIAL: From Micronycteris megalotis: Holotype male and allotype female (slides) from host no. 7961, taken from culvert under Borinquen Highway near Empire Range, 24 October 1961, C. M. Keenan and R. L. Wenzel. In the collection of Chicago Natural History Museum. Para- types. — 4 males, 1 female, in road culvert one-half mile SE of Empire Range (Canal Zone) , 23 October 1961, R. L. Wenzel and C. M. Keenan ; 2 males (2 bats) , "French Vault," west bank of Canal, 300 yards east of Empire Range, 23 October 1961, same collectors; 2 males, 2 females (5 bats), same data as holotype; 1 male, 1, sex unknown (2 bats), hollow espave tree, Huile (Canal Zone), 19-20 October 1961, same collectors; 1 male, Loma Boracho (Canal Zone) , 20 September 1959 ; 1 male, Casa Tilley, Cerro Punta (Chiri- qui), 23 January 1960; 1 male, Isla Taboga (Panama), 21 October 1959; 1 female, Guanico (Los Santos) , 27 January 1962. From Micronycteris nice- fori: 3 males, Almirante (Bocas del Toro) , 12 February 1960. From Micro- nycteris sylvestris: 2 males, Armila (San Bias), 12 March 1963, C. 0. Handley, Jr. and F. M. Greenwell. From Saccopteryx bilineata: 4 males (2 bats) , hollow tree, Chepo Road (Panama) , 8 October 1959. EL SALVADOR. — From Saccopteryx bilineata: 1, 5 km. E of La Libertad (La Libertad), 24 October 1953, H. Felten [SNI]. From a mixed collection of Saccopteryx bilineata and Glossophaga s. leachii: 10, coastal forest, Hacienda Nancuchi- name, San Marcos Lempe (Usulutan), 25 March 1954, H. Felten [SNl]. GUATEMALA. — From Saccopteryx bilineata: 7, Finca Santa Cristina, 6 mi. S of Democracia (Escuintla), 20 September 1948, L. de la Torre, CNHM Guatemala Zoological Expedition (1948). Paratypes to be deposited in the collections of Chicago Natural History Museum ; the United States National Museum; the Gorgas Memorial Laboratory, Panama; the Environmental Health Branch, U. S. Army, at Corozal (Canal Zone) ; and the Senckenberg Naturf orschende Institut, Frankfurt a/Main. OTHER MATERIAL EXAMINED : A badly damaged specimen, without host, "Rio Waupes, BRAZIL," collected by the Olalla brothers [LI-13, MCZ] ; an additional male lacking head but probably this species, from Micronycteris WENZEL, TIPTON, AND KIEWLICZ I STREBLID BATFLIES 627 megalotis, Casa Tilley, Cerro Punta (Chiriqui), PANAMA, 6 March 1962. REMARKS : The specimens from Guatemala have the anterior margin of the postvertex somewhat more pointed (fig-. 123F) than those from Pan- ama, (fig. 123E) and they are on the average somewhat smaller. However, we consider them to be the same. The length and disposition of the occipital setae vary, within both populations. S. alvarezi seems to normally occur on species of Micronycteris and on Saccopteryx bilineata, bats which belong to different families. This is prob- ably accounted for by the fact that they often roost in close proximity. S. bilineata roosts on the shaded outside of logs, tree trunks, and branches, and in houses, caves, etc., while M. megalotis (and perhaps other species of Micronycteris) roosts inside logs, stumps and tree holes, and in houses, culverts, etc. S. alvarezi is named for Mr. Vicente Alvarez, of the Environmental Health Branch, USAFSC, at Corozal (Canal Zone) , in grateful appreciation of his valued assistance on this project. Genus Paraeuctenodes Pessoa and Guimaraes Paraeuctenodes Pessoa and Guimaraes, 1937, Ann. Fac. Med. Sao Paulo, 12, (2), pp. 257-258. Jobling, 1939, Arb. Morph. Tax. Ent., 6, (3), pp. 269 (discuss.), 270 (keyed). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact, Fis. y Nat., 3: 417 (keyed) ; 1942, Bol. Ent. Venez., 1: 85 (keyed). Jobling, 1947, Parasitology, 39: 322 (keyed). Hoffman, 1953, Mem. Congr. Cient. Mex., 7: 179 (keyed), 189. Type-species: Paraeuctenodes longipes Pessoa and Guimaraes, 1937. This genus contains a single described species, P. longipes Pessoa and Guimaraes (op. cit., p. 258) . The holotype female was taken from Loncho- glossa ecaudata at Ipiranga, Sao Paulo (Sao Paulo) , Brazil, and the allotype male from Phyllostomus hastatus at Sao Paulo. Paraeuctenodes resembles Strebla in having well-developed dorsal postgenal sclerites and in lacking a remiform postgenal seta. However, it resembles Anastrebla in lacking metatibial macrosetae and in having elongated hindlegs. The f rontoclypeus is entire as in Anastrebla and in some species of Strebla. There are two specimens of this genus in the collection of Chicago Natural History Museum. They appear to represent two new species. One was taken from Carollia p. perspicillata, near San Rafael, TRINIDAD, and the other (doubtfully associated with Carollia p. azteca) was collected in a small cave northeast of Cueva de Lanquin (Alta Vera Paz), GUATEMALA. Thus, it seems highly probable that a species of this genus will be found in Pan- ama. Anastrebla Wenzel, new genus Type-species: Anastrebla modestini, new species. Strebla (not Wiedemann, 1824), Speiser, 1900, Arch. Naturg., 66A, Bd. I, pp. 38-43, 63 (cit.), 65 (keyed). Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 26 (keyed), 31 (cit.). Kessel, 1924, Parasitology, 16: 406-409 (char.); 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 29 (char.). Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 653 (part.). Curran, 1934, Man. Fam. Gen. N. Am. Dipt., p. 479 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28: 355 ff. (morph., syst. pos.), 370 628 ECTOPARASITES OF PANAMA (keyed). Pessoa and Guimaraes, 1937, Ann. Fac. Med. Sao Paulo, 12: 235 ff. (comp. notes). Jobling, 1939, Arb. Morph. Tax. Ent., 6: 269 (discuss.). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed); 1942, Bol. Ent. Venez., 1: 85 (keyed). Jobling, 1949, Parasitology, 39: 322 (keyed). Hoff- mann, 1953, Mem. Congr. Cient. Mex., 7: 178 (keyed), 188 (cit.). Maa, 1963, Jour. Med. Ent., 1: 385. The name Strebla (q.v.) must be used for the species assigned to the genus Euctenodes Waterhouse by Waterhouse, Kessel, and subsequent authors. The genus Anastrebla is proposed for Strebla of Speiser (loc. cit.) and subsequent authors, not Wiedemann, 1824. DIAGNOSIS: With the characters attributed to Strebla (not Wiedemann, 1824) by Kessel (1924) and by Jobling (1939 and 1949). In general, with the characters of Strebla Wiedemann, but differing from that genus chiefly in that the hind legs are noticeably more elongated; the metatibiae are slender and lack macrosetae. Postgenae (fig. 39D) poorly developed dorsally (well developed in Strebla, fig. 122) and posteriorly bearing a remiform scale (lacking in Strebla). REMARKS : The species of Anastrebla superficially resemble those of Para- euctenodes and have similar hindlegs, but the postgenae of Paraeuctenodes are similar to those of Strebla, and, similarly, lack a remiform scale. The species of this genus appear to be restricted to bats of the subfamily Glossophaginae. The three species known to us are from Anoura cultrata, A. geoffroyi, A. aculeata, and Lonchophylla robusta. The species treated as Strebla vespertilionis by Speiser (1900, pp. 38-41 ; pi. 4, figs. 1, 2) and Kes- sel (1924, p. 413, figs. 1, 5, 6; 1925, p. 24 [part.], pi. 3, fig. 20, pi. 4, fig. 24) was collected from Lonchoglossa ecaudata. We have not seen specimens from this host, but it is clear that the species is closely related to, if not identical with Anastrebla modestini n. sp. The streblid figured by Jobling (1929, fig. 4; 1949, fig. 3F) as Strebla vespertilionis is also a species of Anastrebla and is distinct from any known to us. It is characterized by exceptionally short festoon spinelets on the postvertex and occipital plates. Mr. Jobling (pers. comm.) has informed us that his illustration is based on a specimen, without host or locality data, that was given to him by Hugh Scott. KEY TO THE SPECIES OF ANASTREBLA 1. Eyes with a single hyaline cornea, without externally distinct facets. Postvertex and occiput with relatively short, heavy spine-like setae that are not noticeably tapered. Host: Lonchophylla robusta nycteridis n.sp. Eyes with seven or eight externally distinct facets, their corneae separate and distinct. Setae of postvertex and occiput longer, distinctly tapered apically, with long points. Hosts : Anoura spp 2 2. Prescutum (fig. 138A) sparsely setose, the median setose area usually not ex- tending beyond the anterior suture. Wing vein Ri without setae on basal half or more, sixth vein with a few setae apically near the third crossvein, and usually with one to three proximal to mid-length and one or more beyond mid- dle. Median setose area of sternum I narrow, apical margin with about 15 setae modestini n.sp. Prescutum (fig. 138B) more densely setose, the median setose area extending be- yond the anterior pigmented suture. Ri and sixth longitudinal wing veins bare at base only. Setose area of sternum II broader, apical margin with 20-22 setae mattadeni n. sp. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 629 Anastrebla nycteridis Wenzel, new species. Figure 139A, B. Closely related to A. mattadeni n.sp., but differing as follows : the eyes are not distinctly faceted and projecting but have only a few vague facets that are not distinct in alcohol-preserved specimens, but can be detected in slide preparation (fig. 139A) ; the laterovertices are much more elongated; the spinelets of the postvertex and occiput are relatively short and stout ; the male gonapophyses are heavier and much straighter, only feebly curved nearly to apex (fig. 139B) . The female of nycteridis is unknown. DESCRIPTION (Male) : Head. — Anterior division of laterovertices with seven setae, the postero-median seta a very conspicuous macroseta but shorter than in mattadeni and modestini n. spp., not reaching posterior margin of occiput; posterior division with two setae. Dorsal sclerite of postgena with a long stout seta, no longer than and inserted anterior to the remiform seta. Postvertex and occiput with ten stout spinelets (fig. 139A), the inner four pairs subequal, the outer one distinctly shorter. Eyes a single elongate hyaline lens, facets not externally distinct, though they may be discerned in slide preparations at high magnifications (fig. 139A). Ventral ante-ctenidial area about as wide as long (24:23). Thorax. — Prescutum with anterior transverse suture. Chaetotaxy much as in mattadeni; with three irregular transverse rows of setae along side, the setose area ex- tending further anteriorly along mid-line. Twelve antescutellar setae, the middle four only slightly longer than longest discal scutal setae, the outer longer antescutellars stouter, subspinif orm ; discal setae forming a single transverse row at mid-line and three at the sides. Wings. — Setae lacking along base of Rs (basal half or less) a short basal area on fourth longitudinal vein as well as the usual basal area on the sixth longitudinal vein. Rs distinctly longer (20:15) than distance from fork to crossvein r-m. Legs. — Hind femora longer than thorax, very similar in chaetotaxy to those of mattadeni. Abdomen. — Inner dorsal margins of tergum I +11 with three very fine setae. Sternum II much as in mattadeni, but median setose area extending only a little more than halfway to base; with about 18 marginal setae. Hypopygium essentially as in mattadeni. Gonapophyses (fig. 139B) similar to those of mattadeni and modestini (fig. 139D, F), but straighter and heavier. Measurements: BL TL WL ww Male 2.40-2.42 0.80-0.82 1.70-1.95 0.74-0.93 TYPE MATERIAL : Holotype male (slide) from Lonchophylla robusta (host no. 5434), Almirante (Bocas del Toro), 28 January 1960, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Para- type male, same host, La Laguna (Darien) , elevation 2900 feet, GML ; in the collection of the United States National Museum. Anastrebla modestini Wenzel, new species. Figures 138A ; 139C, D. Distinguished from mattadeni by the different shape of the laterovertices and the shorter setae of the postvertex and occiput (fig. 139C), and espe- cially by the much sparser mesonotal chaetotaxy (fig. 138A) . DESCRIPTION : Head. — Anterior division of laterovertex with ten setae, the postero- median one a very conspicuous macroseta; posterior division with two long bristles. Dorsal sclerite of postgena with a long macroseta, inserted anterior to remiform scale, and extending posteriorly beyond margin of head. Postvertex and occiput, together, with five pairs of heavy sub-equal but narrowly pointed spine-like setae, the pair on postvertex slightly longer than the occipitals, the outer pair only slightly shorter than the others. Eyes with about seven or eight distinct facets visible from above. Ventral ante-ctenidial area wider than long (26:20). Thorax. — Prescutum with a second transverse suture; a pair of widely separated 630 ECTOPARASITES OF PANAMA short setae on anterior margin; epaulets consisting of four setae; setal arc consisting of four or five strong setae that are appreciably longer than the discals ; three irregular transverse rows of discal setae between the transverse and anterior sutures, the setose area not extending beyond the anterior suture. Antescutellar row consisting of 10-12 setae, the middle five shorter than the others which are stouter. Discal setae half as long to slightly longer than antescutellars, limited to a single transverse row at middle and two rows laterally. Wings. — Very long; Ri lacking setae on basal half or more in Fig. 138. Thorax, dorsal view. A, Anastrebla modestini, new species, male paratype from Anoura geoffroyi lasiopyga (no. 6218), Casa Lewis (Chiriqui). B, A. mattadeni, new species, paratype from A. g. lasiopyga (no. 5343), Rancho Grande (Aragua), VEN- EZUELA. mattadeni; Rs bare on basal third to half; sixth longitudinal vein with a few setae apically near third crossvein, and usually with one to three proximal to mid-length and one or more beyond middle, occasionally with more setae but these always much more widely separated than setae of other veins; Rs about three times as long (26:8.5) as distance from fork to crossvein r-m. Legs. — Markedly elongated, chaetotaxy very much as in mattadeni. Abdomen. — Very much as in mattadeni. The median setose area of sternum I narrow and extending anteriorly only a little, beyond middle. Apical margin with only about 15 setae, including two macrosetae; four setae between the macrosetae. Female: Only one or two longer setae along inner edges of dorsal setose area of dorsal con- nexivum, the apical one a macroseta. Chaetotaxy of tergum VII and supra-anal plate as in mattadeni. Ventral arc with a pair of longer setae on posterior margin, and a pair of minute ones on anterior margin; not produced, but with a microsetose non- sclerotized extension as in mattadeni. Male: Hypopygium as in mattadeni. The gon- apophyses strongly tapered, and rather strongly curved apically; scattered microsetae along sides on about apical fourth; macroseta inserted at about basal fourth, very long, extending to apex; accessory seta well developed. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 631 Measurements: BL TL WL ww Male 1.98-2.17 0.77-0.82 1.92-2.06 0.77 Female 2.36 0.85 2.14 0.71 TYPE MATERIAL: PANAMA. — Holotype male from Anoura geoffroyi lasiopyga (host no. 6218), Casa Lewis, Cerro Punta (Chiriqui), elevation above 5000 feet, 4 May 1960, C. M. Keenan and V. J. Tipton. In the collec- tion of Chicago Natural History Museum. Paratypes. — From A. g. lasiopyga: a male, Cerro Hoya (Los Santos), elevation 1500-3200 feet, 11 February 1962. GUATEMALA. — From A. g. lasiopyga: a male, Santa Elena (Chimaltenango), 10,000 feet elevation, 26 January 1934, F. J. W. Schmidt, Field Museum-Leon Mandel Guatemala Zoological Expedition. From Anoura sp. (CNHM cat. no. 73362) : a male and female, San Lorenzo, 4 mi. NE of Volcan de Jumay ( Jalapa) , elevation 3750 feet, 15 February 1952, L. de la Torre, CNHM Guatemala Zoological Expedition (1951-52). MEXICO. — From A. g. lasiopyga: 1, Santa Lucia (Sinaloa), elevation 3200 feet, 28 July 1963, J. Knox Jones. TRINIDAD. — From Anoura geoffroyi [geoffroyi'] : a male, Aripo Cave, 19 March 1955, W. G. Davis [RML no. 33590] ; a male, Tamana Caves, 11 December 1954, C. C. Sanborn, CNHM Trinidad Zoological Field Trip. Paratypes to be deposited in the collections of Chicago Natural History Museum ; the United States National Museum ; and the Environmental Health Branch, USAFSC, at Corozal (Canal Zone). OTHER MATERIAL EXAMINED : 3, without host identification, from Cueva Nanarita, Hacienda El Marne (Santa Ana) , EL SALVADOR, 960 meters eleva- tion, H. Felten [SNl]. REMARKS: A. modestini appears to be restricted to Anoura geoffroyi. It is named for Mr. Sulpice Modestin of the Environmental Health Branch, at Corozal (Canal Zone), in appreciation of his assistance in the field. Anastrebla mattadeni Wenzel, new species. Figures 138B ; 139E, F. Easily distinguished from A. nycteridis n.sp. and modestini n.sp. by the long slender setae of postvertex and occiput (fig. 139E) ; from modestini which has similar eyes and laterovertex, it may further be separated by its very different mesonotal chaetotaxy (fig. 138B) . DESCRIPTION : Head. — Anterior division of laterovertex with nine or ten setae. Dorsal sclerite of postgena with a conspicuous heavy posterior macroseta which is much longer than the remiform seta and extends beyond occiput. Postvertex and occiput with five pairs of long, tapering macrosetae which are gradually shorter toward sides, the inner pair conspicuously longer than the outer. Eyes strongly projecting, multi-faceted, seven or eight facets visible from above. Ventral ante-ctenidial area much wider than long (29:21.5). Thorax (fig. 138B). — Prescutum with pigmented suture; epaulets diagonal, with four strong short setae; arc consisting of four to five strong conspicuous setae, no discal setae anterior to them; setae of arc nearly twice as long as discals, interval between transverse mesonotal suture and second pigmented suture with four transverse rows of setae toward side, but five or six along middle, where setose area extends anteriorly to a point just beyond level of second suture. Antescutellar row of setae consisting of about 12-14 long very strong setae shorter at middle, longer laterally, the shortest one not quite twice as long as the longest discal scutal setae, of which there are a single irregular transverse row at middle and two to three laterally. Wings. — Ri and Rs without setae 632 ECTOPARASITES OF PANAMA Fig. 139. A and B, Anastrebla nycteridis, holotype; A, laterovertices, postvertex, and occipital plates (of head) ; B, left male gonapophysis. C and D, Anastrebla modestini, new species, same structures; C, male paratype from Anoura g. geoffroyi (C. C. Sanborn no. 552), Tamana Caves, TRINIDAD; D, holotype male. E and F, Anastrebla mattadeni, new species; E, female paratype from Anoura aculeata (no. 5344), Rancho Grande (Aragua), VENEZUELA; F, paratype from Anoura geoffroyi (no. 5343), same locality. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 633 basally, (Rs sometimes without any setae), fourth sometimes bare at base; sixth with- out setae basally, as usual. Rs nearly twice as long (20.5:11-12) as distance from fork to crossvein r-m. Third crossvein strongly bent and recurrent. Protibiae with a double row of 10-12 longer strong setae, the apical ones macrosetae; mesotibiae with 14^16 strong setae ; metatibiae with short setae only. Pro- and mesof emora with a few macro- setae dorsally and laterally on apical half, metafemora with a conspicuous dorsal row of six to eight macrosetae. First segment of all tarsi as long as the next three ; under- side of hind tarsi with a double row of stronger plantar bristles that are longer than the others, the outer row of about four much stronger; lateral to these are minute plantars; succeeding segments with minute plantars. Abdomen. — Dorsal inner margins of lateral lobes of tergum I+II with five to six very long, thin setae; anterior face with about five short setae. Sternum II strongly, densely setose in a median triangular area which extends to about basal fourth; apical margin with 20-22 longer setae along apical margin, two of them macrosetae, one each side of middle, with seven or eight setae between them. Female: Dorsal connexivum with a few longer setae at base and a few in a single row along inner margin of each lateral setose area, the others very short, about half as long as ventral connexival setae; four pairs of widely separated short segmental setae. Tergum VII with a pair of exceptionally long macrosetae; medial and posterior to these a pair of very short setae. Supra-anal plate with four apical macrosetae and just anterior to them a pair of very short widely separated setae. Seventh sternites with 11-12 long setae, five to seven of them long macrosetae, several of these exceptionally long. Ventral arc not produced, though there is a lobe-like, apparently non-sclerotized, microsetose area extending an- teriorly from its dorsal edge; arc with a single pair of short setae. Male: Sternum V not differentiated. Hypopygium well developed, with a short seta and two or three macrosetae on each side of sternum VII+VIII, and a row of about eight macrosetae along dorsum and sides of tergum IX. Measurements: BL TL WL ww Male 2.05-2.14 0.77-0.82 1.84-1.92 0.77-0.82 Female 2.17-2.31 0.82 1.98-2.17 0.81-0.89 TYPE MATERIAL: PANAMA. — Holotype male (slide) and allotype female (alcohol) from Anoura cultrata (host no. 8593) , upper Rio Changena Camp (Bocas del Toro) , elevation 2500 feet, 27 September 1961, C. M. Keenan and V. J. Tipton. In the collection of Chicago Natural History Museum. Paratypes. — From Anoura cultrata: 15 (2 bats), Cerro Mali (Darien), elevation 4100-4800 feet, 2 and 13 February 1964, C. 0. Handley, Jr. ; 2 (2 bats), Cerro Tacarcuna (Darien), elevation 4100-4800 feet, 20 February and 9 March, Handley. From Anoura geoffroyi lasiopyga: 1, Rancho Mojica (Bocas del Toro), elevation 4800 feet, 12 September 1961. VENEZUELA (Rancho Grande [Aragua], 30 March 1961, C. 0. Handley, Jr.). — From Anoura g. lasiopyga: 3(2 bats) . From Anoura aculeata: 5 (2 bats) . From Enchisthenes hartii: 1. Paratypes to be deposited in the collections of Chi- cago Natural History Museum; the United States National Museum; the Environmental Health Branch, USAFSC, at Corozal (Canal Zone) ; the Departmento de Zoologia, Secretaria da Agricultura, Sao Paulo, Brazil ; and the Faculdad de Ciencias, Universidad de Venezuela. REMARKS : This species is named for Mr. Edmond Mattaden of the En- vironmental Health Branch, at Corozal (Canal Zone) in appreciation of his assistance in the field and in the laboratory. A. mattadeni is recorded from three species of Anoura. However, we suspect that it is primarily a parasite of A. cultrata and A. aculeata and that the records from A. g. lasiopyga may be in error, either because of misidenti- 634 ECTOPARASITES OF PANAMA fication of the host or contamination in the field. The record from Enchis- thenes hartii is obviously doubtful and probably represents a contamination. Genus Metelasmus Coquillet Metelasmus Coquillet, 1907, Ent. News, 18: 292. Costa Lima, 1921, Arch. Esc. Sup. Agric. Med. Vet., Nictheroy, 5: 26 (keyed), 32 (cit.). Kessel, 1925, Jour. N. Y. Ent. Soc., 33: 13 (keyed), 31. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 654. Curran, 1934, Fam. Gen. N. Am. Dipt., p. 477 (keyed) ; 1935, Amer. Mus. Novit., no. 765, p. 5 (keyed). Jobling, 1936, Parasitology, 28, (3), p. 356 ff. (morph.), 370 (keyed); 1939, Parasitology, 31: 494 (synonymizes Lemosia Pessoa and Galvao, 1936) ; 1939, Arb. Morph. Tax. Ent., 6, (3), p. 269 (discuss.), 270 (keyed). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 417 (keyed); 1942, Bol. Ent. Venez., 1: 85 (keyed). Jobling, 1949, Parasitology, 39: 322 (keyed). Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 178 (keyed), 189 (cit.). Lemosia Pessoa and Galvao, 1936, Revista Ent., 6: 243 (type-species: Lemosia setosa Pessoa and Galvao, 1936). Type-species: Metelasmus pseudopterus Coquillet, 1907. This genus contains a single species, M. pseudopterus Coquillet, the only brachypterous species of the subfamily Streblinae. Metelasmus re- sembles Anastrebla in possessing a remiform scale at the posterior margin of the dorsal sclerite of the postgena, and in lacking macrosetae on the meta- tibiae. However, the frontoclypeus has apical detached plates as in many species of Strebla. Metelasmus pseudopterus Coquillet. Figures 39A, B ; 140. Metelasmus pseudopterus Coquillet, 1907, Ent. News., 18: 292, fig. — Sapucay, Paraguay, from Artibeus lituratus. (type 10293, United States National Museum). Seguy, 1926, Ency. Ent., (B), 11, Dipt., 3: 192-194, fig. 1. Stiles and Nolan, 1931, Bull. Nat. Inst. Hlth., Wash., no. 155, p. 654. Jobling, 1936, Parasitology, 28, (3), pp. 370-374 (redescr.), figs. 2, 3A-E. Jobling, 1939, ibidem, 31: 494 (synonymizes Lemosia setosa Pessoa and Galvao, 1936) ; 1939, Arb. Morph. Tax. Ent., 6 (3), p. 269 (discuss.), 270 (keyed). Bequaert, 1940, Rev. Acad. Colomb. Cienc. Exact., Fis. y Nat., 3: 418; 1942, Bol. Ent. Venez., 1: 87 (records). Jobling, 1949, Parasitology, 39: 316 ff., fig. 3C. Grandi, 1952, Introduz. Stud. Ent., 2: 478. Hoffmann, 1953, Mem. Congr. Cient. Mex., 7: 189 (records). Lemosia setosa Pessoa and Galvao, 1936, Revista Ent., 6: 244-248, figs. 1-4. PANAMANIAN MATERIAL EXAMINED : 49 flies, 38 lots, from more than 48 bats. From Artibeus j. jamaicensis, 30 lots, 41 flies (40 bats) , as follows : 6 (3 bats), Almirante (Bocas del Toro), 27 January to 1 February 1960; 1, Natural Bridge, Madden Dam (Canal Zone), 31 August 1959; 2, Barro Colorado Island (Canal Zone), 10 December 1956 [USNM] ; 2, Punta Pina (Darien), 24 March 1960; 7 (4 bats), Rio Seteganti (Darien), 31 January to 5 February 1961; 2 (2 bats), Rio tuira, 25 and 26 February 1958, P. Galindo [GML] ; 6 (4 bats), La Laguna (Darien), 2 to 15 June 1963, GML; 5 (5 bats), Tacarcuna (Darien), 20 June to 12 July 1963, GML; 8 (7 bats), Cerro Hoya (Los Santos) , 8 to 24 February 1962. From Artibeus lituratus palmarum: 2 (2 lots) , Barro Colorado Island (Canal Zone) , 2 and 3 January 1957. From Carollia perspicillata azteca: 2 (2 bats) , Almirante (Bocas del Toro), 24 and 30 January 1960; 1, Tacarcuna (Darien), 2 September 1958, GML. From Vampyressa nymphaea: 1, Rio Mono Camp (Darien), 25 July WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 635 Fig. 140. Metelasmus pseudopterus Coquillet, male, dorsal view. From Jobling (1936). 1963, GML. Without host: 2 (2 lots), Tacarcuna (Darien), 20 June 1963, GML. NON-PANAMANIAN MATERIAL EXAMINED: BRAZIL. — From "ArtlbeUS jamaicensis literatus": 3, Joinville (Santa Catharina) , 8 May 1929. GUATE- MALA (CNHM Guatemala Zoological Expedition, 1948). — From Artibeus j. jamaicensis : 1, Cueva de Los Ladrones, Finca Los Arcos (Escuintla), 27 636 ECTOPARASITES OF PANAMA September 1948, L. de la Torre. From Sturnira lilium parvidens: 1, Finca El Zapote, Zapote (Escuintla), 2400 feet elevation, 14 July 1948, R. D. Mitchell and L. de la Torre. MEXICO. — From Artibeus jamaicensis : 1, Cueva de Zapaluta (Chiapas), 5000 feet elevation, 26 June 1960, D. C. Carter. From Enchisthenes hartii: 1, La Catarina, 2 miles north of Ciudad Guzman (Jalisco), 11 December 1954, L. de la Torre, CNHM Mexican Zoological Field Trip (1954). REMARKS : M. pseudopterus appears to be a parasite of stenodermine bats, especially of the genus Artibeus. The records from Carollia may represent contaminations. The specimen taken from Enchisthenes hartii in Guate- mala may represent a second, very closely related new species. HOST PARASITE RELATIONSHIPS Definitions The semantic difficulties in discussing host-parasite relationships are considerable. Terms such as principal, primary, true, secondary, original, exceptional, and normal, as applied to hosts reflect the attempts to categorize relationships that in a sense cannot be categorized. Yet these terms may be valuable, if their use is clearly understood. We have used the terms host specificity and host-parasite specificity "in relation to the [taxonomic] range of hosts" (Rogers, 1962, p. 219). How- ever, as Caullery (1952, p. 182) has emphasized, one must recognize the distinction "between specificity in fact and in principle." 12 It is also nec- essary to distinguish between apparent specificity, based on limited obser- vations, and specificity based on extensive observations throughout the geo- graphic range of the parasite. Several terms have been used to describe the range of hosts of a para- site: monoxenous for species which are restricted to a single host; oligo- xenous for species which have moderate specificity (Rogers, loc. cit.) ; and polyxenous for species with low specificity. The term oligoxenous has been applied both to species that normally occur on two or more unrelated hosts and to those which occur on closely related ones. Perhaps the term stenoxenous should be used for the latter. Thus, Trichobius joblingi, Speiseria ambigua, and Strebla carolliae, which are characteristic parasites of species of Carollia, would be stenoxenous, while Trichobius dugesioides, which is a characteristic (see below) parasite of Trachops cirrhosus but also occurs on Carollia p. azteca and possibly C. subrufa, would be oligo- xenous (see "Host specificity," below). Oligoxenous and polyxenous and perhaps even stenoxenous species would in most instances probably have preferred hosts, but a preferred host may not be the most suitable host, i.e., the host on which the parasite has 12 "The first is specificity shown by the direct observation of natural phenomena; the second that which results from experiment." (Caullery, loc. cit.) WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 637 the greatest reproductive success (see Haas, 1965). Given sufficient time and opportunity for adaptation by the parasite, the most suitable host or hosts would in most instances become the preferred host or hosts in a given geographic area. Except in rare instances, use of the term "preferred" for a host has been inferential and not based on extensive observation or ex- perimental evidence. We have used the terms principal or primary to indi- cate the host to which the largest number of records apply. It may or may not be the most suitable host or the preferred host but may simply reflect the known records. Needless to say, these reflect the history and accidents of collecting, and relative abundance of hosts. When Jobling (1949) summarized what was known about the host- parasite relationships of New World Streblidae, he referred to Streblidae that normally occur on more than one host as being heteroxenous. We do not use this term because it is generally applied to parasites which com- plete their cycle of development only by passing through (or on) two or more (intermediate and definitive) successive hosts (see Caullery, op. cit., p. 127) . This term would properly apply to most ticks. We found it convenient to use Caullery's (op. cit.) distinction between normal and exceptional hosts. The latter would run the gamut from transitory or temporary hosts to hosts of very low preference. We would not include disturbance transfers and contaminations under this category. Similarly, we referred to parasites which regularly and consistently occur on a host (though not necessarily with a high degree of preference for it) as its characteristic or normal parasites, the others as exceptional parasites. Species of the same genus, which occur together on the same host species, we refer to as synoxenous, and those which occur on different hosts in the same or different geographic areas as alloxenous. Alloxenous species may be either sympatric or allopatric. We suggest that the terms host limitation and host-limited be used to refer to the extent to which the parasite, throughout its life cycle, is di- rectly dependent upon and associated with the host. Thus, holometabolous insects with free-living young, like fleas, exhibit a low degree of host limi- tation. Batflies, which are pupiparous and generally leave the host only long enough to pupiposit or mate, exhibit a high degree of host limitation. Anoplura and Mallophaga are host-limited, since all stages of the life cycle are directly dependent upon the host. Host Specificity Perhaps the greatest single need in determining the degree of host specificity in fact, is for sufficiently large and carefully made, geographically representative collections that enable one to statistically evaluate the frequency with which a parasite is found on a given host or hosts. The literature is replete with host-parasite lists which are not and cannot be evaluated. For groups like the Anoplura and the Mallophaga which are host-limited and exhibit an extraordinarily high degree of host specificity, the problem is of an entirely different order than for Siphonaptera which are much less host-limited, or for the Streblidae. The Streblidae are for 638 ECTOPARASITES OF PANAMA the most part winged and thus quite mobile. Further, their hosts tend to roost in association with other species. The number of exceptional associ- ations and of contaminations due to pooled collecting are relatively high, even for species which prove to be monoxenous. Thus, in the absence of quantitative data, it is difficult to determine the degree of host specificity, even if the parasite is monoxenous or oligoxenous, especially if their hosts are ubiquitous. For such hosts may not only acquire the parasites of other hosts, but their parasites in turn may transfer, with varying degrees of ease, to other hosts with which the normal host roosts (hosts whose ecological adjustments most nearly approach those of the normal host) or to hosts which are taxonomically related. The Streblidae of bats of the genus Carollia illustrate these points. In Panama, three species of Carollia were collected. Carollia perspicillata is one of the commonest and most widespread bats in the lowland tropics of Central and South America. It is tolerant of a wide variety of lowland ecological conditions. Because it is catholic in its selection of roosting sites, it comes to roost in association with a wide variety of bats. In Trini- dad alone, where the bat fauna (62 species) is somewhat limited by com- parison with the mainland, Goodwin and Greenhall (1961) recorded roost- ing associations of C. p. perspicillata with 23 other species of bats. The same ecological amplitude is shown by C. p. azteca in Panama. Carollia castanea, on the other hand, though a common lowland species, is more limited ecologically, and thus in its roosting sites, than C. p. azteca. Carollia subrufa is also an abundant bat in certain lowland localities, but again is more restricted ecologically than C. p. azteca. It also occurs at higher elevations than either azteca or castanea, especially above 3000 feet, where the number of species, of both bats and streblids, are sharply reduced. Thus, by comparison with azteca, the opportunities for parasite exchange between C. subrufa or C. castanea and other bats are greatly reduced. The records of Panamanian Streblidae reflect the differences in eco- logical amplitude of these hosts. Fifteen species were collected from C. p. azteca, four from C. subrufa, and three from C. castanea. Three — Speiseria ambigua, Strebla carolliae, and Trichobius joblingi — occurred on all three hosts. As can be seen from table 6 and fig. 141, they were the predominant flies on Carollia p. azteca. They were the only species on C. castanea. These data suggest that they are characteristic or normal parasites of bats of the genus Carollia, possibly of all Carolliinae, and that other species (all of which show high numerical values for other hosts) are either exceptional parasites, or represent contaminations or errors. If this is true, then these three species should occur only infrequently, as exceptional parasites or contaminations, on hosts other than Carollia. This is indeed the case. Conversely, this apparently is also true for the 12 other species reported from Carollia. Nearly all of them are from hosts with which Carollia may roost, and they show the same high incidence of parasitization for their normal hosts, as do the species characteristic for Carollia on Carollia. The bar graphs in fig. 141 show the incidence of parasitization by streblid species, of Carollia p. azteca and seven hosts (among many) with which it WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 639 TABLE 6. STREBLIDAE FROM BATS OF THE GENUS CAROLLIA, ACCORDING TO SPECIES A B C D E Number Percent Percent of Number of Percent of of bats of bats streblid-in- Streblidae all Streblidae parasitized parasitized fested bats taken on host Carollia p. azteca 339 considered ; 284 (83.8%) parasitized by Streblidae Trichobius joblingi Speiseria ambigua Strebla carolliae Trichobius dugesioides Strebla altmani Trichobius yunkeri Metelasmus pseudopterus Aspidoptera busckii Trichobius macrophylli Aspidoptera delatorrei Mastoptera guimaraesi Nycterophilia parnelli Trichobius dugesii johnsonae " sparsus 282 99 100 38 19 3 3 2 83.18 29.20 29.49 11.20 5.60 0.88 0.58 n 0.29 99.29 34.85 35.21 13.38 6.69 1.05 0.70 0.35 1104 160 121 53 29 3 74.44 10.78 8.15 3.57 1.95 0.20 0.13 0.06 Carollia castanea 36 considered; 34 (99.4%) parasitized by Streblidae Trichobius joblingi Strebla carolliae Speiseria ambigua 34 6 4 94.44 16.66 11.11 100.00 17.64 11.76 78 6 6 86.66 6.66 6.66 Carollia subrufa 30 considered; 19 (63.3%) parasitized by Streblidae Trichobius joblingi Speiseria ambigua Strebla carolliae Trichobius dugesioides 18 4 1 1 60.00 13.33 3.33 3.33 94.73 21.05 5.26 5.26 66 4 1 1 91.66 5.55 1.38 1.38 commonly roosts, in Panama. The eight species of bats belong to the family Desmodidae and four subfamilies of Phyllostomidae. The implications need no further explanation. The very low incidences mostly represent transi- tory transfers, or other nonspecific associations, but some undoubtedly represent errors or contaminations due to collecting techniques. It is to be expected that a large percentage of exceptional species will be reported 640 ECTOPARASITES OF PANAMA II -s . . s-s- I I -s I I II I II -s I -s a 3 | £ "I i I -g u WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 641 from ubiquitous hosts, because of the more numerous opportunities for transfer of parasites from other bats. There will also be a larger number of contaminations and errors recorded for them, simply because more are collected. It will be noted that some of the parasites listed in fig. 141 (like Trichobius yunkeri, T. johnsonae, and Nycterophilia parnelli) are charac- teristic parasites of other hosts, like Pteronotus spp., which are not included in the figure. Two species of Streblidae, Strebla altmani and Trichobius dugesioides, which appear to be characteristic parasites of other hosts, occurred in sig- TABLE 7. INCIDENCE OF TRICHOBIUS JOBLINGI (j.) AND T. DUGESIOIDES (d.) ON CAROLLIA PERSPICILLATA AZTEC A AND TRACHOPS CIRRHOSUS CIRRHOSUS IN PANAMA. A B C D* Percent of Number of Percent (of Mean no. hosts flies all Streblidae of flies parasitized collected on host) per host j. d. ;. d. j. d. ;. d. 263 1.59 84.02 Trachops cirrhosus 55 bats considered; 54 (98.2%) para- sitized by Streblidae Carollia p. azteca 339 bats considered; 284 (83.77%) para- sitized by Streblidae *Mean no. per host = mean no. of each species per host bat parasitized by that species. 5.45 67.27 83.18 11.20 1104 121 74.44 3.57 0.09 3.87 4.69 0.18 nificant numbers on C. p. azteca, S. altmani appears to be characteristic of both Lonchorhina aurita and Macrophyllum macrophyllum, related genera of Phyllostominae. Although it was taken on 5.6% of the Carollia p. azteca considered, it apparently did not occur on Carollia in the absence of one of the normal hosts. T. dugesioides appears to represent a more complicated situation. It parasitized 67% of the specimens of Trachops cirrhosus examined (table 7) and constituted 84% of the Streblidae on that host, but it parasitized only 11.2 % of Carollia p. azteca and constituted only 3.57% of the Streblidae on that host. Records indicate that it is an ex- ceptional parasite of other hosts. Trachops cirrhosus and Carollia p. azteca often roost in the same sites. There appears to be very little transfer of T. joblingi from its normal host, C. p. azteca, to Trachops, while the reverse appears to be true for dugesioides from Trachops to Carollia p. azteca. One might assume from the data that the incidence of T. dugesioides on C. p. azteca is to be ex- plained by a simple transfer from Trachops, when the two hosts roost to- gether. This does not appear to be the case. Most of the specimens of 642 ECTOPARASITES OF PANAMA dugesioides that occurred on Carollia p. azteca were from bats that roosted in caves and a mine shaft, typical sites for Trachops cirrhosus. Interest- ingly, no Trachops were recorded from these collecting sites. On the other hand, quite a different situation prevailed among hosts collected from a culvert along Chepo Road. Here, a series of seven Trachops cirrhosus were infested with an average of 17 dugesioides per bat parasitized by that species, as opposed to an average of 4.69 for all localities. Seventeen Carollia p. azteca were collected in this same culvert. Not a single speci- men of T. dugesioides was taken from them ! The reasons are not apparent, but several possibilities are suggested. Fifteen of the 17 Carollia were females ; it may be that dugesioides is not attracted to females. However, in other localities where Carollia was parasitized by dugesioides, there was no observable difference in parasitization according to sex of the host. One may suggest, too, that transfer does not easily take place in an "unnatural" roosting site such as a culvert. Still another possibility is that this streblid has such a strong preference for Trachops that it will parasitize another host only if it emerges from the puparium in the absence of the normal host, for example, if the Trachops colony has moved to another roosting site. If this is the case, it would be most interesting to determine whether or not host-conditioning of these parasites takes place. Analyses of host associations, similar to those shown in table 6 were used to evaluate the relationships of most of the common hosts and their Streblidae. These were the bases for determining which flies are normal or characteristic for a host in the host-parasite list (p. 663) . This is a simple approach. Much more sophisticated statistical tech- niques will be needed for investigating many of these problems. For ex- ample, we have not taken into consideration the size of the sample. Data have been lumped to a considerable extent. We have eliminated from con- sideration mixed collections or pooled collections, or collections which we definitely know to have been contaminated. The columns indicate different indices that can be used as a guide in evaluating the degree of host speci- ficity. In general, it was our experience that if the values in Columns B, C, and E were below 1%, the records represented contaminations and errors in labeling, or disturbance transfers. Slightly higher values appeared to represent disturbance transfers or transitory associations of a non-specific nature. In most instances normal associations, including those of low host preference, gave higher values. In general, each of the figures appears to be a valid index as to the de- gree of specificity. In some instances, the percentage for each species of the total number of Streblidae collected from the host, gave a better ap- preciation of the relationship to the host. In others, especially when popu- lations of Streblidae on the host were small, the figures in Columns B and C gave a clearer picture. In instances where the host is parasitized only under special circumstances, the figure in Column C was more instructive. For example, Strebla alvarezi was taken on species of Micronycteris and on Saccopteryx bilineata, two genera of bats which often roost in close prox- imity (seep. 627). WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 643 The incidence of parasitization of these hosts by S. alvarezi (table 9) was very low. It was only 12.09% for the 124 specimens of Micronycteris megalotis collected, and even lower for Saccopteryx bilineata. The figure for Trichobius keenani, a species known to us only from species of Micro- nycteris, is even lower. Thus, the figures given in Column B for the over-all incidence of S. alvarezi could be interpreted to mean that megalotis is an exceptional host of this fly. However, Micronycteris species roost in small groups, usually in relatively exposed situations (see pp. 627 and 647), con- ditions probably not suitable for the maintenance of large populations of Streblidae. If this is true, then bats from many roosting sites would be negative for Streblidae, even though the host might elsewhere be parasitized by species that are restricted to it, or to it and to closely related species. It would seem, then, that a more significant indicator of the host relation- ship would be a figure which at least to some degree takes this into account, e.g., one which shows the incidence of each species on those bats that were parasitized by Streblidae. Further, this eliminates from consideration host specimens which for one reason or another were not examined, or were not etherized and examined promptly enough to recover the Streblidae. The figures in Column C, then, more nearly approximate those in E and probably give a truer picture of the specificity, though not of the incidence of parasitization within the total population of the host (see p. 647) . We are in need of acceptable indices of host specificity that take into account the variables mentioned. Their development and use would be facili- tated by the use of modern data processing techniques. Analysis of the field data for all of the Streblidae indicates that in Panama about 55 % of the species are known only from a single host ; an- other 15% appear to be monoxenous, but occasionally occurred as excep- tional parasites on other hosts; about 13.5% were stenoxenous, i.e., they occurred on bats of a single genus; and 15% were oligoxenous and nor- mally occurred on bats of two or more genera. A few are unassignable. None were polyxenous. This is in closer agreement with the conclusions of Ross (1961) than of Jobling (1946), though Ross dealt with only a few species, from a rela- tively limited area. Jobling concluded that only nine of the 36 species of New World Streblidae recognized by him were monoxenous, and that the rest were "heteroxenous," i.e., they normally occurred on more than one species of bat. His conclusions were the natural consequence of having to deal with unreliable host data and the limited collections available at the time. This made it nearly impossible to discriminate closely related species. His interesting ideas on the relationships between the roosting habits of the hosts and the Streblidae they harbored are generally valid, but his statement that a very large number of the rare forest bats are free of Streblidae is not entirely true. Also contrary to his belief, some species of Streblidae, e.g., Trichobius macrophylli and members of the Trichobius caecus group, may regularly occur in large numbers on the host, while others may do so in exceptional cases. 644 ECTOPARASITES OF PANAMA Although most of the Panamanian Streblidae appear to be monoxenous or stenoxenous, a preliminary study of extensive collections from Mexico indicates that toward the edge of the distribution of the family, the species tend to be less host specific, at least those that occur on cave bats. Seasonal movements, including migration, appear to be more common among bats of the temperate zone. It seems probable that cave Streblidae would have a better chance of maintaining themselves throughout the year, if they could live on whatever host was predominant at a given time. In other in- stances, essentially tropical hosts living near the periphery of their range in ecologically "marginal" situations, would probably be less uniformly distributed and less abundant. Other things being equal, ecological poly- valence (including host preference) would presumably be advantageous to parasites of such hosts. This would agree with findings in some other groups of ectoparasites, as well. Parasite Faunules Because of the high degree of host specificity, it is possible, in most cases, to identify a host by its Streblidae. This is true even when the para- sites are not strictly monoxenous, because the composition of the streblid faunule of a host is distinctive. For example, Strebla carolliae, Trichobius joblingi, and Speiseria ambigua constitute a faunule that is characteristic of bats of the genus Carollia. If Trichobius dugesioides is also present, the faunule is characteristic of Carollia perspicillata azteca and possibly of C. subrufa. Similarly, Trichobius costalimai, Trichobioides perspicillatus and Strebla hertigi form a characteristic faunule of Phyllostomus discolor, both P. d. discolor and P. d. verrucosus. The streblid faunule may differ between "subspecies" of the host. For example, Trichobius longipes, Mastoptera minuta, and Strebla consocius constitute the faunule on Phyllostomus hastatus hastatus, at least in north- ern South America, while T. longipes, Mastoptera guimaraesi and Strebla mirabilis constitute the faunule on P. h. panamensis in northwestern South America and Panama. In the lowlands of Panama (and possibly Colom- bia), this faunule may include Strebla hertigi, as well. The faunule may also vary geographically in the same subspecies. For example, in Costa Rica and Nicaragua, Strebla mirabilis apparently disappears from P. h. hastatus and is entirely replaced by S. hertigi. In Panama, the number of characteristic species per host varied from one to five. Not all of them necessarily occur on each host bat or in all roosting sites or localities. Below, we have listed the number of host species col- lected in Panama that fell into each category. No. of characteristic streblid species per host': 012345 No. of host species: 51 14 21 7 6 1 These figures will change with further collecting. For example, it will probably be found that fewer than 14 hosts harbor only a single species, and that some of the hosts which were negative do have Streblidae. Undoubtedly, many factors determine the nature of the streblid faunule on any one bat, including evolutionary history of host and parasite, physical WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 645 and biotic factors of the environment (including the host), biology of host and parasite, and size of host and parasite, to list a few. In general, the largest number of species of Streblidae are recorded from hosts that are ecologically labile, especially ubiquitous hosts and/or hosts that tend to roost in sizable (sometimes very large) colonies. Among such hosts in Panama are Phyllostomus h. panamensis, Carollia perspicillata azteca, Artibeus jamaicensls, and Pteronotus parnellii fuscus. On the other hand, species of the genus Tonatia, which may harbor four or more species of Streblidae each, live in separate family groups ! Carollia p. azteca had the largest number of characteristic species of any host bat in Panama. Synoxenous Parasites Although a little more than 70 % of the host species collected harbored two or more species of Streblidae, and 35 % harbored three or more, it is only rarely that an individual host bat normally harbors two species of the same genus simultaneously. When this is so, the flies are usually morpho- logically different types, presumably specialized for living on different body regions of the host. In those rare instances in which this does not appear to be the case, one of the two species appears to be secondarily acquired from an ecologically and/or taxonomically related host, and most if not all of these exchanges appear to be unilateral or non-reciprocal. We have already noted that Trichobius joblingi, a normal parasite of Carollia, and T. dugesioides, a characteristic parasite of Trachops (and possibly of Chrotopterus) , occur together on Carollia p. azteca and, some- times on C. subrufa. On the other hand (see above), joblingi rarely trans- fers from Carollia to Trachops. Likewise, Strebla altmani may occur to- gether with S. carolliae on Carollia p. azteca but only when the latter is associated with Lonchorhina aurita, one of the normal hosts of altmani; but S. carolliae, the characteristic Strebla of Carollia, rarely transfers to Lonchorhina under the same circumstances. Trichobius johnsonae is a characteristic parasite of Pteronotus par- nellii fuscus, and T. yunkeri of Pteronotus suapurensis and P. psilotis. In one instance, 27 specimens of johnsonae and 26 of yunkeri were recorded on a single specimen of P. suapurensis. It cannot be determined from the field data whether or not P. p. fuscus was also present. It may be that the collection was not taken from a single host species or individual, as represented. Trichobius sparsus normally occurs in small numbers together with T. johnsonae on Pteronotus parnellii fuscus. However, these two species of Trichobius belong to different groups and probably are specialized in their feeding habits. The only other synoxenous species of New World Streblidae that we know are the two species of Strebla, mirabilis and hertigi, that occur to- gether on P. hastatus panamensis in Panama, and possibly Colombia. S. mirabilis typically occurs in larger numbers (see following paper, table 11 and fig. 147) as the "dominant" species, and hertigi in smaller numbers as the "subordinate" species. However, in Costa Rica and Nicaragua, mira- 646 ECTOPARASITES OF PANAMA TABLE 8. INCIDENCE* OF FOUR SPECIES OF STREBLIDAE ON PHYLLOSTOMUS HASTATUS PANAMENSIS COLLECTED AT FIVE LOCALITIES IN PANAMA. Localities Chepo Road (Panama) (hollow tree in forest) 8 Oct. (16 bats) Chilibrillo Caves (Panama) July & Oct. (17 bats) Madden Dam (Canal Zone) (natural bridge and small cave. 28-29 Sept. (14 bats) Fort Kobbe (Canal Zone) (hollow Ficus tree) 9 Oct. (5 bats) Fort Sherman (Canal Zone) (buildings & bunkers) 4 & 6 April (19 bats) Strebla mirabilis 100.00(200) 76.57(206) 92.85(170) 100.00(77) 100.00(146) Strebla hertigi 87.50(40) 5.38(1) 14.28(1) 0.00 5.26(1) Trichobius longipes 100.00(86) 70.58(45) 28.57(47) 100.00(35) Mastoptera guimaraesi 93.75(113) 5.88(1) 7.13(1) 0.00 47.36(43) 42.10(77) * Shown as percent of hosts parasitized by each species. Figures in parentheses = number of specimens of each species of streblid. bilis does not seem to occur on P. h. panamensis but is entirely replaced on that host by S. hertigi. The limited data suggest that competitive displace- ment is involved. The problem is discussed in the following paper. Ecology of Parasitization Little is known about the population ecology and dynamics of Streb- lidae, or their relation to parasitization of the bat hosts. However, as was pointed out (p. 426), population densities may be high for Streblidae of bats like Pteronotus, which roost in relatively large colonies, usually in long established sites, and low (or the parasites absent) in new or temporary sites. In general, too, as Jobling (1949b) indicated, Streblidae tend to be absent from those forest bats which roost in small numbers in exposed situations. Thus, species of Emballonuridae, in general, have few or no Streblidae, or possibly acquire them (see p. 627) from other ecologically associated bats whose roosting sites may be slightly more favorable for Streblidae. Bats like Thyroptera, which roost singly or in very small num- bers in rolled leaves of Heliconia and bananas, lack these flies. Their roosting habits probably render them unsuitable as hosts for Streblidae. However, they may have Nycteribiidae of the genus Hershkovitzia™ This 13 The Nycteribiidae appear better able to maintain themselves on hosts that roost apart from others, in relatively restricted sites and in small numbers. As a family, they also have a broader climatic tolerance and distribution than do Streblidae. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 647 may be the situation, too, in Anthorhina and Mimon, the only Phyllostomidae known to us that normally harbor Nycteribiidae. Closely related phyl- lostomine genera are parasitized by Streblidae and have distinctive fau- nules. Even bats of the genus Tonatia, which roost in family groups, have well-developed streblid faunules. However, these bats usually roost in the interior of Nasutitermes nests which had been hollowed out and used as nests by parrots, and then abandoned. In such an enclosed "home," it is likely that the microclimate and other conditions are suitable for Streblidae. Further, the Streblidae would be relatively restricted, to both bat and roosting site, and the hazards of being separated from the host would be correspondingly reduced. In bats whose roosting sites and habits fall between the extremes repre- sented by Pteronotus and Thyroptera, one would expect to find considerable variance in both the incidence and density of Streblidae, especially if the host roosts in a wide variety of situations. Phyllostomus hastatus pana- mensis appears to be such a host. Table 8 shows the incidence of the four characteristic species of Streblidae of that host, based on collections made at five Panamanian localities. Forest situations appear to be most suitable for parasitization of this host by S. hertigi and it is probable that this fly transfers to it from Phyllostomus d. discolor in these habitats (see follow- ing paper, by Wenzel and Tipton) . Caves appeared to be less favorable sites than the others, for all of the Streblidae of P. hastatus panamensis. This is interesting, because the genera and species groups of Streblidae repre- sented on this host are characteristic of Phyllostominae, bats which as a group tend to be forest dwellers. One can only guess at the factors that completely exclude Mastoptera guimaraesi from P. h. panamensis in a Ficus tree, when this fly was present in large numbers on the same host roosting in a building, and also para- sitized 100% of the bats in a hollow espave tree in a forest along Chepo Road. Further, at Fort Sherman, only four guimaraesi were recovered from 13 bats collected in one building, whereas 73 were taken from six bats collected in two others ! Similarly, only 14.57^ of 124 specimens of Micronycteris megalotis col- lected at 27 sites throughout Panama, were parasitized by Streblidae (table 9). Yet, in an area of a few square miles near the Empire Range (Canal Zone), the senior author and Charles M. Keenan found that 64.7 % of them were infested. A thousand yards (or less) away, along the road to Cocoli, the bats were equally as numerous but all were negative for these flies. More bats were collected at these sites than for which data are given, but the differences are nonetheless evident. The area in which the parasitized bats occurred included a considerable amount of forest in which there were logs and large espave and other trees with basal tree holes, both suitable roosting sites for this bat. In the area along the road to Cocoli, where no parasitized bats could be found, the forest was mostly young "second growth." No tree holes were seen there, and all of the bats taken were roosting in culverts and similar man-made sites. As pointed out (p. 627), Micronycteris megalotis roosts in small num.- 648 ECTOPARASITES OF PANAMA TABLE 9. INCIDENCE OF STREBLIDAE ON MICRONYCTERIS MEGALOTIS MICROTIS IN PANAMA A B Number of Percent of megalotis megalotis parasitized parasitized Empire Range Area 6 sites : 17 bats Strebla alvarezi Trichobius keenani Total, excluding Empire Range Area 21 sites : 107 bats Strebla alvarezi Trichobius keenani Total localities 27 sites: 124 bats Strebla alvarezi Trichobius keenani 11 10 3 7 5 2 18 15 5 64.70 58.82 17.64 6.54 4.67 1.86 14.51 12.09 4.03 C* Percent of parasitized megalotis 90.90 27.27 71.42 28.57 83.33 27.77 D E Number of Percent of Streblidae all taken Streblidae on host 15 8 20 13 65.21 34.78 50.00 50.00 60.60 39.39 *Percent of those megalotis parasitized by Streblidae that were parasitized by S. alvarezi or T. keenani, respectively. bers, in relatively exposed situations. In natural habitats, it roosts just inside tree holes and hollow logs, but in habitats disturbed by man, it also roosts in culverts, buildings, and similar sites. In the Empire Range area, parasitized individuals were taken in all of these roosts. We think it prob- able that the parasites of this bat can establish and maintain themselves only in relatively undisturbed areas, where the host populations are able to roost in natural and relatively stable sites such as tree holes, and where populations of the host have maintained themselves over a considerable period of time. This would explain the absence of the parasites in such heavily disturbed areas as the one outside the Empire Range area near Cocoli. Extensive and intensive collecting, with appropriate ecological observa- tions, population estimates of the hosts, and other data, will provide valu- able insights, but autecological studies of hosts and parasites are essential to an understanding of the problems of parasitization. Taxonomic Concordance It is premature to attempt a detailed study of the concordance between the classification of the Streblidae and that of their hosts. The taxonomy and morphology of the Streblidae are too imperfectly known to permit one to delineate probable phyletic relationships within the family. Still to be investigated are puparial characters, internal morphology, morphology of the reproductive apparatus, and chromosome karyotypes, among others. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 649 Nonetheless, a tentative grouping (table 10) of the genera and species groups of the New World Streblidae, based largely on structure of the abdomen and head, reveals a number of interesting correlations with the classification of the hosts. In the Trichobius pallidus, caecus, and major groups, the laterovertices and occipital lobes or plates of the head are poorly differentiated. They are better differentiated, but still poorly sclerotized, in Joblingia and Anatrichobius. They are well sclerotized in other genera. In Series 3, they are well differentiated, and in Series 4 (Streblinae), they reach their maximum development and differentiation. One may question the validity of these groupings, since any of the characters on which they are based could evolve independently. We tenta- tively regard the insertion of the accessory setae posterior to the macrosetae on the male gonapophyses as a generalized condition, and the free cerci of the female as a derived condition. In some instances, the gonapophyseal setae are poorly differentiated ; and it cannot be determined which, if any, are accessory and which are macrosetae. Likewise, it is sometimes very diffi- cult to determine whether the female cerci are free or not. It will be nec- essary to further investigate these and many other characters. However, with the exception of Series 3, the series seem to be internally consistent, even though the inter and intra series relationships are not clear. If the groupings have validity, then it appears that Trichobius is not a homogeneous genus. This is certainly true. The genus is sufficiently poly- morphic that one can justify creating a series of separate genera or sub- genera for most of the Trichobius groups and perhaps even further subdivide the major group. Some of these appear to be ancestral to dis- tinctive evolutionary lines. For example, it is clear that Joblingia and Anatrichobius are brachypterous derivatives of the major-corynorhini subgroup of the major group of Trichobius. It is also reasonably clear that Megistopoda, Paratrichobius and Neotrichobius evolved from species of, or closely related to, the Trichobius phyllostomae group; that the dugesii group is closely related to and probably derived from the longipes group; that the genus Trichobioides evolved from a species of, or related to the dugesii group ; and further that the brachypterous Mastoptera prob- ably evolved from a species related to Trichobioides perspicillatus. In their male and in some female characters, the Streblinae appear similar to the caecus group of Trichobius. However, they are so highly spe- cialized in their head characters, and differ sufficiently in most female char- acters that, at present, we cannot demonstrate a relationship. Within the Trichobiinae, the least specialized streblids appear to be the pallidus, caecus and major groups of Trichobius. The male of Synthesiostrebla has not been examined. The general morphology and female genital structure sug- gest that the genus may be related to the pallidus group. The female of Stizostrebla longirostris has not been studied, either, but its morphology suggests that it is closely related to Pseudostrebla. Super- ficially, Parastrebla handleyi appears very similar to Pseudostrebla, but in its female genital structure is very similar to members of the Trichobius 650 ECTOPARASITES OF PANAMA TABLE 10. TENTATIVE GROUPING OF NEW WORLD STREBLIDAE. (* •= recorded from Panama. ? = assignment dubious.) DIVISION I Subfamily NYCTEROPHILIINAE. Form flea- like. Abdominal terga I and II distinctly separated lateroventrally, fused dorsally; sterna I-VI of male represented by nor- mally developed, similar sclerites. Male genital apparatus external. Nycterophilia Ferris, p. 432 coxata Ferris, p. 434 *fairchildi n.sp., p. 436 *natali n.sp., p. 438 *parnelli n.sp., p. 434 DIVISION II Subfamilies TRICHOBIINAE, STREBLINAE. Form not flea-like. Abdominal terga I and II completely fused ; male sterna dissimilar, III and IV absent, V and VI often absent. Male genital apparatus internal. SERIES 1 (TRICHOBIINAE). Male. — Ac- cessory gonapophyseal setae inserted pos- terior to the macrosetae. Female. — Ab- dominal cerci articulated or fused with ventral arc; postgenital sclerite usually present. Trichobius Gervais, p. 442 pallidus group, p. 447 pallidus (Curran), p. 447 caecus group, p. 448 caecus Edwards, p. 450 *galei n.sp., p. 449 *johnsonae n.sp., p. 455 *yunkeri n.sp., p. 453 ? Synthesiostrebla Townsend, p. 429 amorphochili Townsend, p. 429 Trichobius Gervais major group, p. 456 ladamsi Augustson, 1943 corynorhini Cockerell, 1910 hirsutulus Bequaert, 1933 major Coquillet, 1899 pseudotruncatus Jobling, 1939a (= kesselae Jobling, 1938) *sparsus Kessel, p. 457 sphaeronotus Jobling, 1939a truncatus Kessel, 1925 Joblingia Dybas & Wenzel, p. 507 *schmidti Dybas & Wenzel, p. 507 Anatrichobius n.g., p. 502 *scorzai n.sp., p. 503 ?Parastrebla n.g., p. 578 *handleyi n.sp., p. 579 Trichobius Gervais phyllostomae group, p. 496 *brennani n.sp., p. 497 phyllostomae Kessel, p. 498 *vampyropis n.sp., p. 500 Megistopoda Macquart, p. 540 (= Pterellipsis Coquillet, 1899) *aranea (Coquillet), p. 542 pilatei Macquart, p. 541 *proxima (Seguy), p. 543 *theodori n.sp., p. 545 Paratrichobius Costa Lima, p. 518 *dunni (Curran), p. 527 *longicrus (M. Ribeiro), p. 521 *lowei n.sp., p. 528 *sanchezi n.sp., p. 530 *salvini n.sp., p. 532 Neotrichobius n.g., p. 536 *stenopterus n.sp., p. 539 SERIES 2 (TRICHOBIINAE). Male.— Ac- cessory gonapophyseal setae inserted an- terior to macrosetae. Female. — Cerci united with ventral arc; postgenital scle- rite absent except in Speiseria. Trichobius Gervais uniformis group, p. 459 *keenani n.sp., p. 462 *lionycteridis n.sp., p. 464 *lonchophyllae n.sp., p. 461 *uniformis Curran, p. 459 Speiseria Kessel, p. 549 *ambigua Kessel, p. 549 WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 651 TABLE 10 (continued) SERIES 3 (TRICHOBIINAE). Head without a complete ventral ctenidium. Male. — Ac- cessory gonapophyseal setae inserted an- terior to the macrosetae. Female. — Cerci free; postgenital sclerite absent. Noctiliostrebla n.g., p. 560 aitkeni n.sp., p. 567 *maai n.sp., p. 569 dubia (Rudow), p. 563 megastigma (Speiser), p. 564 *traubi n.sp., p. 565 Paradyschiria Speiser, p. 571 fusca Speiser, p. 573 *lineata Kessel, p. 574 parvula Falcoz, p. 574 *parvuloides n.sp., p. 575 ***** Trichobius Gervais longipes group, p. 465 *bequaerti n.sp., p. 471 *costalimai Guimaraes, p. 471 *dunni n.sp., p. 474 *dybasi n.sp., p. 469 *longipes (Rudow), p. 465 ( = mixtus Curran) *mendezi n.sp., p. 469 dugesii group, p. 476 diphyllae n.sp., p. 492 *dugesii Townsend, p. 478 *dugesioides n.sp., p. 488 furmani n.sp., p. 490 *joblingi n.sp., p. 481 *macrophylli n.sp., p. 486 *parasiticus Gervais, p. 494 *urodermae n.sp., p. 476 Trichobiodes n.g., p. 510 *perspicillatus (Pessoa & Galvao), p. 511 Mastoptera n.g., p. 512 *guimaraesi n.sp., p. 514 *minuta (Costa Lima), p. 515 Exastinion (Pessoa & Guimaraes) , p. 558 *clovisi (Pessoa & Guimaraes), p. 558 Aspidoptera Coquillet, p. 551 *busckii Coquillet, p. 555 *delatorrei n.sp., p. 557 phyllostomatis (Perty), p. 553 Pseudostrebla Costa Lima, p. 582 *greenwelli n.sp., p. 582 *ribeiroi Costa Lima, p. 582 ? Stizostrebla Jobling, p. 586 longirostris Jobling, p. 586 Eldunnia Curran, p. 586 *breviceps Curran, p. 588 SERIES 4 (STREBLINAB). Head with a complete ventral ctenidium. Male. — Ac- cessory gonapophyseal setae inserted pos- terior to the macrosetae. Female. — Cerci free; postgenital sclerite usually absent. Strebla Wiedemann, p. 591 *altmani n.sp., p. 623 *alvarezi n.sp., p. 625 *carolliae n.sp., p. 619 *christinae n.sp., p. 606 consocius n.sp., p. 600 *diaemi n.sp., p. 599 diphyllae n.sp., p. 613 *galindoi n.sp., p. 604 *hertigi n.sp., p. 596 *hoogstraali n.sp., p. 603 *kohlsi n.sp., p. 618 machadoi n.sp., p. 607 mexicana Rondani, p. 610 *mirabilis (Waterhouse), p. 615 tonatiae (Kessel), p. 602 * vesper tilionis (Fabricius), p. 609 Paraeuctenodes Pessoa & Guimaraes, p. 627 longipes Pessoa & Guimaraes, p. 627 Anastrebla n.g., p. 627 (— Strebla of authors, not Wiedemann, 1824) *mattadeni n.sp., p. 631 *modestini n.sp., p. 629 *nycteridis n.sp., p. 629 Metelasmus Coquillet, p. 634 *pseudopterus Coquillet, p. 634 The species arrangement is alphabetical. The page on which treated or cited is given for each genus and species. The year of publication is added for species not cited in the text. 652 ECTOPARASITES OF PANAMA major group, especially T. truncatus. The male of Parastrebla is unknown. The genital apparatus of Paradyschiria and Noctiliostrebla appears to be less specialized than in the other Trichobiinae, though it may have un- dergone simplification. Until they can be studied in greater detail, these two genera are placed in Series 3 with some reservations. In female genital structure (see pp. 519, 549), Speiseria is clearly re- lated to the Trichobius phyllostomae group and related genera (Series 1). However, it differs from them in the arrangement of the paired (male) gonapophyseal setae. It can hardly be regarded as related to the Trichobius uniformis group, with which it is placed here. The disposition of its gona- pophyseal setae may have evolved independently of other groups that have the same arrangement. Because the Streblidae exhibit a high degree of host-limitation and specificity, the groups of Streblidae proposed above, should, if they have any validity, show some concordance with the host taxa. A number of them do, as will be shown below. In figures 142-146, characteristic host associations were used. Division I. NYCTEROPHILIINAE Of seven species (four described) of Nycterophilia, five occur on Chilonycterinae and Natalidae (fig. 142) ,u An undescribed species, closely related to one that occurs on Mormoops (Chilonycterinae) has been taken from Platalina (Glossophaginae). N. coxata occurs on Macrotus (Phyl- lostominae), as does a related undescribed. species from the West Indies. All the hosts of Nycterophilia are primarily cave inhabitants. This would facilitate transfer of these flies, both on a temporary and an evolutionary basis, from Chilonycterinae and Natalidae to other cave-inhabiting bats, such as Leptonycteris and Macrotus. Division II. TRICHOBIINAE and STREBLINAE Series 1. Two species (one undescribed) of the Trichobius pallidus group (fig. 142) occur on bats of the family Furipteridae. The species of the caecus group are closely related to them but differ in being somewhat more specialized. Their eyes are reduced to a single facet, the median thoracic suture is reduced to a short fork, and the first tarsomere possesses a spe- cialized comb scale. There are several undescribed species from Natalus and Pteronotus but these have not been studied in enough detail to include in the figure. The host distributions of these groups of Trichobius and of Nycterophilia support the view of Dalquest and Werner (1954) and de la Torre (1962) that the Chilonycterinae should be regarded as a separate family of 14 The dendrogram of presumed relationships of the families and subfamilies of New World Chiroptera in figures 142, 143, and 145, is based in part upon a sketch provided by Dr. Karl Koopman. We have inserted the Emballonuridae and Vespertilionidae, not included by Dr. Koopman and have changed the position of the Desmodidae. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 653 Stenoderminae sturnirinae DESMODIDAE Glossophaginae NATALIDAE o • MOLOSSIDAE VESPERTILIONIDAE Carolliinae Phyllonycterinae Chilonycterinae oo® Trichobius caecus group Fig. 142. Dendrogram of families and subfamilies of New World bats, showing host distribution of the species of Nycterophilia and of the Trichobius pallidus and caecus groups. Unless otherwise indicated each symbol represents a separate species, (u = undescribed species. Vespertilionoidea rather than as a subfamily of Phyllostomidae. On the basis of facial histology (the presence of certain glands), Dalquest and Werner (op. cit.) placed them nearer the Vespertilionidae than to the Furipteridae and Natalidae. De la Torre (unpub. thesis) states that "In skull structure and general morphology the chilonycterid group appears to be much closer to the family Natalidae and to other related genera (Furip- terus, Amorphochilus, and Thyroptera) than to the phyllostomid group." This view is more consonant with the streblid distribution than is that of Dalquest and Werner. The Trichobius major group, and the related brachypterous genera 654 ECTOPARASITES OF PANAMA Stenoderminae Sturnirinae DESMODIDAE Glossophaginae NATALIDAE VESPERTILIONIDAE ooooo o o • Phyllonycterinae Chilonycterinae O NOCTILIONIDAE EMBAUONURIDAE Trichofaius major group Anatrichobius Joblingia Fig. 143. Dendrogram of families and subfamilies of New World bats, showing host distribution of the species of the Trichobius major group and of the derived genera, Joblingia and Anatrichobius (Series 1). Each symbol represents a separate species. Joblingia and Anatrichobius (fig. 143), occur primarily on cave-dwelling Vespertilionidae, with the exception of a few more specialized forms which occur on other cave bats. These include T. sparsus on Pteronotus parnellii (Chilonycterinae) and T. sphaeronotus on Leptonycteris nivalis (Glosso- phaginae). Trichobius adamsi on Macrotus (Phyllostominae), is doubt- fully included in this group. The species with the most generalized female genital structure are primarily warm temperate North American and subtropical in distribution, either latitudinally or altitudinally. Joblingia has been taken only at altitudes above 5500 feet in Panama, Costa Rica, and Guatemala. Anatrichobius has been taken at similar elevations in tropical South America and at lower elevations in the southern latitudes. WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 655 The Streblidae treated thus far appear to be the most generalized in genital and abdominal structure of all the New World Streblidae. It is in- teresting to note that they occur chiefly on Vespertilionoidea, which, like the Emballonuroidea, are generally regarded as among the most generalized Microchiroptera. Further, these flies occur chiefly on cave bats. It has been suggested (Jobling, 1949b; Theodor, 1957) in regard to the origin of batflies, that they evolved from guano-inhabiting flies of caves. Few New World Streblidae occur on Emballonuroidea, with the exception of Para- dyschiria and Noctiliostrebla on the Noctilionidae, and several species on Emballonuridae. This is probably because most New World Embal- lonuridae roost in very small numbers, in exposed situations. Those on the Emballonuridae are related to forms that occur on Phyllostomidae, and at least one host (see p. 627), shares a species with ecologically associated phyllostomine bats. The Trichobius phyllostomae group and the presumably derived genera, are also placed in Series 1, but do not appear to be closely related to the other streblids of that series. All of the described species 13 occur on genera that de la Torre (op. cit.) places as two compact and related groups of Stenoderminae (fig. 144). 16 Their host distribution not only indicates that these Streblidae are closely related and have primarily radiated on the Stenoderminae, but also supports de la Torre's grouping of these hosts. Of special interest is the distribution of the four species of the Trichobius phyllostomae group on Stumira, Vampyrops, and Artibeus, and similarly, of the Megistopoda species on Sturnira and Artibeus. Aspi- doptera, which belongs to Series 3, is similarly restricted to Sturnira, Artibeus (lituratus and jamaicensis) and possibly Chiroderma. Since Sturnira apparently does not roost with these bats, though they have been observed feeding with them (de la Torre, pers. comm.), the parallel oc- currence of closely related species of three genera of Streblidae on bats of the genera Sturnira and Artibeus, and of a species of one of these genera on Vampyrops, gives credence to de la Torre's contention (op. cit.) that the Sturnirinae should be placed with the Stenoderminae and not retained as a separate subfamily. Series 2. The species of the uniformis group are very closely related. Three of the four occur on three genera of Glossophaginae (fig. 144), re- spectively, and the fourth (keenani) on species of Micronycteris. In Pan- ama, Micronycteris megalotis, one of the hosts of keenani, was frequently found roosting with Glossophaga soricina, and it is possible that such an ecological association may, historically, explain this distribution of the flies. 13 An undescribed species of Paratrichobius has been taken from Choeronycteris mexicana in Arizona. 18 Dr. de la Torre has added several genera which he inadvertently omitted from his dendrogram, which appeared as fig. 4 in his unpublished thesis (1961), and has kindly given us permission to reproduce it. We have used the same dendrogram, with modifica- tions, in fig. 146. 656 ECTOPARASITES OF PANAMA Series Trichobius unKormis group Speiseria ambigua Trichobius phyllostomae group Paratrichobius Neotrichobius stenopterus Megistopoda Vampyrum Barticonycteris Micronycteris Macrotus Chrotopterus Tonatia Trachops Lonchorhina Macrophyllum Phylloderma Mimon Anthorhina Phyllostomus j Lionycteris '" Glossophaga Lonchophylla Platalina Lonchoglossa Monophyllus Anoura Hylonycteris Lychonycteris Leptonycteris Musonycteris Choeronycteris -Phyllostominae ^Glossophaginae J Choeroniscus Erophylla Phyllonycterisj Carollia L Rhinophylla J Brachyphylla ~1 Chiroderma Vampyriscus Vampyressa Mesophylla Vampyrodes Sturnira Vampyrops Enchisthenes Uroderma Artibeus Pygoderma Ardops Phyllops Ariteus Stenoderma Centurio Sphaeronycteris Ametrida f-Phyllonycterinae Carolliinae -Stenoderminae Fig. 144. Dendrogram of relationships of genera of phyllostomid bats, showing host as- sociations of Streblidae of the Trichobius phyllostomae group and related genera, and of the T. uniformis group. (1 = longicrus group, u = undescribed species.) WENZEL, TIPTON, AND KIEWLICZ : STREBLID BATFLIES 657 It should be noted that the three host genera of Glossophaginae belong to a cluster of genera that de la Torre considers to be closely related. The Streblidae on other genera of Glossophaginae are not at all closely related to the uniformis group. The genus Speiseria cannot be placed satisfactorily with either the uniformis group or the phyllostomae group and related genera, though it seems to be more closely related to the last-named. It is placed in Series 2, with the uniformis group, because of the arrangement of its accessory gonapophyseal setae. In all of its other characters, including the presence of a female postgenital sclerite, it is closer to the phyllostomae group and related genera. Although it appears to be primarily a parasite of species of Carollia, we have seen several collections taken from Vampyrops vittatus, one from Panama, the others from Guatemala. Interestingly, the problem of the relationships of Speiseria is much the same as that of the relation- ship of its principal hosts, the Carolliinae. Series 3. It should be noted again that the Streblidae of Series 1 and 2 — in which the female cerci are articulated or fused with the ventral arc — are largely restricted to or are characteristic parasites of non-phyllostomine bats. On the other hand, most of the genera of Series 3 are characteristic parasites of Phyllostominae, with the exception of Aspidoptera on Steno- derminae, Noctiliostrebla and Paradyschiria on Noctilionidae, and some species of the Trichobius longipes and dugesii groups. The two latter groups have scattered representatives on a miscellaneous assortment of hosts, probably derived through ecological association. These hosts in- clude Molossidae, Stenoderminae, Glossophaginae, Carolliinae, and Desmodi- dae, but apparently no Vespertilionoidea or Emballonuroidea. The distribution of the longipes and dugesii groups is shown in fig. 145. Several undescribed species are included. With the exception of T. dunni and two other closely related (undescribed) species from Molossidae, the members of the Trichobius longipes group (p. 465) are restricted to Phyl- lostomus and Tonatia, as are the species of Mastoptera. These are gen- erally regarded as closely related genera of bats, along with Mimon. No Streblidae, only Nycteribiidae, are known from Mimon. None are known from the genus Vampyrum, either. In the dugesii group, the species of the dugesii complex (p. 476) are found on Macrophyllum (Phyllostominae), Glossophaga (Glossophaginae) Carollia (Carolliinae), and Uroderma (Stenoderminae). Three unde- scribed species are known to us from Anoura and Choeronycteris (Glossoph- aginae) and Artibeus (Stenoderminae). Their close relationship with species of the Trichobius longipes group suggests that they are derived from them and have secondarily radiated onto both phylogenetically and ecolog- ically related bats. Of the species in the parasiticus complex (p. 487), T. dugesioides occurs on Phyllostominae (Trachops) and Carolliinae (probably secondarily, on Carollia), while three others, including an undescribed species from Diaemus, are parasites of Desmodidae. A fifth species, fur- mani, has also been taken on Desmodidae and on Glossophaga (Glossopha- 658 ECTOPARASITES OF PANAMA Stenoderminae 0,0 Sturnirinae Phyllonycterinae Glossophaginae 0000 U U f? Phyllostominae OO OO® O • S.consocius Fig. 147. Occurrence of species of Strebla on Phyllostomus hastatus and Phyllostomus discolor in Central America and northern South America, based on collections studied. of individuals of the two flies, the difference is even more striking. It will also be noted (table 11) that hertigi was absent or nearly so, from those colonies of P. h. panamensis that roosted in non-forest sites like caves and buildings. This suggests that this bat is a suitable host for hertigi only where ecological conditions of its roosting sites are similar to those of P. discolor. It further suggests that P. h. panamensis acquires hertigi in those situations where it can come into contact with discolor and /or its parasites, probably through roosting sites. From table 11 it will be seen that in the cases cited for Panama, the mean number of Strebla per host bat examined (column D) ranged from 7.7 to 15.4 for mirabilis and 0 to 2.5 for hertigi. If the mean number per bat is calculated for only those bats parasitized by each species (column E) the highest for hertigi is 2.85. It will be noted that the highest (mean) number of mirabilis per bat were taken from hosts on which no hertigi were found. In Costa Rica and Nicaragua, where P. h. panamensis reaches the northern limit of its distribution, the picture appears to be very different. Here, WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 685 fO w O e rQ 03 O ^5 6 "^ C3 SH d) o • 8 C 8 2 i- oS S w o 0 «H 8 T3 O "^ OJ ^ « O t^ HJ i— 1 ^ CO "o Csi • OS N « C in oq co' i o N ^, (Panam m 2 o 'd oS O T3 03 O> S« T) Canal Z 0) o> J-l •+J oo «^> o3 C 03 O e bunkers co 'ANAMA -= a 0 X o ft 0) 6 (Hollow tree Chilibrillo CE Madden Dam (Natural brl cave) Fort Kobbe ( (Hollow Fici Fort Sherma «B to &JO c is 'B M T3 O o 686 ECTOPARASITES OF PANAMA none of the specimens collected were parasitized by S. mirabilis ! The only species of Strebla present on that host was hertigi. All of the six host bats included in the sample from Nicaragua were parasitized by Streblidae, but only four (66%) by Strebla hertigi. On these four bats (column E), the number of hertigi per bat was 9.74 ! Thus, when mirabilis was absent, the numbers of hertigi per bat increased to a level nearly comparable to those of mirabilis on P. h. panamensis in Panama. We are not certain that Strebla mirabilis is absent from P. h. panamensis throughout Costa Rica and Nicaragua. Interestingly, a single collection from Sibube (Bocas del Toro), Panama (near Costa Rica) had three specimens of hertigi and two of mirabilis. One individual of P. h. pana- mensis from Armila (San Bias), had five hertigi and no other Streblidae. Why is S. mirabilis replaced by hertigi on Phyllostomus h. panamensis in Costa Rica and Nicaragua? Several answers may be suggested. The simplest is that toward the northern limit of its range, the roosting sites and/or population structure of the host are not as suitable for Strebla mirabilis as they are further south, nearer the epicenter of the host's range, 12 and thus mirabilis cannot maintain itself on this bat at these latitudes. Another possibility, which assumes that Phyllostomus discolor is the "reser- voir" of the hertigi that parasitize P. h. panamensis, is that the hertigi on P. discolor discolor differ from those on P. discolor verrucosus.13 If this is so, it may be that the population on P. discolor discolor is better adapted to P. h. panamensis than is S. mirabilis, and thus competitively displaces mirabilis on that host in Costa Rica and Nicaragua. The differences in the Strebla populations on P. h. panamensis do appear to coincide with the geographic separation of the two races of P. discolor (fig. 147) . However, if the single small collection from P. h. panamensis taken at Sibube is in- dicative, then there may be a geographic gradient in the ratio of mirabilis to hertigi on that host. This would lend support to the first explanation. Un- fortunately, nearly all of the Panamanian collections are from the Canal Zone and the Province of Panama. Extensive intermediate collections are necessary for an understanding of this problem. Whatever the explanation, the salient fact is that in Panama — where the two species of Strebla coexist on the same host — the mean number of hertigi per host is very small compared with that on its normal host, and with that on P. h. panamensis in Costa Rica and Nicaragua where it does not coexist with mirabilis. These data strongly suggest that in Panama some kind of competitive interaction between hertigi and mirabilis keeps the population of hertigi on P. h. panamensis at a lower level than when mirabilis is absent. They further suggest that mirabilis is absent from this host in Costa Rica and Nicaragua because of competitive displacement. 12 One could similarly argue that at these latitudes climatic or other ecological con- ditions are not suitable for mirabilis, though the occurrence of this species even farther north at higher elevations, on Trachops cirrhosus is then difficult to explain (see p. 617). 13 We are unable to detect morphological differences between specimens of hertigi from these two subspecies. WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 687 If this is so, it may also explain the failure of either hertigi or mirabilis to parasitize P. h. hastatus in the presence of S. consocius (fig. 147). However, one can also argue that this host is physiologically or ecologically (specifically?) so distinct as to be unsuitable for either hertigi or mirabilis, even though mirabilis is present on Trachops,1* and hertigi on Phyllostomus d. discolor in areas occupied by P. h. hastatus. "Delousing" the hosts and infesting them with known numbers of para- sites may provide evidence of the third type, i.e., the evidence derived through experimental manipulation. The host bats are easily kept alive in the laboratory (See Wenzel, Tipton, and Kiewlicz, p. 425, this volume) . Barnes (1965, pp. 274-276) has suggested that two California fleas of the genus Anomiopsyllus which live in the nests of the wood rat, Neotoma fuscipes are mutually exclusive (competitive displacement, sensu DeBach, op. cit.). A rather similar host distribution was noted by Tipton and Mendez (pp. 317-318, this volume) for Kohlsia mojica and traubi (and perhaps keenani) on Peromyscus n. nudipes in Panama. While some dif- ferences in geographic and ecological distribution appear to be involved, these may be factors in the process of competitive displacement (see DeBach, op. cit. p. 204). Complementary distributions on quite a different taxonomic level, may also bear analysis in this connection, for example: the world-wide host and geographic distributions of Streblidae and Nycteribiidae ; the relative absence of Streblidae and Nycteribiidae on Molossidae, which are parasi- tized by Polyctenidae ; or the absence of dermanyssid mites on oryzomyine rodent hosts, which are normally heavily parasitized by laelaptid mites of the genus Gigantolaelaps (see Yunker and Strandtmann, this volume, p. 83). III. Epidemiological Considerations Of approximately 212 terrestrial mammals reported for Panama (in- cluding man and 10 introduced species) , ectoparasites are recorded for 155. Undoubtedly many more species occur than are indicated in the compre- hensive host-parasite list (p. 797). The collections here reported were not made with statistical analysis in mind, and thus were not uniform in sam- pling, recording techniques, or in coverage. For example, relatively few collections of ectoparasites were made in the subtropical zone (approx. 2500- 5000 feet). Because of this, we have had to carefully select our data in treating various aspects of host-parasite relationships. Nonetheless, with these and other factors taken into account, it is evident that certain hosts acquire a disproportionately large number of parasites in comparison with others. Large numbers of species of ectoparasites were recorded for certain euxenous (= hospitable) hosts, as opposed to apoxenous (= inhospitable) hosts, which had few or none (fig. 148). For example, 15 or more species of ectoparasites were reported from 18 host species : 41 from the opossum, 14 See Wenzel, Tipton, and Kiewlicz (p. 617, this volume) for a discussion of the taxonomic status of "mirabilis" on Trachops. 688 ECTOPARASITES OF PANAMA Didelphis marsupialis; 37 from the spiny rat, Proechimys semispinosus ; 31 from the short-tailed bat, Carollia perspicillata azteca; 29 each from the spiny pocket mouse, Heteromys desmarestianus and the deer mouse, Pe- romyscus n. nudipes; 25 from the cotton rat, Sigmodon hispidus; 24 each from the tree squirrel, Sciurus granatensis, and the coati, Nasua nasua; 23 each from the rice rat, Oryzomys capita and the cane rat, Zygodontomys microtinus; 21 from Homo sapiens; 19 each from the porcupine rat, Hop- 30-i 25- 20- 15- 10- 5 - I •••• I . 20 25 30 35 40 Number of parasite species Fig. 148. Numbers of species of mammal hosts, according to numbers of species of ectoparasites taken from each in Panama. Negative hosts are not included. lomys gymnurus, the spiny pocket mouse, Liomys adspersus, and the rice rat, Oryzomys caliginosus; 18 from the four-eyed opossum, Philander opossum, and 15 each from the brown mouse, Scotinomys xerampelinus, the nine- banded armadillo, Dasypus novemcinctus, and the fruit bat, Artibeus j. jamaicensis. In general, analysis of the other hosts indicates that the num- ber of nonspecific (exceptional) parasites decreases with increasing ecologi- cal specialization and/or geographical restriction of the host. The species with the largest number of parasites reported for it is the common opossum, Didelphis marsupialis. Of 41 species reported for this host from Panama, probably no more than four or five are characteristic parasites and none appear to be monoxenous ! Four are characteristic para- sites of marsupials, but not of Didelphis alone. These are a dermanyssid mite, Ornithonyssus wernecki; a tick, Ixodes luciae; and a flea, Adoratop- syllai. copha; the fourth is a flea, Juxtapulex echidnophagoides, which occurs on both marsupials and the armadillo, Dasypus novemcinctus, in Panama. It WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 689 appears then, that Didelphis must acquire most of its parasites from other hosts. It is a truly ubiquitous, ecologically broadly tolerant animal, which ranges from sea level to above 5000 feet elevation, in Panama. It moves through and between a number of ecological formations, both horizontally and altitudinally, and thus comes into contact with many other components of the communities of which it is a part, including other hosts and their parasites. Some of these parasites move onto Didelphis, in varying degrees of association, mostly in small numbers, as opportunity per- mits or circumstances require. An individual opossum carries a much smaller number of species than is apparent from the comprehensive host- parasite list, the faunule varying from habitat to habitat, and especially at different elevations. The fleas reported from Didelphis illustrate this point (see Tipton and Mendez, this volume, p. 326). Below 2500 feet, they were: an agouti flea, Rhopalopsyllus a. tupinus; a paca flea, R. L lugubris; an armadillo flea, R. cacicus saevus (probably through use of armadillo burrows) ; and Polygenis klagesi, from the ubiquitous spiny rat, Proechimys semispinosus. Between 2500 and 5000 feet elevation, they were Polygenis r. beebei (from species of Oryzomys, especially O. caliginosus and 0. capita) ; and the marsupial flea Adoratopsylla i. copha. At elevations above 5000 feet, they were: the rabbit flea, Hoplopsyllus glacialis exoticus; Juxtapulex echidnophagoides, shared by Didelphis and the nine-banded armadillo; a squirrel flea, Pleochaetis d. dolens; and more abundantly than at lower elevations, a marsupial flea, Adoratopsylla i. copha. Throughout its range, Didelphis was occasionally parasitized by the cat flea, Ctenocephalides f. felis. This kind of pattern is reflected in the other groups of ectoparasites collected from the opossum, too. While its ecological tolerance and vagility obviously influence the num- ber of parasites it acquires, Didelphis may tolerate a greater variety of parasites than do many other hosts. It is a primitive animal that is only superficially specialized. Many parasites adjust to specialized differences of the hosts. It is significant that so many parasites of New World mar- supials show little specificity to host species, but rather to marsupials as a group. The large number of ectoparasites it acquires may partly reflect an easier "penetration" of Didelphis by non-specific parasites, due to the relative absence of competition from forms which are more narrowly ad- justed to it or from forms which may be adjusted to it only in certain en- vironments. The interrelationships between Didelphis and other hosts are far more numerous and complex than indicated by the above discussion. In Panama more than 70 hosts were recorded for the 37 non-marsupial parasites that were reported from Didelphis. These included 11 species of birds and reptiles and three of bats. Thus, the number of possible interrelationships through exchange of ectoparasites and/or micro-organisms, directly, or in- directly through "intermediary" hosts, is enormous. Didelphis could well play an important role in the dissemination of ecto- and endoparasites between animals that are ecologically more re- stricted. This might also be true of some of the other euxenous hosts. 690 ECTOPARASITES OF PANAMA While there appears to be little taxonomic relationship between these "car- rier" hosts, all have one feature in common with Didelphis, namely that they contact many components of a community, but in different ways. Among the bats, Carollia perspicillata azteca (see Wenzel, Tipton, and Kiewlicz, this volume, p. 638) roosts in a wide variety of sites and with a variety of hosts. Tree squirrels, on the other hand, contact a wide range of components through their foraging and nesting activities, both on the forest floor and in the tree strata. Many of the euxenous hosts are ubiquitous in yet another sense. Some, like Proechimys semispinosus, Peromyscus nudipes, Oryzomys capito and O. albigularis may be abundant more or less uniformly distributed (per- vasive) forest animals, and the grassland Sigmodon hispidus and Zygo- dontomys may be similarly distributed. Some of these hosts exhibit considerable sociability and even commensalism. Further, populations of such common and widespread species may be restricted to "pockets" in areas where their habitats are discontinuous. In the presence of an extra- ordinary abundance of food, such a population may increase far beyond the ordinary carrying capacity of the pocket. With the exhaustion of this extra- ordinary increment of food, the excess numbers of rodents may spill over into neighboring areas in outbreaks that are referred to as ratadas or "rat plagues" in rural South America.15 From the lists of ectoparasites given for them (p. 797), it is obvious that hosts like these must have contacts with many other host species or their runways, nests, etc., and/or their parasites. Further, rat plagues or ratadas must provide unusually favorable circumstances for exchange of both ecto- and endoparasites and other micro-organisms. It is quite likely that they may acquire pathogens from ecologically more restricted hosts and ectoparasites, and thus become "carriers" or even reservoirs, in the epi- demiological sense. Although no arthropod vectors have been demonstrated, the ecology of the Beni (Bolivia) epidemic of haemorrhagic fever emphasizes the impor- tance of this type of host. Proechimys guyannensis, and Calomys callosus, two rodents that have been incriminated in the epidemiology of this disease (Runs, 1964), are typical ratadas forms (Hershkovitz, pers. comm.). Proechimys semispinosus (see above) is one of the notable euxenous hosts in Panama. For ectoparasites one could assemble a graph very similar to that (fig. 148) shown for the hosts. The largest number of host species were recorded for Acarina, especially chiggers and ticks, and a few fleas like Ctenocephali- des felis. Brennan and Yunker (this volume, p. 235) recorded 35 hosts each for the chiggers Eutrombicula goeldii -and E. alfreddugesi, in Panama. The epidemiological importance of such non-specific (promiscuous) or of 15 Hershkovitz (1962) has given an extended discussion of pocket populations and ratadas. He cites instances in which ratadas of South American cricetines are correlated with cyclic fruiting and seed production of bamboos. As many as 17-20 years may elapse between fruiting. WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 691 polyxenous ectoparasites, especially the immature stages of heteroxenous ticks, is well documented in the literature on arthropod-borne diseases. If survey and sampling techniques are sufficiently refined, it should be possible, through modern data processing, to analyze many of the complex interrelationships between hosts, parasites, and the extra-host environ- ment. This should not only lead to a better understanding of the population dynamics, but may make it possible to evaluate hosts and ectoparasites of a given area in terms of their potential epidemiological importance. IV. Altitudinal distribution In the following discussion we refer to tropical, subtropical, and montane zones in the sense of Holdridge and Budowski (see fig. 149 and Fairchild, "Introduction," p. 5, this volume). This differs from the classification of Goldman and Zetek (1926) and that of Fairchild (loc. cit.). The tropical zone of Fairchild correlates more closely with many parasite distributions than does that of Holdridge and Budowski. When we describe groups as being temperate in distribution, we are referring to their climatic ad- justment without restricting them geographically. A. Mites and Ticks Family Laelaptidae Subfamily LAELAPTINAE. Most of the laelaptine mites occurred in the tropical and subtropical zones. Some, like Laelaps nuttalli and Echinolae- laps echidninus (both introduced) , were taken only near sea level. Others, like Haemolaelaps glasgoivi, were taken from sea level to 7500 feet. Tipton, Altman, and Keenan (this volume, p. 34) suggest that this species is com- posite. Most species of Gigantolaelaps were tropical and subtropical, like their oryzomyine hosts. However, G. inca (described from Peru) was taken only above 5000 feet, as was Eubrachylaelaps jamesoni (described from Mexico) . The native species of Laelaps showed considerable differences in their altitudinal ranges. Some, like pilifer and dearmasi, occurred only in the tropical and subtropical zones, while others, like thori, were taken be- tween 2000 and 7800 feet. Families Dermanyssidae, Trombiculidae We cannot assess the altitudinal distributions of these two families. In general, the Dermanyssidae appeared to be tropical and subtropical in dis- tribution, though Ornithonyssus bacoti, the Tropical Rat Mite, was taken from sea level to 5000 feet. The Tropical Fowl Mite, O. bursa, occurred at low elevations, while O. sylviarum, the Northern Fowl Mite, was taken at 5700 feet on Volcan Chiriqui, the southernmost record of this species. Family Spinturnicidae The altitudinal distribution of these mites was very similar to that of the Streblidae (see below) . As in the case of the streblid Joblingia, the spin- turnicid genus Paraspinturnix on Myotis n. nigricans was taken only in the lower montane zone. 692 ECTOPARASITES OF PANAMA Superfamily Ixodoidea The altitudinal distribution of the ticks is discussed by Fairchild, Kohls, and Tipton on pp. 168-170, this volume. Most species of Amblyomma oc- curred in the lowlands and those of Ixodes in the highlands. B. Rove Beetles Family Staphylinidae The parasitic amblyopinine staphylinid beetles are primarily temperate in their distribution, both altitudinally and latitudinally in South America. The Middle American species, including those taken in Panama, are from the montane zones. The distribution shown for Amblyopinus tiptoni (fig. 150) is typical. C. Batflies (Diptera) Family Streblidae The Streblidae are primarily tropical and subtropical in distribution, with very few species in the warm temperate and none in the cool temperate regions. This is strikingly illustrated by their altitudinal distribution in Panama. Of the 66 species recorded from Panama, 49 (ca. 75.4% ) were re- stricted to the tropical zone, or nearly so. The ranges of a few of these ex- tend into the lower elevations of the subtropical zone. About nine species (12.3%) were either restricted to the subtropical zone or ranged from the tropical or subtropical zones to the lower altitudes of the lower montane. Three of these — Anastrebla mattadeni, A. modestini, and Exastinion clovisi — occurred only on bats of the genus Anoura, which are primarily subtropical. The other six species were parasites of fruit- eating bats (Phyllostomidae: subfamilies Stenoderminae and Sturnirinae). Four species (6.2% ) were taken only in the lower montane zone. These were Joblingia schmidti from Myotis n. nigricans, Anatrichobius scorzai from M. chiloensis, Trichobius keenani from Sturnira ludovici, and T. vampyropis from Vampyrops vittatus. No species were taken in the upper montane zone. A few species had a considerable altitudinal range. Three (4.6%) occurred with their hosts, Desmodus r. rotundus and Trachops c. cirrhosus from sea level to nearly 5600 feet. Paratrichobius "longicrus" showed a similar altitudinal range, but occurred on different stenodermine hosts at different elevations. It may be a composite species (see Wenzel, Tipton, and Kiewlicz, this volume, p. 519) . For the most part, the altitudinal range of a host and its Streblidae coincided closely. An outstanding exception to this is the very restricted distribution of Joblingia, schmidti, as compared with its host Myotis n. nigricans. As Handley points out (p. 770) this bat is probably a composite species. According to him (pers. comm.) the montane population is prob- ably a separate species, while the lowland population in Panama may consist of several cryptic species. This probably explains the puzzling distri- bution of species of Basilia (Nycteribiidae) on this host in Panama and else- where in Latin America. WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 693 Family Nycteribiidae Although the Nycteribiidae appear to tolerate cooler climates and pene- trate further into the temperate regions than do the Streblidae, this was not reflected in the altitudinal distributions of the seven species that have been collected in Panama. All are from the tropical zone. D. Fleas The altitudinal distribution of the fleas is shown in figure 149. It should be emphasized that the ranges of a number of the species would be extended by further collecting and also that the chart does not reflect relative abun- dance at different altitudes. Although some species were taken throughout a considerable altitudinal range, they were obviously most abundant in a much narrower one. This is reflected in the data given by Tipton and Mendez (beginning on p. 325, this volume) . Family Pulicidae Tunga penetrans is the only New World species of the genus that is not restricted to Southern Brazil (Sao Paulo, Bahia, Goyaz). It occurs from South America to Mexico and has a correspondingly broad altitudinal range as well (fig. 149). The known species of Rhynchopsyllus have the same altitudinal distribution as their host bats (genus Molossus). Juxtapulex echidnophagoides, known from Costa Rica (+4300 feet elev.) , and Panama, has an altitudinal range similar to that of the batfly, Joblingia, as does Hoplopsyllus glacialis exoticus, from Panama. All of the 859 specimens of Juxtapulex collected in Panama were from above 5000 feet elevation. The altitudinal ranges of Pulex irritans and P. simulans are interesting. They are discussed by Tipton and Mendez (this volume, p. 293), who feel that simulans may be lowland and irritans highland in Middle America. Family Rhopalopsyllidae These fleas are principally South American. Most of the South Ameri- can genera are decidedly temperate in distribution, but the two large genera, Rhopalopsyllus and Polygenis, are represented in the subtropical and tropi- cal zones. Nine species of these two genera were taken in Panama. Seven were primarily tropical and subtropical in distribution and two were taken only in the lower montane. Family Ceratophyllidae Species of five genera were collected in Panama. Ceratophyllus altus, the only species of the genus known from Panama, occurs in the lower montane zone. Dasypsyllus gallinulae perpinnatus, another bird flea, is altitudinally and latitudinally temperate, and occurs from Western North America to Panama and probably South America (Johnson, 1957). It was taken in the lower montane zone of Panama. Dasypsyllus I. venezuelensis occurs in the same zone, but at slightly lower elevations. The other three genera of Ceratophyllidae that were taken in Panama are Jellisonia, Pleo- chaetis, and Kohlsia. These three genera of rodent fleas are centered in the 694 ECTOPARASITES OF PANAMA jlUOUj)03S E||AsdOX)|d u9>|unA e||9zua/v\ egdoo -| BOjfOUJ s|L|dBj6 *6 ,, sjsuajanzB EJSIHOX SU9JOP 'P • -ds • aBuosuqof Bjuosjnar SjSU8|9nZ3U9A '| n BSOI09dS ' sijqn6n| snujdn; -B sn||Asdo|Bdoijy isa6e|)| juunp snoijoxa -6 sn||AsdO|doH suB|nujjs ,, ! X9|Hd a s 83 a OH NOI1VAH3 WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 695 warm temperate and temperate zones of Middle America and Mexico, and this is reflected in their altitudinal distribution. A few species were taken in the subtropical, but none in the tropical zone. Family Ischnopsyllidae The bat fleas were represented by two lowland South American genera, Hormopsylla and Ptilopsylla, both of which occurred at or near sea level, and by Sternopsylla distincta speciosa, which occurred in the lower mon- tane zone. Sternopsylla is closely related to the other two genera and occurs as S. distincta distincta in central and southern United States. Family Hystrichopsyllidae Adoratopsylla i. copha, taken in subtropical and lower montane zones, is a South American race of a species belonging to a South American genus. It occurs in Mexico as A. i. intermedia. Strepsylla dalmati and Wenzella yunkeri, both known only from the upper montane zone, belong to genera that are endemic to the highlands of Middle America. Family Stephanocircidae Plocopsylla scotinomi, the only known Middle American species of the temperate South American subfamily Craneopsyllinae, was taken only in the upper montane zone. In general, ectoparasite taxa of South American relationships occurred below and those of holarctic affinities above 5000 feet elevation. V. Altitudinal Concordance Between Hosts and Parasites The altitudinal distributions of host-limited parasites like the Spin- turnicidae and of nearly host-limited ones like the Streblidae and Nycteri- biidae paralleled those of their hosts quite closely. The collections of biting and sucking lice were not adequate to determine to what extent this was true for them. It is not true of groups like the fleas, ticks, and the laelaptine mites, which in general are not host-limited. The data for the Laelaptinae must be treated with considerable caution, because some of the species may be composite (see Tipton, Altman, and Keenan, this volume, p. 31). In figure 150, we have shown the altitudinal distribution of a series of representative mammal hosts and some of their characteristic non-host- limited parasites. The altitudinal ranges must not be taken too literally, for several reasons. First, they do not reflect the relative abundance of the hosts and parasites at different elevations. The data on altitudinal distribution of the fleas given by Tipton and Mendez (beginning on p. 325, this volume) give an indication of this, but the nature of the field data does not permit detailed analyses of this type. Second, the data are lumped for all of Panama. Regional and edaphic climatic differences greatly modify the fauna and flora and thus affect the altitudinal range of both host and parasite in a given locality, as do other environmental factors. Environ- ments on the drier Pacific slopes of the mountains are quite different from 696 ECTOPARASITES OF PANAMA WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 697 those on the more humid Atlantic slopes. Similarly, the habitat at the sum- mit of a mountain 4000 feet high differs from that at the same altitude on a mountain that is 10,000 feet high. Carefully documented field collections and ecological observations should provide an appreciation of these factors. In spite of the limitations of the data, it is clear that the altitudinal ranges shown for many parasites do not coincide with those of the hosts. Nor does the altitudinal range of a parasite on a given host necessarily coincide with the entire altitudinal range of the parasite. In some instances, as in the case of the flea Jellisonia johnsonae (fig. 150) these differences could reflect sampling techniques or local edaphic differences. On the other hand, the flea Strepsylla dalmati was taken on Peromyscus n. nudipes from about 5500 feet to the altitudinal limit of this host at 8000 feet ; but it was taken above this altitude on other hosts, like Reithrodontomys and Scoti- nomys. Similarly, Pleochaetis altmani was taken on Scotinomys xeram- pelimis and Reithrodontomys creper, between 5600 and 8000 feet (fig. 150) , but a few specimens were taken on Reithrodontomys sumichrasti at 10,300 feet. Other authors have also indicated that parasites may not have as broad distributions as the hosts. This may be an attribute of non-host-limited forms. The population density of most hosts must decrease as one moves away from optimum habitat conditions, and the opportunities for their non- host-limited parasites to encounter a suitable host probably decrease corre- spondingly, or even disproportionately because of their lesser vagility. Un- der these circumstances, one might expect selection to narrow the ecological range of a parasite so that it corresponds more nearly to the optimal en- vironmental conditions of the most suitable host(s). In this respect, it should be remembered that homoiothermal hosts would have relatively greater ability to move into climatically less suitable habitats than would poikilothermal non-host-limited parasites. It must also be noted that distribution may be correlated with subspecies of the host. For example, no fleas were taken on Sciurus granatensis below 5000 feet. All, including those apparently specific to this host, were from the subspecies S. granatensis chiriquensis, above 5000 feet. Similarly, Wen- zella yunkeri was taken only from Heteromys desmarestiamcs chiriquensis, but not from the other subspecies that occurred at lower elevations. It is clear that in the future, host identifications to subspecies must be given whenever possible. Correlation between host subspecies and parasites and altitudinal data may in some cases indicate that the taxonomic status of the host merits further investigation. VI. Faunal Relationships Few groups of ectoparasites are well enough known for the Neotropical Region as a whole to permit an evaluation of the faunal relationships of their Panamanian representatives. Though it is not well known for either Middle or South America, we believe that the flea fauna of Panama and cer- tain other critical areas has been sampled well enough to give a general picture of the distribution of the families and genera. Very large and repre- sentative collections of batflies (chiefly Streblidae) are at hand from Middle 698 ECTOPARASITES OF PANAMA America, and representative samples have been secured from northern South America and to some extent Peru, though the fauna of the Amazon basin is still largely unknown. The distribution of the amblyopinine beetles (Staphylinidae) and the laelaptine mites also contribute some interesting points that bear on faunistics and zoogeography. In the following discussion we refer to all of the North American con- tinent between the northern boundary of Mexico and the southern boundary of Panama as Middle America, and to the area north of Mexico as North America. We have arbitrarily used the term in this geographic sense as did Baker (1963) , rather than in a zoogeographic sense (see Hershkovitz, 1958) . However, we have followed Hershkovitz (op. cit.) in his use and definition of the Patagonian Subregion. In the following discussion, it should be remembered that of 50 genera and ± 300 species of cricetine rodents recorded from South America, all but two recent invaders (see Hershkovitz, this volume, p. 738) are complex penis types and no parasites have been reported for these two. Thus, when we refer to South American complex-penis-type Cricetinae, we do not infer that there are South American simple-penis-type hosts for which ectopara- sites have been reported. Both groups are well represented in Middle America. A. Mites Family Laelaptidae Subfamily LAELAPTINAE. The geographic and host distribution of the lae- laptine mites are instructive. Eight genera were taken from rodents in Panama. Four of them belong to a complex of six closely related genera (Tipton, 1960, p. 258). These are Laelaps, Tur, Mysolaelaps, and Echino- laelaps. Laelaps and Echinolaelaps are cosmopolitan and occur mostly on Murinae, Microtinae and complex-penis-type Cricetinae. They do not oc- cur on peromyscines. Except for the introduced L. nuttalli, the species were taken almost exclusively on Oryzomys and related complex-penis-type Crice- tinae. The species of Tur occur almost entirely on caviomorph rodents of the family Echimyidae, of South American origin. Mysolaelaps also is neotropical and occurs on South American complex-penis-type cricetines and on caviomorphs. Two related genera, Longolaelaps and Tricholaelaps occur on Murinae (Rattus) in Sumatra. Haemolaelaps is a cosmopolitan genus with more than 60 species which are associated with a wide range of hosts, but chiefly sciuromorph rodents (Tipton, op. cit., p. 242). H. glasgoivi, the only species taken in Panama, occurred on a wide variety of small inammals, including rodents of the three suborders and marsupials (table 12). The 14 described species of Gigantolaelaps, chiefly South American, are almost exclusively parasites of oryzomyine Cricetinae. The five species taken in Panama were originally described from South America; all have been taken in Venezuela, four of them also in Brazil, the fifth in Peru. All except one occurred below 5000 feet in Panama. Gigantolaelaps inca, de- scribed from Peru, was taken only above 5000 feet, chiefly on Oryzomys WENZEL AND TIPTON I HOST-PARASITE RELATIONSHIPS 699 albigularis and O. alfaroi (fig. 150) sparingly on Peromyscus and Didelphis. The genus Eubrachylaelaps is closely related to Haemolaelaps, Cavilae- laps (on South American caviomorphs) and Gigantolaelaps (Furman, 1955; Tipton, op. cit.). The eight described species occur mostly on the simple-penis-type cricetines, Peromyscus and Neotomodon, in the subtropi- cal and temperate altitudes and latitudes from Panama to California. Eubrachylaelaps rotundus Fonseca has been reported only from South America (Brazil and Venezuela) from a variety of sciuromorph and com- plex-penis-type myomorph (cricetine) rodents and marsupials. About half of the specimens taken in Venezuela (Furman and Tipton, 1961) were from Zygodontomys. This appears to represent a host transfer for Eubrachy- laelaps. The total numbers of Laelaptinae collected are tabulated for each genus of mites and of hosts, in table 12. We are aware that this method of presentation has many weaknesses, e.g., it does not show the host asso- ciations by species. Nonetheless it does show the preponderant associa- tions of Eubrachylaelaps with Peromyscus, of Gigantolaelaps with complex- penis-type Cricetinae, especially Oryzomys; of Laelaps with caviomorph rodents and complex-penis-type Cricetinae; the promiscuity of Haemo- laelaps glasgowi; the restriction of Steptolaelaps to the sciuromorph Hetero- myidae, and of Tur to the caviomorph Echimyidae. To summarize — of the eight genera that occur in Panama, three (Lae- laps, Haemolaelaps, Echinolaelaps) are cosmopolitan; three (Mysolaelaps, Gigantolaelaps, Tur) are primarily South American, and are associated in Middle and North America with rodents of South American derivation or affinities ; while Eubrachylaelaps and Steptolaelaps appear to be Middle and North American in origin. The principal hosts of one South American genus (Tur) are caviomorph rodents. The principal hosts of the other two, as well as of two of the cos- mopolitan genera, are primarily Murinae, and Microtinae, and complex- penis-type Cricetinae. Most Murinae and the Microtinae have complex penes, too. The occurrence of the preponderance of the Laelaptinae on rodents of this type suggests that Hershkovitz's treatment (1962) of the Cricetinae as a subfamily of the Muridae is sound. It is of special sig- nificance that many of the host associations of the genera of Laelaptinae are at a suprageneric level, as evidenced strikingly by the species of Tur on Echimyidae (Caviomorpha) and of the genus Steptolaelaps on Hetero- myidae (Sciuromorpha) . Thus, the differences in host associations between the related genera Eubrachylaelaps and Gigantolaelaps, with Eubrachy- laelaps on simple-penis-type Cricetinae (Peromyscini, see Hooper and Mus- ser, 1964, p. 54) and Gigantolaelaps only on complex-penis-type Cricetinae (chiefly oryzomyine genera) suggest a long period of geographic isolation and of differentiation of the two groups of hosts and parasites. If we have belabored this point, it is because we believe it is important to establish the point — as evidenced by their mites and other ectoparasites, as well as by their distribution — that the simple-penis-type Cricetinae do appear to constitute a group, phyletically distinct from the other New World 700 ECTOPARASITES OF PANAMA TABLE 12. NUMBERS OF LAELAPTINE MITES COLLECTED IN PANAMA, ACCORDING TO GENERA OF HOSTS AND MITES. (introduced species not included) Hosts Order MARSUPIALIA Family Didelphidae Philander (opossum) Metachirus (nudicaudatus) Didelphis (marsupialis) Order RODENTIA Suborder Sciuromorpha Family Sciuridae Sciurus (granatensis) Family Heteromyidae Liomys (adspersus) Heteromys (2 spp.) Suborder Myomorpha Family Cricetidae Subfamily Cricetinae *Oryzomys (10 spp.) *Nectomys (alfari) *Zygodontomys (microtinus) *Sigmodon (hispidus) Tylomys (panamensis) Peromyscus (2 spp.) Reithrodontomys (3 spp.) Scotinomys (2 spp.) Suborder Caviomorpha Family Echimyidae Proechimys (semispinosus) 1610 Hoplomys (gymnurus) 18 * = Complex-penis-type Cricetinae. s. .2 a S § _o .e P* us o to A d 8 co "~* d to A & a en to e A _g to CO A e on » S o o ~2 » *> « Hosts I £ ^ 5; % h Order RODENTIA Suborder Sciuromorpha Family Sciuridae Sciurus (2 spp.) 129 20 .. 245j 3 Family Heteromyidae Heteromys (desmarestianus) 80 1 . . 1 . . 59 Suborder Myomorpha Family Cricetidae Subfamily Cricetinae *Oryzomys (3 spp.) 296 14 1 20 *Nyctomys (sumichrasti) 4 . . 3 Peromyscus (2 spp.) 329 409 9 122 14 Scotinomys (2 spp.) 38 3 45 41 5 Reithrodontomys (3 spp.) 107 1 3 96 8 . . * = complex-penis-type Cricetinae. f = Pleochaetis d. dolens. The remaining genera — Pleochaetis, Jellisonia and Kohlsia — are of spe- cial interest, because they appear to be primarily temperate genera which are restricted to or centered in the highlands of Middle America, chiefly on Peromyscus and related simple-penis-type Cricetinae and tree squirrels of the genus Sciurus. Their host associations are shown in table 16. Only two species of the genus Pleochaetis are recorded from the (south- western) United States. One of these, P. sibynus, also occurs in Mexico. The other is a subspecies of P. equatoris, a species of unusually wide dis- tribution. The nominate form occurs from Peru and Ecuador to Mexico, though it was not taken in Panama. Pleochaetis dolens is known from Ecuador (P. d. quitanus) and Panama to Mexico (P. d. dolens). The only other species known from Panama is P. altmani. It was taken chiefly from species of Reithrodontomys, Scotinomys xerampelinus, and Peromyscus n. nudipes, while P. dolens dolens was taken chiefly from Sciurus granatensis chiriquensis and Peromyscus nudipes nudipes, with scattered records from other hosts. The other Middle American species have been taken chiefly from Peromyscus. Two species are known only from South America, P. smiti and P. apollinaris, both from Colombia. They have been taken from various hosts, chiefly complex-penis-type Cricetinae. WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 711 With a single exception, the 17 species of the genus Kohlsia are limited to Middle America. Six species have been recorded from Panama, only one from South America. This latter species, K, campaniger was taken from "Hesperomys" sp. in Ecuador. Its generic assignment is doubtful (Traub, 1952). The species of Kohlsia appear to be parasites chiefly of Peromyscus and other simple-penis-type Cricetinae and of Sciurus. Of the Panamanian species, K. azuerensis was taken from Peromyscus flavidus; K. graphis graphis was taken only from Sciurus granatensis chiriquensis ; of 18 specimens of keenani, eight were from simple-penis-type cricetine rodents (Peromyscus and Scotinomys) and ten from complex-penis-type cricetines (Oryzomys) ; 19 K. mojica is known only from Peromyscus n. nudipes; K. tiptoni from Sciurus granatensis chiriquensis; traubi chiefly from Peromys- cus n. nudipes, with scattered records from other hosts. There are ten known species of Jellisonia. Nine have been reported from Mexico; of these, ironsi Eads and bullisi Traub and Johnson have also been taken in southwestern United States. Jellisonia johnsonae is known only from Panama. The genus has not been reported from South America. Most of the species have been taken from Peromyscus and related genera of simple-penis-type cricetine rodents, like Baiomys and Reithrodontomys, but one has been taken from Microtus (Microtinae) according to Traub (1952). Family Ischnopsyllidae Six genera of bat fleas have been reported from the New World. Three of these (Hormopsylla, Ptilopsylla, and Rothschildopsylla) are known only from the tropical lowlands. Rothschildopsylla is known only from South America. Three species of Hormopsylla are known from South America, and one (kyriophila) from Panama. They are parasites of molossid bats, as is true of the two species of Ptilopsylla, dunni from Panama and leptina from Brazil. The genus Sternopsylla is obviously South American in its relationships. The single species is closely related to those of the pre- ceding two genera (Hopkins and Rothschild, 1956) and like them occurs on molossid bats. It is represented by the subspecies S. distincta texana in the United States and Mexico, S. distincta speciosa in Panama and Peru, S. d. distincta in Paraguay and Parana, Brazil. Sternopsylla d. speciosa was taken in the lower montane zone in Panama (fig. 149). The three genera of bat fleas that have been taken in Panama thus appear to be South American in their relationships. Myodopsylla and Nycteridopsylla, the other two genera that occur in the New World, are holarctic. Nycteridopsylla does not occur in Middle or South America. The North American Myodopsylla collinsi has been taken as far south as Chocoyos (Dept. of Chimaltenango) Guatemala, in the montane zone. Three species of Myodopsylla have been described from 19 The few species of Oryzomys that were parasitized by them had such a scattered representation of flea species, that none can be regarded as characteristic hosts for these fleas. 712 ECTOPARASITES OF PANAMA South America, but none have been reported from between Guatemala and Colombia. The species of both genera parasitize vespertilionid bats of the genus Myotis. It seems likely that a species of Myodopsylla may be found in the highlands of Panama. The distribution pattern of this genus in the New World somewhat parallels that of certain nycteribiid batflies of the genus Basilia, which also occurs chiefly on Myotis. Family Pygiopsyllidae This family is known from the Australian, Oriental, Palaearctic (one species) , Ethiopian and Neotropical Regions. Three subfamilies are recog- nized, two with only one genus each, the third (Pygiopsyllinae) with ±15 genera. Only the Pygiopsyllinae are represented in the New World, by one genus, Ctenidiosomus. The three described species are mostly from complex-penis-type cricetine rodents (Oryzomys, Thomasomys, Rhipidomys, Neomys). They are montane (Andean) in distribution, with species from Peru, Ecuador and Colombia. Undescribed species are known from South America. One has recently been collected in Costa Rica (det. Traub ; Truxal, pers. comm.) ! It was not taken in Panama, though it may occur there. Family Hystrichopsyllidae The Hystrichopsyllidae occur chiefly in the Holarctic Region. At first glance, it might appear that the South American forms represent a recent intrusion of northern hemisphere groups into South America. An analysis of the distributions and relationships does not support this. The South American representatives belong to three subfamilies, as follows : Subfamily HYSTRICHOPSYLLINAE. Ctenoparia is the only genus of the tribe Ctenopariini. The two known species occur on complex-penis-type cricetine rodents in Chile and Argentina. Subfamily CTENOPHTHALMINAE. No representatives of this subfamily have been taken in Middle America, except Ctenophthalmus. There are three genera in South America: Chiliopsylla (allophylla Rothsch.), whose host is dubious ; Neotyphloceras, with two species that occur almost exclu- sively on complex-penis-type cricetine rodents in Argentina and in the Andes north to Colombia; and Agastopsylla, with six species all in the Patagonian Subregion, at high elevations or far southern latitudes, chiefly on complex-penis-type cricetines. Chiliopsylla and Neotyphloceras are the only genera of the tribe Neotyphloceratini, while Agastopsylla is the only genus of the tribe Agastopsyllini. The distribution and taxonomic position of these genera, including those whose species are on cricetines suggest a con- siderable period of isolation in South America. Subfamily DORATOPSYLLINAE. Smit,(1962) iists six genera, represent- ing four tribes : the Idillini, with a single genus Idilla from marsupials in Australia; the Acedestini, with a single monotypic genus Acedestia from marsupials in Australia ; Tritopsyllini, with a single genus Adoratopsylla, from marsupials in South and Middle America; and Doratopsyllini with Doratopsylla, Corrodopsylla, and Xenodaeria from insectivores. Xenodaeria is known only from Sikkim. The holarctic Doratopsylla and Corrodopsylla occur on shrews. The southernmost record of these is a species of Corrodop- WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 713 sylla from Guerrero, Mexico (Traub, pers. comm.). No fleas of this genus were found on the four specimens of the shrew Cryptotis sp. examined in Panama. Three species of the subgenus Adoratopsylla have been taken on South American marsupials, mostly at southern latitudes in Brazil and at inter- mediate elevations in Venezuela and Colombia. The single species of the subgenus Tritopsylla is represented in South America by four subspecies, one of which, A. (T.) intermedia copha, occurs in Panama. A fifth sub- species has been taken in Mexico. We have dwelt at some length on these doratopsylline fleas because Dora- topsylla and Adoratopsylla have been considered to be very closely related genera, and it seemed that the occurrence of Adoratopsylla on marsupials might represent a recent host divergence from the Holarctic Doratopsylla. But, as pointed out by Smit (op. cit.) , the geographic and host distributions of the genera of Doratopsyllinae and the tribal allocations of the genera suggest that the host associations of these fleas are very old. Thus the occurrence of Adoratopsylla in Middle America most likely represents an intrusion from South America. Subfamily NEOPSYLLINAE. None of the genera of this subfamily have been taken in South America. Only two genera have been reported from Middle America : a species of Meringis from Mexico and seven species of Strepsylla, six of them from Mexico and one from Panama. The species of Strepsylla are montane and occur chiefly on species of the genus Peromyscus and re- lated simple-penis-type Cricetinae. The genus appears to be endemic to Middle America. Subfamily RHADINOPSYLLINAE. This holarctic subfamily is not represented in South America. The genus Wenzella, which represents a taxonomically isolated tribe (Traub, 1953; Hopkins and Rothschild, 1962), is known from only two species, W. obscura Traub from Guatemala and yunkeri n.sp. from Panama. Both were taken at altitudes above 6000 feet on Heteromys des- marestianus (Heteromyidae). The genus appears to be an old endemic of the Middle American highlands. The Heteromyidae, too, are primarily Middle American in distribution. Family Stephanocircidae Two subfamilies are recognized. The Stephanocircinae include a single genus (Stephanocircus) with five species, all from Australia and Tasmania, and occuring chiefly on marsupials, but also on Rattus. The Craneopsyllinae, with seven genera and 27 described species are known only from South America, with the exception of Plocopsylla scotinomi from Panama. Nearly all of them are from the Patagonian Subregion. Many are found at ele- vations of 10,000 to 16,000 feet in Peru.20 Although the species of Plocopsylla parasitize caviomorph and myo- 20 Because of distinctive conditions in Peru, temperatures at these high altitudes are more comparable to those at lower altitudes in Panama. 714 ECTOPARASITES OF PANAMA morph rodents and marsupials in South America, they appear to occur most frequently on complex-penis-type Cricetinae. The principal host of P. scotinomi was clearly Scotinomys teguina, a simple-penis-type cricetine. This probably represents a host transfer, similar to that undergone by the species of Amblyopinus (rove beetles) in Middle America. Summary Perhaps the most notable feature of the flea fauna of Panama (and all Middle America) is that there are no endemic tropical lowland genera. The tropical-subtropical lowland fauna is with few exceptions, predomi- nantly South American, much of it probably of recent origin. Most of it has close relationships — even to the subspecies level — with coastal Colombia, Ecuador, and Peru. Above 5000 feet the fauna is largely Middle and North American or Holarctic in its relationships, but, excluding introduced spe- cies, it has few genera in common with the fauna north of Mexico, except for species that occur on ubiquitous or highly vagile hosts, like some bats, rabbits, and birds. Of the 25 species taken above 5000 feet (fig. 149) 13 (52%) belong to genera that are known only from or are largely centered in Middle Amer- ica : Pulex, Juxtapulex, Strepsylla, Wenzella, Kohlsia, Pleochaetis, and Jelli- sonia. Of 16 species that were taken only above 5000 feet, the percentage was considerably higher (approx. 69 %) . Two others were obviously of northern origin (Hoplopsyllus glacialis and Ceratophyllus altus) . The main point illustrated by these data is that in Middle America there are a number of genera that are endemic or nearly so and nearly all are found at higher altitudes and/or northern latitudes, and chiefly on host genera that are restricted (or nearly so) to Middle and North America. Of the 18 species that were restricted to the montane zones, or nearly so, only five can be regarded as South American derivatives. They are Poly- genis atopus, Rhopalopsyllus I. cryptoctenes, Sternopsylla speciosa (bats), Dasypsyllus I. venezuelensis (birds), and Plocopsylla scotinomi. Two others, which occur in Middle America but were not taken in Panama, are Hectopsylla knighti (birds) from Mexico and Ctenidiosomus sp. from Costa Rica. With the exception of Plocopsylla, none of the South American genera occurred above 6000 feet in Panama. Only three species of Middle or North American genera occurred below 4000 feet, none below 2000 feet. No endemic genera of close South American relationships are known to occur in the montane zones of Panama or elsewhere in Middle America. The converse is true of South America. The fleas of South America like those elsewhere, are primarily temperate in distribution. Thus, in South America they occur chiefly in the Patagonian Subregion and in the Cordilleras. With the exception of a few obviously recent arrivals, a few highly vagile forms, and some introductions, the fauna consists chiefly of endemics or of groups, like the Rhopalopsyllidae, which are predominantly centered in South America but may have dispersed out- ward. Among these latter are cold temperate forms like Parapsyllus (pen- guin fleas), or genera with warm-adapted species like Rhopalopsyllus and Polygenis, which occur in Middle and North America, chiefly on hosts of WENZEL AND TIPTON I HOST-PARASITE RELATIONSHIPS 715 South American origin. Only a very few temperate South American forms like some bird fleas (e.g., Dasypsyllus lasius) and PlocopsyVa scotinomi and Ctenidiosomus sp. occur in Middle America. The relationships of the South American fleas are chiefly with the southern hemisphere of the Old World, especially with the Australian Region, but also with the Ethiopian and Oriental Regions. A few South American genera like Ctenoparia and Adoratopsylla belong to hystrichopsyllid groups like the Hystrichopsyllinae and Doratopsyllinae which are predominantly Holarctic. However, these genera belong to endemic tribes and are probably relicts. Very few representatives of the Middle American fauna have penetrated South America, except a few species of Pleochaetis, and possibly Kohlsia and a few other recent invaders like Cediopsylla and Euhoplopsyllus, on such vagile hosts as rabbits and possibly squirrels. Since 1), there are no endemic tropical lowland genera in Middle America; 2), the native lowland fleas are overwhelmingly South American or of South American affinities, and 3), there are very few representatives of Middle and North American genera in South America, then it appears that recent dispersals of small mammals and their fleas have been predominantly from South into Middle America rather than the converse. An earlier (perhaps pre-Pleistocene) dispersal may account for endemic species of Rhopalopsyllus and Polygenis in Middle America. Very few montane forms appear to have moved in either direction across the isthmus. We believe that dispersal across the "bridge" by some of these temperate genera 21 may have been facilitated by lower tempera- tures than exist at present (see Nygren, 1950; Dorf, 1959, map 5). This may well have been one of the principal factors which permitted dispersal of many forms between the two continents, rather than the elevation of the Bolivar trough. VII. Zoogeographic Conclusions With the exception of some conspicuously recent immigrants 22 into South America from Middle America, the distribution patterns shown by the ectoparasites hold true for most of the hosts, too. It is especially strik- ing in the case of the cricetine rodents, those of the montane region being largely simple penis types (Peromyscini), which have barely penetrated South America, while cricetine hosts of the lowland subtropical zones are largely complex penis types of South American relationships, e.g., Oryzomys, Sigmodon, and Zygodontomys. Among the ectoparasites, we have found little evidence of endemism in the tropical lowlands of Middle America, except north of Panama, and there, entirely at the species level. It is significant that nearly all of the lowland parasites belong to: 1), genera which otherwise occur only in the lowland 21 For example, such South American genera as Ctenidiosomus and Plocopsylla (fleas) and Amblyopinus (staphylinid beetles), and Middle American genera like Pleochaetis and Kohlsia (fleas). 22 Like rabbits, some squirrels, cats, deer, shrews, spiny mice (Heteromys) , camels, bears, etc. 716 ECTOPARASITES OF PANAMA tropics of South America, e.g., the bat fleas Hormopsylla, Ptilopsylla, and Rhynchopsyllus ; or 2) , "expanding" South American genera, like Polygenis and Rhopalopsyllus, which on that continent have both temperate and warm- adapted species. We do find ample evidence in the fleas, mites, and Streblidae, of the existence of a distinctive Middle American ectoparasite fauna, but in the subtropical and especially the temperate altitudes and latitudes. It is char- acterized by the flea genera Pleochaetis, Kohlsia, Jellisonia, Wenzella, Strep- sylla, and probably Juxtapulex; by the streblid genus Joblingia and the gen- eralized members of the Trichobius major group ; and by laelaptid mites of the genera Steptolaelaps and Eubrachylaelaps. Deserts may have isolated many of these genera from North America (north of Mexico) and they may also have been a barrier to the southward dispersal into Mexico of more recent northern temperate groups. They appear to have been more effective barriers for non-host-limited parasites than for their hosts or than for host-limited parasites, probably because of the narrower climatic toler- ances of the non-host-limited forms. What is especially interesting is that these Middle American groups of ectoparasites are largely restricted to simple-penis-type cricetine rodents, especially Peromyscus, Reithrodontomys, and Scotinomys in Panama, as well as other genera like Baiomys and Neotomodon farther north. These hosts, too, are primarily temperate (altitudinally and latitudinally) in distribution. These distribution patterns suggest that the tropical lowland fauna has probably moved between Panama and South America with relative ease, for a considerable time. On the other hand, the fauna of the subtropical and temperate (montane) zones of Middle America shows an increasing degree of endemism, and thus of isolation from South America, the higher the altitudes. This suggests that the extensive "sea" of lowland tropical rain forests in Panama, especially in the Darien, may have been more effec- tive in isolating the distinctive temperate Middle and South American ectoparasites and their hosts than was the waterway between Panama and South America (the Bolivar Portal). Portals and lowlands north of the Isthmus of Panama probably also served as isolating barriers, but we are not concerned with them in this discussion. It is difficult to reconcile these conclusions with the view of Simpson (1950) and Patterson (1957) that the ancestors of the complex-penis-type South American Cricetinae immigrated into South America over a land bridge which arose in the Pliocene-Pleistocene, and subsequently : 1 ) , evolved into an array of 50 genera and ±300 species; 2), acquired a considerable Table 17. Explanation of abbreviations. AUS. = Australian region; M.AM. = Middle America; cos. = Cosmopolitan; ETH. = Ethio- pian region; HOL. = Holarctic Region; N.AM. = North America, north of Mexico; NE. = Nearctic Subregion; NW. = New World; OR. = Oriental Region; PAL. = Palaearctic Sub- region ; PAN. = Panama ; s.AM. = South America ; so. ISLDS. = Islands of the Southern hemisphere. WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 717 Table 17. Numbers in South Pulicidae TUNGINAE PULICINAE Pulicini Spilopsyllini Malacopsyllidae Rhopalopsyllidae RHOPALOPSYLLINAE Parapsyllini Rhopalopsyllini Ceratophyllidae CERATOPHYLLINAE FOXELLIINAE Leptopsyllidae AMPHIPSYLLINAE Ischnopsyllidae Vermipsyllidae Pygiopsyllidae Hystrichopsyllidae HYSTRICHOPSYLLINAE Hystrichopsyllini Ctenopariini STENOPONIINAE NEOPSYLLINAE Neopsyllini Phalacropsyllini ANOMIOPSYLLINAE Jordanopsyllini Anomiopsyllini RHADINOPSYLLINAE Corypsyllini Rhadinopsyllini Wenzellini DORATOPSYLLINAE Doratopsyllini Tritopsyllini CTENOPHTHALMINAE Ctenophthalmini Carterettini Neotyphloceratini Agastopsyllini Stephanocircidae CRANEOPSYLLINAE of Genera (bold face) and Species (roman) of Native Fleas Middle, and North America, according to Family. S.AM. PAN. MEX. M.AM. N.AM. Distribution 3/14 2/2 2/2 3/3 NEO., PAL. 2/3 2/5 2/5 2/2 M.AM., HOL., ETH., AUS. 2/3 1/1 2/4 2/4 2/6 HOL., NEO. 2/2 S.AM. 8/39 . . . . . . . . S.AM., AUS., & SO. ISLDS. 3/38 2/7 2/3 2/11 1/2 NEO., NE. 4/12 5/12 13/43 13/54 17/111 HOL., ETH., OR., NEO. 2/5 2/5 2/18 NE. 5/9 1/3 1/2 3/3 1/1 1/1 1/5 1/1 2/3 1/4 7/26 1/1 •• 5/6 HOL. 2/2 3/4 3/8 COS. 1/5 HOL. 1/1 AUS., OR., I 2/3 2/3 2/7 HOL. S.AM. 1/1 1/1 1/2 HOL. 1/1 1/1 2/2 HOL., OR. 2/7 2/6 5/38 HOL. 1/1 NE. •• 5/26 HOL. 2/10 NE. 1/1 1/1 3/11 HOL. •• 1/2 •• M.AM. 1/1 1/1 2/5 HOL. 1/1 1/1 •• NEO. 1/4 1/4 1/1 HOL. 1/2 NE. S.AM. S.AM. AUS., NEO. NEO. 718 ECTOPARASITES OF PANAMA fauna of South American ectoparasites; 3), moved back (Oryzomys, Sig- modon, etc.) into Middle and North America with their newly acquired para- sites, where 4) , they were isolated long enough to differentiate (as did their parasites) into species that are distinct from the South American ones now inhabiting the lowlands in Panama and elsewhere in Middle America. With very few exceptions, the ectoparasites of these complex-penis-type cricetine rodents belong to families and tribes or genera which are either restricted to or centered in South America, and whose closest relatives are in most cases Old World forms, especially of the Australian Region, but also of the Oriental and (to a much lesser extent) Ethiopian Regions. Only a very few parasites that are identifiable as Middle or North American, and these are obviously fairly recent intruders, occur on these hosts. Among these are the laelaptine mite Eubrachylaelaps rotundus, a few fleas of the genus Pleochaetis, another of the genus Kohlsia,23 and several rabbit fleas of the genera Cediopsylla and Hoplopsyllus (Subg. Euhoplopsyllus) . The geographic and host distributions of lice like Hoplopleura, cited by Vanzolini and Guimaraes (1955) , must be re-examined. The complex-penis-type Cricetinae, like Oryzomys and Sigmodon, that occur north of Panama, can hardly be relicts of an old fauna that dispersed into South America, if their fleas (Polygenis) are an indication. Polygenis is an "expanding" South American genus which has, quite clearly, dispersed into Middle America along with complex-penis-type Cricetinae and cavio- morph rodents from South America. Most of the Panamanian species probably dispersed very recently. In most cases they are not even subspe- cifically distinct from South American forms. Further, there seems to be little other reason to accept the Pliocene- Pleistocene transition as the principal time of dispersal 24 of the ancestral complex-penis-type cricetine rodents (and possibly some other mammals, too) into South America. Even if the fossil Cricetinae known from the Upper Pliocene of the Argentine (see following paper) reflect the first ap- pearance of these rodents in southern South America, this may mark the end of a long "trail" of dispersal and evolution rather than the beginning (see Hershkovitz, pp. 727-732). Because the implications of our distributional data appeared to conflict with prevailing views regarding the dispersal of the Cricetinae into South America, we discussed the problem with Mr. Philip Hershkovitz. His ac- count of their origin, dispersal and radiation (see following paper) gen- erally agrees with our conclusions regarding the ectoparasites. Acknowledgments We wish to express our gratitude to the following persons for reading 23 Many of the Ceratophyllidae are squirrel fleas (Hopkins 1957a). This is true of some species of Pleochaetis and Kohlsia, too. The few South American species of these genera may have immigrated on squirrels. 24 It is important to note that Nygren (1950, p. 2005. See also following paper by Hershkovitz, this volume, p. 725) believes that the Panama land bridge may have been available for passage by terrestrial animals since the close of the Middle Miocene! WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 719 portions of the manuscript and offering suggestions and corrections : Mr. Henry Dybas of Chicago Natural History Museum, for reading the section of the manuscript relating to coexistence and competitive displacement; Profs. Alfred E. Emerson and Lynn Throckmorton, of the Department of Zoology, of the University of Chicago, for reading the section on host specificity; Col. Robert Traub, U. S. Army (Ret.), School of Medicine, the University of Maryland, for reading the entire manuscript, and especially the portions dealing with the fleas; and, Mr. Philip Hershkovitz, Chicago Natural History Museum, for reading the entire paper as it relates to mam- mal hosts. The geologic history, zoogeography and ecology of the cricetine hosts have been discussed at length with Mr. Hershkovitz. Abstract The authors discuss host specificity, coexistence and competitive displacement, alti- tudinal distribution, some epidemiological considerations, and faunal relationships and their zoogeographic implications. The need for new approaches to field sampling and analysis of host and ectoparasite populations is emphasized. It is suggested that the degree of host-parasite specificity in ectoparasites is largely correlated with the degree to which the parasite is host-limited, i.e., restricted to the host, during its life cycle. Thus, a high degree of host specificity occurs most commonly among hemimetabolous groups whose life cycle is spent on the host ; among holometabolous forms whose free-living early stages have been eliminated through ovoviviparity and thus re- main closely associated with the host most of the time; or holometabolous forms which are closely confined with the host by the physical nature of its "home", as e.g., in kangaroo rat burrows, pocket gopher burrows, etc. It is further suggested that in host-limited forms, homozygosity for host specificity is achieved quickly, and likewise, speciation and niche specialization, because of inbreeding. This is in contrast with the situation in those ectoparasites which are non-host-limited, like most fleas and heteroxenous ticks, in which there is extensive outbreeding. It is further suggested that polyhaematophagy is selected for in these non-host-limited forms. In such non-host-limited forms either a broadly adaptive genetic variability or balanced polymorphism as regards host specificity would greatly increase the chances of host-finding and thus of survival. A case of coexistence and possible competitive displacement among streblid batflies parasitic on Phyllostomus hastatus is discussed. The data indicate that the altitudinal distribution of host-limited forms parallels that of the hosts closely, while there is a notable lack of concordance between that of non-host-limited parasites and their hosts. The altitudinal distributions of the ectoparasites when correlated with their system- atics and geographic and host relationships indicate that : 1 ) , the tropical lowland faunae of Panama and northern South America are virtually identical, but endemism increases correspondingly with increase in altitude, and is marked in the temperate zones, both altitudinally and latitudinally ; 2), this is largely true of the lowlands north of Panama, too, but here species endemism is evident; 3), the temperate (including the montane) fauna of South America is largely precinctive with considerable endemism at the family, subfamily and tribal level, and is Old World, especially Southern Hemis- phere in its relationships; 4), the montane fauna of Middle America likewise shows con- siderable endemism, but chiefly at the generic level, and its relationships are overwhelm- ingly with the Holarctic Region; 5), very little interchange is evident between the temperate faunae of the two continents, excepting parasites of such vagile hosts as birds, bats, and squirrels; 6), recent dispersals of ectoparasites of small mammal hosts, espe- cially Cricetinae, appear to have been chiefly from South to Middle, rather than from Middle to South America. The data appear to conflict with the views of Simpson (1950) and Patterson (1957, fig. 9), regarding the dispersal and radiation of the Cricetinae in South America, during the Pliocene-Pleistocene. It is suggested that dispersal of these rodents into South America took place in the Miocene or earlier. References BAKER, ROLLIN H. 1963. Geographical distribution of terrestrial mammals in Middle America. Amer. Midi. Nat., 70: 208-249, 1 text fig., 25 tables. BARNES, ALLEN M. 1965. Three new species of Anomiopsyllus ( Siphonaptera : Hystrichopsyllidae). Pan-Pac. Ent., 41, (4), pp. 272-280, 2 text figs. CARSON, HAMPTON L. 1959. Genetic conditions which promote or retard the formation of species. In Genetics and Twentieth Century Darwinism. Cold Spg. Harb. Symp. Quant. Biol., 24:87-105, 5 text figs. CAULLERY, MAURICE 1952. Parasitism and Symbiosis. London, Sidgwick and Jackson Ltd. xii + 340 pp., 80 text figs. CLAY, THERESA 1949. Some problems in the evolution of a group of ectoparasites. Evolution, 3: 279-99, 4 text figs., 9 tables. 1957. The Mallophaga of birds. In First Symposium on host specificity among parasites of vertebrates. Neuchatel. Pp. 120-158, 2 pis., 3 text figs., 10 tables. DEBACH, PAUL 1966. The competitive displacement and coexistence principles. Ann. Rev. Ent., 11: 183-212. DORF, ERLING 1959. Climatic changes of the past and present. Contr. Mus. Palaeont., Univ. of Mich., 13: 181-210, 1 pi., 3 text figs., 7 maps. EMERSON, ALFRED E. , 1954. Dynamic homeostasis: a unifying principle in organic, social, and ethical evolution. Scient. Month., 78: 67-85. 1960. The evolution of adaptation in population systems. In Evolution after Darwin. University of Chicago Press. 1 [The evolution of life, its origin, history and future] , pp. 307-348. FAIRCHILD, GRAHAM B. 1966. Introduction. In Ectoparasites of Panama, pp. 1-7. 720 WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 721 FURMAN, DEANE P. 1955. Revision of the genus Eubrachylaelaps (Acarina: Laelaptinae) with descrip- tions of two new species from Mexico. Ann. Ent. Soc. Amer., 48, (1-2), pp. 51-59, 18 text figs. FURMAN, DEANE P., AND TIPTON, VERNON J. 1961. Acaros parisitos Laelaptinae (Acarina: Laelaptidae) de Venezuela. Mem. Soc. Cienc. Nat. La Salle, 21, (60), pp. 166-212, 12 pis. GOLDMAN, E. A., AND ZETEK, JAMES 1926. Panama. In [Shelford, V. E., editor] Naturalists' Guide to the Americas. Baltimore, Williams & Wilkins Co. Pp. 612-622, 1 fig. GUIMARAES, LlNDOLPHO R. 1966. Nycteribiid batflies from Panama. In Ectoparasites of Panama, pp. 393- 404, text figs. 35-37. GUIMARAES, LINDOLPHO R., AND D'ANDRETTA, M. A. V. 1956. Sinopse dos Nycteribiidae (Diptera) do novo mundo. Arq. Zool., Sao Paulo, 9: 1-184, 232 text figs., 5 maps, 3 tables. HAAS, GLENN E. 1965. Comparative suitability of the four murine rodents of Hawaii as hosts for Xenopsylla vexabilis and X. cheopis (Siphonaptera). Jour. Med. Ent., 2, (1), pp. 75-83. HERSHKOVITZ, PHILIP 1958. A geographic classification of neotropical mammals. Fieldiana: Zoology, 36: 579-620, text figs. 7-8, 18 tables. 1962. Evolution of neotropical cricetine rodents (Muridae) with special reference to the phyllotine group. Fieldiana: Zoology, 46. 524 pp., 123 text figs., 65 tables. 1966a. Mice, land bridges, and Latin American faunal interchange. In Ectopara- sites of Panama, pp. 725-751, text figs. 151-4. 1966b. South American swamp and fossorial rats of the scapteromyine group (Cricetidae, Muridae) with comments on the glans penis in muroid taxonomy. Zeitschr. Saugetier Kunde, 30, heft. 1 (In press). HOLDRIDGE, R. R., AND BUDOWSKI, G. 1955. Report of an ecological survey of the Republic of Panama. (Technical Coop- eration program of the I.A.I.A.S. of the O.A.S.). Unpublished. HOLLAND, GEORGE P. 1964. Evolution, classification, and host relationships of Siphonaptera. Ann. Rev. Ent., 9: 123-146, 1 text fig. HOOPER, EMMET T., AND MUSSER, GUY G. 1964. The glans penis in neotropical cricetines (Family Muridae) with comments on classification of muroid rodents. Misc. Publ. Univ. Mich. Mus. Zool., no. 123. 57 pp., 9 text figs. HOPKINS, G. H. E. 1957a. Host associations of Siphonaptera. In First Symposium on host specificity among parasites of vertebrates. Neuchatel. Pp. 64-87. 1957b. The distribution of Phthiraptera on mammals. Ibidem, pp. 88-119. HOPKINS, G. H. E., AND ROTHSCHILD, MIRIAM 1953. An illustrated catalogue of the Rothschild Collection of fleas (Siphonaptera) in 722 ECTOPARASITES OF PANAMA the British Museum (Natural History), etc. 1. Tungidae and Pulicidae. xiv + 362 pp., 45 pis., 466 text figs. 1956. Idem, 2. Coptopsyllidae, Vermipsyllidae, Stephanocircidae, Ischnopsyllidae, Hypsophthalmidae and Xiphiopsyllidae. xi + 446 pp., 32 pis., 708 text figs. 1962. Idem, 3. Hystrichopsyllidae (Acedestiinae, Anomiopsyllinae, Hystrichopsyl- linae, Neopsyllinae, Rhadinopsyllinae and Stenoponiinae). ix + 559 pp., 10 pis., 1050 text figs. JOHNSON, PHYLLIS T. 1957. A classification of the Siphonaptera of South America. Mem. Ent. Soc. Wash., no. 5. 6 -f 298 pp., 114 pis. JOHNSON, PHYLLIS T., AND LAYNE, JAMES N. 1961. A new species of Polygenis Jordan from Florida, with remarks on its host re- lationships and zoogeographic significance. Proc. Ent. Soc. Wash., 63: 115-123, 10 text figs. RUNS, MERLE 1964. Ecological studies on the Department of Beni, 1963 to 1964. 8 ms. pp., 1 table (unpublished report given in La Paz, July 26, 1964). LEWONTIN, R. C. 1957. The adaptations of populations to varying environments. In Population studies: animal ecology and demography. Cold. Spg. Harb. Symp. Quant. Biol., 22: 395-408, 7 text figs. MACHADO-ALLISON, CARLOS E. 1963. Revision del genero Amblyopinodes Seevers, 1955 (Coleoptera, Staphylinidae) . Acta Biol. Venez., 3: 371-416, 108 text figs. MAYR, ERNST 1957. Evolutionary aspects of host specificity among parasites of vertebrates. In First symposium on host specificity among parasites of vertebrates. Neuchatel. Pp. 7-14. MEAD-BRIGGS, A. R., AND RUDGE, A. J. B. 1960. Breeding of the Rabbit Flea, Spilopsyllus cuniculi (Dale) : requirements of a "factor from a pregnant rabbit for ovarian maturation." Nature, 187, (4743), pp. 1136-1137, 2 figs. NYGREN, W. E. 1950. Bolivar geosyncline of northwestern South America. Bull. Amer. Assn. Petr. Geologists, 34: 1996-2006, 3 text figs. PATTERSON, BRYAN P. 1957. Mammalian phylogeny. In First symposium on host specificity among para- sites of vertebrates. Neuchatel. Pp. 15-49, 11 text figs. SEEVERS, CHARLES H. 1955. A revision of the tribe Amblyopinini : staphylinid beetles parasitic on mam- mals. Fieldiana, Zoology, 37: 211-264, figs. 28-44. SIMPSON, GEORGE G. 1950. History of the fauna of Latin America. Amer. Scientist, 38, (3), pp. 361-389, 1 text fig. SMIT, F. G. A. M. 1963. A description of the male of Idilla caelebs, with a discussion of the relationships WENZEL AND TIPTON : HOST-PARASITE RELATIONSHIPS 723 of this species ( Siphonaptera : Hystrichopsyllidae). Proc. Roy. Ent. Soc. Lond., 31:109-114, 2 text figs. TIPTON, VERNON L. 1960. The genus Laelaps, with a review of the Laelaptinae and a new subfamily Alphalaelaptinae. Univ. Calif. Publ. Ent., 16: 233-356, pis. 22-47. TIPTON, VERNON J., ALTMAN, ROBERT M., AND KEENAN, CHARLES M. 1966. Mites of the Subfamily Laelaptinae in Panama. In Ectoparasites of Panama, pp. 23-81, pis. 1-36, tables 1-3. TIPTON, VERNON J. AND MENDEZ, EUSTORGIO 1966. The fleas (Siphonaptera) of Panama. In Ectoparasites of Panama, pp. 289- 385, pis. 47-93, text fig. 34. TRAUB, ROBERT 1952. Atyphloceras tancitari and Jellisonia bonia, new species of fleas from Mexico (Siphonaptera). Amer. Mus. Novit., no. 1558. 19 pp., 24 text figs. 1953. Wenzella obscura, a new genus and species of flea from Guatemala. Jour. Wash. Acad. Sci., 43: 77-85, 10 text figs. WARD, RONALD A. 1957. A study of the host distribution and some relationships of Mallophaga para- sitic on birds of the Order Tinamiformes. Part I. Ann. Ent. Soc. Amer., 50: 335- 353, 1 table, 20 text figs. Part II. Ibidem, pp. 452-459, 1 text fig., 4 tables. VANZOLINI, P. E., AND GUIMARAES, LINDOLPHO R. 1955. Lice and the history of South American land mammals. Rev. Brasil. Ent., 3: 13-45, 4 text figs. WENZEL, RUPERT L., TIPTON, VERNON J., AND KIEWLICZ, ALICJA 1966. The streblid batflies of Panama (Diptera Calypterae: Streblidae). In Ecto- parasites of Panama, pp. 405-670, text figs. 38-146, tables 7-10. YUNKER, CONRAD, AND STRANDTMANN 1966. The Genus Hirstionyssus in Panama. In Ectoparasites of Panama, pp. 725- 751, text figs. 151-154. Mice, Land Bridges and Latin American Faunal Interchange1 PHILIP HERSHKOVITZ 2 REVIEW OF THE HISTORY OF SOUTH AMERICAN MAMMALS The history of South American mammals begins with the Paleocene records from Argentina. The order Marsupialia is represented here by three superfamilies. The didelphoid opossums are known from abundant remains but the family Didelphidae was in North America since the Cretaceous. The carnivore-like borhyaenoids which compose the second superfamily were the principal predators of the South American Tertiary. Fossils of the third superfamily, the small shrew-like caenolestoids, are known from the Eocene of Argentina but species of this group were present in southeastern Brazil during the upper Paleocene. Remains of armadillos which were similar to living species of the edentate order Xenarthra ap- pear in the Argentine Paleocene, while fossils of related glyptodonts, ground sloths, and anteaters, are of a later period in the Tertiary. Members of the extinct order of primitive ungulates, the Condylarthra, and five inti- mately related orders of peculiar South American hoofed animals had al- ready attained a high level of diversity during the Paleocene. The Notoungulata radiated into animals ranging in size and appearance from the rhinoceros-like to the hare-like. The Litopterna evolved into forms resembling horses, camels, and small antelopes. Some of the Astrapotheria have been compared with rhinoceroses and hippopotami. Little is known of the Paleocene Xenungulata but the few recognized members of this order are among the oldest of South American mammals. The proboscidian-like Pyrotheria date from the Eocene and seem to have been the last of the native South American ungulates to evolve. Precisely how and when these animals first arrived in South America is unknown. Except for marsupials and condylarths, there is no record of 1 Preparation of this manuscript was aided by National Science Foundation Grant GB-2059. The figures were drawn by Josephine Stessl. 2 Division of Mammals, Chicago Natural History Museum, Chicago, Illinois. 725 726 ECTOPARASITES OF PANAMA the occurrence of such animals in Middle America or northern South Amer- ica prior to their appearance in southern South America. Knowledge of what are presently regarded as later invasions is similarly deficient. Rodents of the suborder Caviomorpha appeared nearly full bloom in the Oligocene of the southern half of the continent. Their progenitors must have invaded South America earlier but none of the steps leading to the build-up has been discovered. Monkeys of an organization quite like that of living South American primates appeared as suddenly in the upper Oligocene of southern Patagonia. Nothing is known of their platyrrhine ancestry. There is no record of the evolution and dispersal of neotropical Primates from early Tertiary prosimians of the middle latitudes of North America to Oligocene ceboids of the middle latitudes of South America. The next records of new faunal elements in the South American Tertiary date from middle or perhaps early Miocene with the appearance of the dol- phin Proinia True, a genus supposedly of the family Susuidae (order Cetacea) . Two genera of this family survive in South American waters with one, Inia, confined to fresh water. Whatever the systematic position of Proinia (cf. Kellogg, 1942) of the Patagonian marine deposits, the Pliocene Ischyorhynchus Ameghino is a true river susuid and very nearly related to Inia. The La Venta fossils discovered in Central Colombia (cf. Stirton, 1953) give an inkling of the great variety and wide dissemination of South Amer- ican mammals by late Miocene. They include remains of many of the southern forms already known and, in addition, fragments of true cebid monkeys, a bat of the family Phyllostomidae (Chiroptera), and a river manatee of the family Trichechidae (order Sirenia). Late Miocene or early Pliocene saw the arrival of members of the raccoon family, Pro- cyonidae (order Carnivora). Finally in the upper Pliocene, mice of the murid subfamily Cricetinae (Myomorpha, order Rodentia) had already evolved into forms indistinguishable from representatives of living sigmo- donts. Evidently, they thrived in great numbers on the plains of north- eastern Argentina together with herds of peccaries (Tayassuidae, order Artiodactyla) which had migrated from the north. Skunks of the family Mustelidae (order Carnivora) may also have invaded South America at this time but the oldest fossil, discovered in Eastern Buenos Aires, may be early Pleistocene rather than late Pliocene. Thus, by the end of the Tertiary, fossil evidence reveals that South America was inhabited by three superfamilies of the order Marsupialia, two of which survive ; five superfamilies of the order Xenarthra of which two are now extinct; six orders of ungulates, five native and none with living issue; two families of the order Prim'ates; one family of Cetacea with a genus confined to fresh water; one family each of Chiroptera, Sirenia, Carnivora, and two suborders of Rodentia, the still flourishing Caviomorpha and Myomorpha. The oldest records of all but three of these taxa are from the southern half of the continent. The ancestors of all South American orders or families that originated in North America left no sign of their sojourn in Middle America. With exceptions noted, their first passage through northern South America is also unrecorded. It is practically cer- HERSHKOVITZ I LAND BRIDGES, FAUNAL INTERCHANGE 727 tain that additional mammalian families either invaded or arose in northern South America during the Tertiary but disappeared without trace. Still others now living in South America almost certainly arrived or evolved there during the late Tertiary but for lack of fossil evidence may be in- correctly adjudged to be post-Tertiary invaders. Bats may have been the dominant and most diversified of small South American mammals throughout the Tertiary but only one genus of Phyl- lostomidae is known from that period. The other endemic tropical American chiropteran families are the Noctilionidae, Desmodidae, Natalidae, Furip- teridae, and Thyropteridae. They were members of a very ancient fauna but only fragments of some of these have been found in Pleistocene owl pellets. Squirrels (Sciuridae) undoubtedly invaded South America several times from the north. The ancestor of the pygmy squirrel, Sciurillus, may well have been the first to arrive, perhaps simultaneously with Middle Tertiary Primates. Successive invasions of dogs, mustelids, and deer may also have occurred but at a later date beginning perhaps with middle or late Pliocene. Tapirs of the order Perissodactyla are known from the Pleistocene but the first may have reached South America during late Pliocene. The same may apply to bears and cats which are also known from the Pleistocene of South America. Horses and mastodons (order Proboscidea) arrived and disappeared during the Pleistocene but the camels survived.3 One genus of shrews, Cryptotis (Soricidae, Insectivora), one genus of rabbits, Sylvilagus (Leporidae, Lagomorpha) and one genus of spiny mouse, Heteromys (Heteromyidae, Rodentia) are among the latest arrivals and are representa- tives of the only other new mammalian families to invade South America since early Pleistocene. MICE, MOVEMENTS AND BARRIERS The advent of mice, or myomorph rodents, in South America is shrouded in mystery. A long history of differentiation in northern North America, Middle America, and South America preceded their appearance in the upper Pliocene of Argentina. Much of the evolution can be reconstructed from the morphology of living forms but the time of invasion and the routes of dispersal must be inferred from recent distributional patterns. New World mice, the Cricetinae, are recorded from the Oligocene of North America but possibly arose in late Eocene. They may have originated in Eurasia and spread into northern North America, then Middle America and South America. Or, they may have arisen in northern North America and spread northward into the Old World and southward into South Amer- ica. Insofar as the history of living cricetines is concerned, either hypothesis could be valid, but present knowledge favors the concept of a New World 3 The bovine Colombibos atactodontus Hernandez Camacho and Porta, said to be from the upper Pleistocene of Colombia, is based, according to Thenius (1964, p. 275, foot- note 4), on what seem to be the deciduous teeth of domestic cattle. 728 ECTOPARASITES OF PANAMA origin for cricetines. Whether the center was in northern or southern (i. e., Middle American) latitudes of North America cannot be determined now. It is known that the most primitive of living cricetines are sylvan and the 84 74 72 Fig. 151. — Axis of Bolivar syncline (bars) and western Tertiary borderland (diagonals) in Ecuador and Colombia with extension in Panama. (Adapted from Nygren, 1950, fig. 1) HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 729 ancestral form must also have been sylvan with all the morphological characters distinguishing them from pastoral forms (cf . Hershkovitz, 1962, p. 16). Judged by the North American fossil record, it may be assumed that all well defined Recent genera of pastoral cricetines were already dif- ferentiated in the Pliocene. Whatever its geological history during the Tertiary, Middle America must have served variously as a transition zone, a center of evolution and a staging area. From Middle America, cricetines and other New World mammals spread into higher northern latitudes in the wake of retreating glaciers and as suitable habitats became available. Their movements south- ward were largely controlled by geographic, tectonic, and ecological factors. Spread into South America was accomplished by means of over-water movements, i. e., island hopping and waifing, by swimming or fording; or passively, by shifting land connections or cutoffs. It was finally accom- plished via the Isthmian land bridge which only recently became complete at its South American terminal (fig. 152). Actual means or routes of dispersal used by the oldest migrants are hypothetical and need not have been the same or used at the same time by different individuals of the same species or different species. A broad reconstruction of the history of mammalian faunas in Latin America has been presented by Simpson (1950). His account served as basis for orientation and has been universally accepted as an article of faith. I reproduce herewith the principal section (pp. 379-381) relating to the intercontinental exchange of faunas and the dissemination of cricetines in South America : "A change like that going on in the Eocene on the World Continent also occurred in South America, but at a greatly later date, in the late Pliocene and Pleistocene . . . Its cause was the rise of the Central American bridge and the consequent irruption into South America of many derivatives from the fauna of the World Continent. This third broad faunal stratum did not come in all at once, in a single wave. Already in the late Miocene a few northern forms appeared, small arboreal placental carnivores more or less related to the raccoon. Not long there- after, apparently in early Pliocene times, some South American animals, ground sloths, reached North America. These forerunners do not seem to indicate a con- tinuous land connection but probably utilized the island chain, gaps in which were closing progressively as the Central American and northwestern South American regions rose relative to sea level. The exact moment when the bridge became com- plete is not established, but this probably occurred during the age called Chapad- malalan in South America and Blancan in North America, placed by some au- thorities as latest Pliocene and by others as earliest Pleistocene. Even then the exchange was at first rather limited in scope and the full surge of intermigration did not occur until somewhat later, in unequivocally Pleistocene times. Soon or late, at least fifteen (possibly sixteen) families invaded South America in this great episode . . . "Invasion occurred in both directions. By a moderate tabulation, fifteen fam- ilies of North American mammals then spread into South America and seven families spread in the reverse direction. The main migrants to the south were rabbits, squirrels, field mice, dogs, bears, raccoons, weasels, cats, mastodons, horses, tapirs, peccaries, camels, and deer, including in most of these cases some variety of related forms." The spectacle of fifteen or sixteen North American families and seven 730 ECTOPARASITES OF PANAMA South American families awaiting the propitious moment "in unequivocally Pleistocene times" to cross the Panamanian bridge is dramatic. Viewed in the light of fossil records and what is known of the ecological preferences, habits, and vagility of the animals concerned, the stage seems real enough but the scene is less a product of fact than fancy. The geological and mammalian history of the Americas indicates that the Isthmian water gap was much less effective in isolating faunas than the climatic barrier across northern Mexico. For most mammals the water gap may not have been more than a minor obstacle and for some such as otters and other aquatic and subaquatic animals, an invitation. It was cer- tainly not as much a deterrent as water barriers crossed by mammals in reaching the Antilles, the Galapagos, the Philippines, Australia, New Guinea, and other islands well off the continental shelves. Evidently, many northern families which spread through the Isthmus to the edge of the gap crossed it opportunely. Herds of wandering mastodons, tapirs, horses, pec- caries, camels and deer would cross such bodies of water easily and routinely just as their living relatives do today. Island chains and rafts provided others with means of access to opposite shores. Unaquatic mammals such as ground sloths and related edentates were crossing the water gap since middle and probably early Pliocene while monkeys crossed at various times since the Oligocene or perhaps late Eocene. Despite the absence of fossil evidence, there is no reason to assume that many more families of contem- poraneous mammals, some better adapted to water than others, were not doing the same. The traffic was both active and passive and from either con- tinent to the other. Rodents habitually crossed water barriers and colonized continental and oceanic islands throughout the world. Latin American cricetines are no exception. The vole-like terrestrial or pastoral species seem to have been even more successful than their aquatic relatives in crossing water barriers and ranging widely. At the time of the Pliocene-Pleistocene transition when, according to Simpson's dictum, the intercontinental faunal exchange was about to begin, complex-penis-type genera and tribes of cricetines (see below) had already become differentiated in South America and reached nearly their present limits of natural dispersal in both North and South America. By late Pliocene, Sigmodon, the most generalized member of the South American sigmodont group, had invaded North America at least as far as Kan- sas (Hibbard, 1937, p. 247). At the southern extreme, late Pliocene Proreithrodon Ameghino (inseparable from Pleistocene Ptyssophorus Ameghino and Tretomys Ameghino apd from Recent Reithrodon Water- house) was already one of the most common rodents of Patagonia (Hersh- kovitz, 1955, p. 639 ff.). Other Pleistocene cricetines of northeastern Ar- gentina are also identical with living forms. The rich and well preserved fossil mammals of the Pleistocene of Minas Gerais, eastern Brazil, discov- ered in the first half of the last century by the Danish paleontologist Wilhelm Lund, are, for the most part, indistinguishable from living mammals of the same region. Closure of the Panamanian portal may have facilitated the crossing of HERSHKOVITZ: LAND BRIDGES, FAUNAL INTERCHANGE 731 Fig. 152. — Rio Atrato-Rio San Juan basin, western Colombia. Land bridge connecting North and South America across the western border of the former Isthmian seaway is the narrow divide between the two rivers. The river channels follow the axis of the Bolivar geosyncline. 732 ECTOPARASITES OF PANAMA some species but there is no evidence that it resulted in a spectacular in- crease in the volume of faunal interchange. THE CRICETINE GLANS PENIS IN TAXONOMY AND NEW WORLD ZOOGEOGRAPHY New and Old World cricetines (Cricetinae) and microtines (Microtinae) are here treated as subfamilies of Muridae. This classification combines, in effect, all categories of the Muridae and Cricetidae of Simpson (1945, pp. 83, 205) or Wood (1955, p. 176) into the single family Muridae as rec- ognized by Ellerman (1941, p. 1). Recent studies of the glans penis and male accessory reproductive glands support this classification and add considerable data to our knowledge of the interrelationships of many of the species, genera and supergeneric assemblages of murids. Two basic phallic types within the Muridae are demonstrated by Hooper (1958, 1959, 1960, 1962), Hooper and Hart (1962), Hooper and Musser (1964a, 1964b) and are discussed by Hershkovitz (1966). The first or complex penis type (fig. 153 A) is characterized by a baculum with typically three terminal digits and paired sac-like sinuses of the spongy tissue of the glans. This type predominates in Old World and South Amer- ican Cricetinae, Murinae, Gerbillinae, Microtinae, and probably others in- cluding otomyines and dendromurines. The simple type penis (fig. 153B) is distinguished by the absence of lateral bacular digits and sacculations in the spongy vascular layer of the glans. It characterizes the essentially Holarctic peromyscine Cricetinae comprising the thirteen genera, Peromyscus, Reithrodontomys, Aporodon, Onychomys, Baiomys, Scot- inomys, Ochrotomys, Neotomodon, Nelsonia, Ototylomys, Tylomys, Neot- oma, and Xenomys. Complex and simple phallic types are not restricted to murids and the simple type is not confined to peromyscines. Hooper and Musser (1964a) demonstrate simple or intermediate phallic types among some Old World cricetines (Myospalax, Mystromys) , South American cricetines (Nyctomys, Otonyctomys, Scapteromys), nesomyines (Macrotarsomys) and microtines (Dicrostonyx) , some species of Microtus, Ellobius (cf. Hooper and Hart, 1962) . Hooper and Musser (1964a, p. 53) also mention a simple type penis among murines but do not identify the species. The simple penis is derived from the complex and arose independently in all major murid categories (cf. Hershkovitz, 1966). Simplification may be effected by reduction and elimination of the lateral bacular digits, as has occurred in all peromyscines, some murines (Bittera, 1918, p. 414), some microtines (Hooper and Hart, 1962), and some South American cricetines (Hooper and Musser, 1964a; Hershkovitz, 1966). It may also result from reduction and elimination of the middle digit only, as in the South American Scapteromys, Phyllotis, and Zygodontomys (Hershkovitz, 1966) ; or by fusion of all digits, as in most Scapteromys (Hershkovitz, 1966). Members of the peromyscine group are a diverse lot judged by external, cranial, and dental characters. The penis, although of the simple type is HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 733 also highly diversified within the group. Hooper and Musser (1964a) divide peromyscines into two groups. Earlier, Hooper (1960) had arranged them in four. The several convergent paths leading to a simple penis increase the probability that peromyscines are polyphyletic. In any case, it is vir- tually certain that peromyscines, individually or collectively, are derived from the same complex penis type stock now represented by South Ameri- can cricetines. These two assemblages, the essentially North American 2mm COMPLEX A 4mm SIMPLE B Fig. 153. — Glans penis of New World cricetines. A, complex type with three digitate baculum in Oryzomys flavescens (after Hooper, and Musser, 1964a, fig. Ih). Glans of A dissected midventrally to expose baculum. B, simple type with simple rod-like baculum in Peromyscus pirrensis (after Hooper and Musser, 1964b, fig. le). Glandes cleared and stained to reveal internal structures. peromyscines with simple penis and the essentially South American crice- tines with complex penis, are the primary indicators of inter- American ex- changes of mammalian faunas during the late Cenozoic. Their separate dispersal patterns form a mosaic of five faunal strata. Old North and Middle American cricetines, representing stratum I (fig. 154) migrated over water, i. e., by island hopping or waifing, into South America and radiated there into representatives of stratum II. Descend- ants of some elements of stratum II returned by over water routes to Middle America and differentiated significantly forming stratum III. Finally, late migrants of South American stock spread over the completed Isthmian land bridge and are identified as stratum IV. North American elements also spread at the same time by the same route into South America and be- 734 ECTOPARASITES OF PANAMA came stratum V. There has been little or no change in the taxonomic grade of either stratum IV or V. Many mammals and other vertebrates belong to the same faunal strata as the cricetines. The following section, however, describes the five strata in terms of cricetines only. LATIN AMERICAN CRICETINE FAUNAL STRATA STRATUM I. OLD MIDDLE AMERICAN COMPLEX PENIS TYPES. Old complex penis types; evolved in situ or migrated from northern North America (fig. 154). The only Middle American members of this stratum are the following two closely related monotypic genera, Nyctomys Otonyctomys These are possibly relicts of the archaic North or Middle American stock from which the complex-penis-type thomasomyines may be derived on the one hand, and from which the simple-penis-type peromyscines diverged on the other. The male genital tract and the accessory reproductive glands (Arata, 1964, p. 14) of Nyctomys sumichrasti and Otonyctomys hatti are reduced or simplified and approach those of peromyscines. Other- wise, they most nearly resemble species of the South American genera Thomasomys and Rhipidomys. The possibility that the stem form of Nyctomys and Otonyctomys evolved in South America and migrated over water into Middle America (see stratum II) must also be considered. The Tertiary history of Middle American cricetines is unknown. There is no suggestion that Eumys of northern North American Oligocene may be ancestral. According to Clark, Dawson and Wood (1964, p. 42) some element of the Eumys complex may have given rise to "hesperomyines." Their phylogenetic chart (op. cit, fig. 5) shows a line leading to Leidymys (Oligocene-early Miocene) through Copemys (late Miocene-early Pliocene) to Peromyscus (upper Pliocene and later). Another branch leads from Leidymys to Miochomys (late Miocene) then directly to Onychomys of the same horizons as Peromyscus. It seems that the "hesperomyines" of these authors equals the peromyscines of neomammalogists. The name pero- myscine, or the tribal form Peromyscini, is zoologically correct and nomen- clatorially valid for North American simple-penis-type cricetines (see nomenclatorial note p. 746) . STRATUM II. SOUTH AMERICAN COMPLEX PENIS TYPES. Descendants of stratum I; migrated over water from Middle America into South America (fig. 154) ; evolution to generic and tribal grades. There are approximately 50 genera and 300 species of South American cricetines. Samplings of a large number of genera (cf. Hooper and Musser, 1964a) show all to be of the complex penis type. The affinities of the re- maining genera are such as to virtually preclude the possibility that any of them are not of the same basic type. Ancestral species arrived in South America during the Tertiary, perhaps from Miocene onward. Subsequent intercontinental movements of complex-penis-type cricetines may have been, until late Pliocene-Pleistocene, almost entirely from South America into North America (see strata III and IV, beyond). HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 735 Fossil cricetines congeneric with living South American forms are known from the Pleistocene of southeastern Brazil and the late Pliocene and Pleistocene of northeastern Argentina. There is no likelihood that the evolution of the more highly specialized and geographically restricted species and generic clusters could have taken place anywhere else than in situ. The evolution of most of these forms can be reconstructed from the extant South American fauna. For a discussion of the adaptive radiation, dispersal, and taxonomy of some of the genera and generic groups see Hershkovitz (1944, 1955, 1960, 1962, 1966). OLIGOCENE; MIOCENE PLIOCENE PLE I STOCENE Fig. 154. — Cricetine rodent faunal strata (Roman numerals) in Middle and South Amer- ica. Broken arrows indicate over water migration, solid arrows over land. Stratum I = old North American complex-penis-type mice. Stratum II = tribes and genera of South American complex-penis-type mice evolved from I. Stratum III = genera and species of Middle American and West Indian complex-penis-type mice differentiated from II. Stratum IV = slightly differentiated Middle American descendents of II. SP = North American simple-penis-type mice. Stratum V = slightly differentiated simple-penis-type migrants in South America. STRATUM III. OLD SOUTH AMERICAN MIGRANTS IN NORTH AMERICA. Early descendants of stratum II; migrated over water into Middle America and the West Indies (fig. 154) ; evolution to species and generic grades. Three genera, Sigmodon, Oryzomys and the extinct oryzomyine genus Megalomys compose this stratum. They are of the complex penis type and derived from South American stock. Sigmodon is represented by several extinct and living species, all closely related and treated here as a single species complex with an intricate geographic history. The genus Oryzomys 736 ECTOPARASITES OF PANAMA of this stratum includes ten species, one extinct, and two subgenera includ- ing the nominate form. The West Indian Megalomys is known from the Pleistocene and the Recent but was exterminated in historic time. Remains of one species have been found near those of ground sloths. The North American migrations of Oryzomys and Sigmodon also coincided largely with the movements of ground sloths, glyptodonts and giant armadillos. The species groups of Sigmodon and Oryzomys are listed below in the order of the magnitude of their geographic range from north to south. Sigmodon hispidus complex The North American range of Sigmodon is greater than that of any other complex- penis-type cricetine. Its present and past distribution in North America points to the cotton rat as one of the earliest over-water migrants from South America. It has been recorded from the upper Pliocene of Kansas (S. intermedius Hibbard, 1937, p. 247) and S. hispidus was in Florida during Pleistocene. Successive invasions from South Amer- ica, or reinvasions of South America from Middle America, may account for the con- tinuous distribution of livirg Sigmodon hispidus in Middle and northern South America. The extinct species of North American Sigmodon are properly components of strata III. On the other hand, Recent and Pleistocene S. hispidus may be a late arrival and an element of strata IV. The equal if not greater probability is, however, that Sigmodon hispidus has been a persistent and pervasive element in all faunas from strata II to V. The sigmodont upper right molar described but not named by Black (1963, p. 485) from the early Miocene Thomas Farm locality in Gilchrist County, Florida, is tantalizing. The enamel pattern of the tooth most nearly resembles that of Sigmodon. The occlusal surface, however, is said to be terraced. The molars of Sigmodon are plane, hence more specialized. The Thomas Farm tooth may represent a true sigmodont, a parallel development or, less likely, an extinct side issue of the -E/wmi/s-peromyscine line. Oryzomys palustris complex palustris (= couesi, azuerensis, cozumelae, gatuncnsis, antillarum) Fossils identified as O. palustris are known from the Pleistocene (Illinoisan?) of Florida (Gut and Ray, 1963, p. 325). Examination of the type of Oryzomys antillarum Thomas from Jamaica shows it to be a weakly differentiated member of the palustris group. It may be treated as a subspecies, Oryzomys palustris antillarum Thomas. In reaching Jamaica as a waif it crossed a much wider water gap than any separating the Americas at the Isthmus during the Tertiary. peninsulae fulgens nelsoni fossilis (extinct) Described by Hibbard (1955, p. 213) from the Pleistocene interglacial (Sanga- mon?) of Meade County, southwestern Kansas. The genus Oryzomys does not exist in Kansas now, but Hall (1955, p. 246) notes a specimen of O. palustris in the U. S. National Museum said to have been collected in Neosho Falls, south- eastern Kansas, by B. F. Goss, in 1859. Oryzomys melanotis If this proves to be a member of the O. alfaroi group it belongs in faunal stratum IV (below). Subgeneric affinities undetermined. Oryzomys bombycinus Subgeneric affinities undetermined but near O. melanotis. Oryzomys aphrastus Known from the type specimen only, a skin with shattered skull. Its nearness to HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 737 Nectomys (Sigmodontomys) alfari suggests that aphrastus might belong to faunal stratum IV. Oryzomys (Micronectomys) dimidiatus Micronectomys is known from the type specimen only. Should representatives of 0. dimidiatus be discovered in South America it would be treated as a member of faunal stratum IV.4 With the possible exceptions noted, the degree of divergence of the Mid- dle American species of Oryzomys from their South American stems im- plies a long isolation. The morphological differences suggest polyphylety and their geographic distribution indicates successive invasions. Mem- bers of the O. palustris complex have spread farthest north and are the most differentiated from South American species. They are known from the Pleistocene of southern and central United States. The more southern species of Oryzomys are more closely related to their South American rela- tives. Megalomys curazensis Described by Hooijer (1959) from the late Pleistocene of Curasao. It was con- temporaneous with Paulocnus Hooijer (1962), a megalonychid ground sloth. Megalomys desmarestii Known from the Lesser Antillean islands of Martinique, Santa Lucia and Barbuda (Pleistocene). The species has been exterminated. Its nearest living relative ap- pears to be the scansorial Oryzomys (Macruroryzyomys) hammondi of northwestern Ecuador. Oryzomys pliocaenicus Hibbard (1939:539) from the Edson Quarry Pliocene of Kansas is not an Oryzomys according to Hibbard (in litt.). Dr. Hibbard adds that he re-examined the type in 1952 and believes it may prove to be a species of Bensonomys Gazin. Eligmodontia arizonae Gidley, 1922, from the San Pedro Valley Pliocene in Arizona, was made type of the genus Bensonomys by Gazin (1943, p. 489 ) . It has nothing to do with Eligmodontia Waterhouse, a genus of gerbil- like mice of the southern half of South America.5 STRATUM IV. LATE SOUTH AMERICAN MIGRANTS IN MIDDLE AMERICA Late descendants of stratum II; spread over Panamanian land bridge into Middle America (fig. 154) ; low grade subspeciation. Thirteen complex-penis-type species representing seven cricetine genera are common to North and South America. Their geographic and genetic continuity can be traced across the Panamanian land bridge. Sigmodon hispidus Zygodontomys brevicauda (see Hershkovitz, 1962:203, for taxonomy) Rhipidomys scandens (genus unrevised and Colombian representative of scandens not presently determined) Nectomys alfari 4 "Oryzomys (Micronectomys) borreroi" Hernandez, from the Rio Chucuri Valley, Santander, Colombia, is undoubtedly an Oryzomys but, judged by the original descrip- tion, it is certainly not a Micronectomys. 5 Simpson (1945, p. 84) overlooked the Gazin revision and includes "Eligmodontia" in the upper Pliocene fauna of North America. 738 ECTOPARASITES OF PANAMA Neacomys tenuipes (includes pictus) Rheomys trichotis (includes thomasi, underwoodi, hartmanni and raptor) Oryzomys alfaroi Oryzomys capito (= talamancae) Oryzomys albigularis (= devius, pirrensis) Oryzomys concolor (= tectus) Oryzomys bicolor (= endersi, trabeatus) Oryzomys caliginosus Oryzomys nigripes complex (subgenus Oligoryzomys; includes fulvescens, delicatus, victus, longicaudatus, delticola and others) Of the species listed above, only Sigmodon hispidus has a range extend- ing north of the Middle American province. Its distribution through f aunal strata II-IV inclusive, is discussed in the preceding section. The vole-like Sigmodon and Zygodontomys are true pastoral forms. The ichthyomine Rheomys trichotis is also derived from a pastoral stock but is highly spe- cialized for aquatic life and fish eating. The remaining species are sylvan. Rhipidomys is arboreal, Nectomys subaquatic, Neacomys terrestrial and the species of Oryzomys vary from strictly terrestrial (O. albigularis and the vole-like O. caliginosus) to the arboreal O. concolor. Members of the un- revised Oryzomys nigripes complex range from southern Mexico to the straits of Magellan. They are terrestrial and scansorial and live in for- ests, woodlands, bordering fields and open savannas from sea level to above tree line in the Andes. Oryzomys victus Thomas, from Saint Vincent, Les- ser Antilles, is a well differentiated Oligoryzomys, geographically nearest 0. delicatus of Trinidad. Unless recently imported, it represents faunal stratum IV in the West Indies. Faunal flow of stratum IV from South America to North America via the Isthmian land bridge seems to have been greater in volume and richer in variety than the flow of stratum V in the opposite direction. STRATUM V. LATE MIDDLE AMERICAN MIGRANTS IN SOUTH AMERICA Late descendants of North American simple penis types (peromyscines) ; spread over Panamanian land bridge into South America contemporaneously with IV (fig. 154) ; low grade subspeciation. The thirteen (to sixteen) recognized genera of autochthonous North American cricetines, not including Nyctomys and Otonyctomys (see stratum 1, above) , are peromyscines. They are characterized by a simple penis and include about 150 species. The geographic range of each of seven of the genera, namely, Tylomys, Ototylomys, Aporodon, Reithrodontomys, Peromyscus, Scotinomys, and Neotoma, extends into or is restricted to Mid- dle America. Most of the fifty to sixty species assigned to these seven genera in Middle America are unrevised. Only the following two reached South America. Tylomys nudicaudatus (includes gymnurus, panamensis, watsoni, tumbalensis) Aporodon mexicanus The two species are scansorial, the first frankly arboreal. Both range into the Andes of Colombia and northern Ecuador, and neither is more than subspecifically differentiated from its nearest Middle American relative. As in the case of the contemporaneous spe- cies of stratum IV (above), the geographic and genetic continuity between Middle and South American representatives of these species point to the land bridge migration route. HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 739 Peromyscus pirrensis Goldman appears to be a relict species confined to the Panamanian side of the Serrania del Darien. Hall and Kelson (1959, p. 653, map 374) indicate without documentation that the species occurs on the eastern or Colombian side of the Serrania. Absence of simple-penis-type cricetines in South America except for the two recent invaders proves conclusively that this group originated in North America, perhaps in Mexico. As compared with Tertiary over water crossings, faunal flow across the Isthmian bridge from North America into South America was anticlimactic. WATER GAPS, BRIDGES, AND TERTIARY FAUNAL INTERCHANGE The American continents during Tertiary were at times connected, at others separated, by a waterway or a complex of islands and straits. The final and crucial barrier separating the faunas of Middle America and South America was the Bolivar geosyncline (fig. 151). According to Nygren (1950:1998, abstract) : "the Bolivar geosyncline extends through coastal Ecuador and Colombia from southwestern Ecuador to the Golfo de Uraba. Six cross-basin highs separate the deeps within the trough. Marine sedimentation began in the south in the middle Eocene, gradually encroached northward, and continued intermittently on into the upper Miocene. Several unconformities of varying importance are present. Post- Miocene sedimentation is mostly non-marine. During the early Tertiary the sediments were largely derived from the west, but after the lower Miocene they were mostly from the east. Migration of terrestrial animals could have taken place through this area during the periods from upper Cretaceous to middle Eocene, middle Oligocene, lower Miocene, middle Miocene, and from upper Miocene to Re- cent [italics mine]." The western borderland of the Bolivar geosyncline is described by Nygren (p. 2005) as being a "rather wide lowland area up until upper Middle Miocene [when it] was sub- merged by down-faulting . . . except for a little narrow strip of land close to the present coast line. Remnants such as submerged peaks near Esmeraldas, Ecuador, Gorgona Island, and peaks at Cabo Corrientes still remain . . . The eastern borderland was an area of low relief with embayments extending to the Cauca Valley in upper Oligocene time." During middle Miocene, the "eastern borderland was raised into the high peaks of the western Cordillera [op. cit. p. 2003]." The north-south flowing Rios Atrato and San Juan now drain the Co- lombian or Chocoan portion of the main axis of the Bolivar Trough (fig. 152). The Atrato empties into the Golfo de Uraba in the Caribbean and the San Juan flows into the Bahia de Buenaventura in the Pacific. Except for the low, narrow divide between their heads, the Rios Atrato-San Juan now constitute the last ivater barrier separating Middle and South Ameri- can mammals. As the channels of the Atrato-San Juan became increasingly narrower through sedimentation more animals succeeded in crossing by ferrying, swimming or fording. As the meanders of the rivers became more numer- ous and sharper, more frequent cutoffs resulted and more habitats with their occupants passed from one side of the water gap to the other. Notwith- standing the accelerated rate of exchange, the mammalian fauna of the west 740 ECTOPARASITES OF PANAMA bank of the Atrato-San Juan is still predominantly Middle American and the entire coast from the Carribean Golfo de Uraba in Colombia to the Pacific Golfo de Guayas in Ecuador is zoogeographically an integral part of the Middle American Province (Hershkovitz, 1958, p. 596). The western or Middle American end of the Isthmian region was prob- ably undergoing comparable changes in geomorphology. There may have been one or several successive straits across the Isthmus of Panama (cf. Lloyd, 1963, p. 88; Fischer and Pessagno, 1965, p. 433). Tertiary mam- mals on any part of the shifting intercontinental region were, in effect, straddling the two continents. Sooner or later some became established on the mainland of the opposite continent while others retreated or disappeared entirely. By the time the continents were welded in fact, all mammalian interchanges as we know them today on the generic level, and many of them on the specific level, had already been consummated. LATE PLIOCENE-PLEISTOCENE FAUNAL INTERCHANGE Precise dating of complete closure of the Isthmian water gap would necessarily be arbitrary and as far as mammalian zoogeography is con- cerned hardly more than academic. Nevertheless, this date has been a major concern of students of the interrelationships of continental faunas and its determination by Simpson as the period of transition from upper Pliocene to Pleistocene governed the thinking of all zoogeographers. Simpson's (1940, p. 694) reasons are expressed in the following quotation: "In the Chapadmalalan [southeastern Buenos Aires Province, Argentina] for the first time many mammals of North American origin appear. They have inevitably undergone some change in the long journey but most or all of them could be derived from middle Pliocene forms in North America. These were the first animals surely to use the bridge. In North America animals of South American origin appear very sparingly in the middle Pliocene, then in more variety in the late Pliocene (Blancan stage). The middle Pliocene immigrants in North America are only a very few small ground sloths which could well have crossed (as did the procyonids in the reverse direction) before the bridge was quite complete and passable for the bulk of the fauna.12 It was thus complete and passable by Blancan in the North American sequence and by Chapadmalalan in the South American. The evidence is strong for the approximate equivalence of Blancan and Chapadmalalan. The Blancan is now considered the typical late Pliocene in North America." 13 [Footnote 13 follows:] "Some students insist that the earliest beds with any immigrant forms in the two continents should be synchronized, sometimes calling the Chapadmalalan middle Pliocene on this basis. Such a criterion, however, would make the Mesopotamian, not the Chapadmalalan, middle Pliocene, a correlation that is extremely improbable. It is more reasonable and more consistent with all other data to admit that the early stragglers crossed before the bridge was complete and to base synchronization on the beginning of extensive faunal interchange, a point already emphasized by Patterson (1937)." The point emphasized by Patterson (1937, p. 379) in his abstract of an unpublished paper is that: "it has generally been assumed that the Americas were reconnected about the beginning of the Pliocene. The vertebrates appear to oppose this view. Apart from raccoons and didelphines, no northern forms occur in the south until the end of the Pliocene, when a whole immigrant fauna suddenly appears. It is therefore held that the reconnection occurred in late middle Pliocene time at the earliest. Had it occurred earlier a more gradual diffusion southward of northern invaders HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 741 should be recorded. The didelphines and raccoons, and the megalonychine sloths found in the Middle Pliocene of North America are regarded as "waif" immi- grants [i. e. over water] ." Patterson offers no evidence, but his sources are evidently the same used in Simpson's revision of the mammal-bearing Tertiary of South America. In this work, Simpson (1940, pp. 678-680) names seven families of North American origin in the Chapadmalalan but does not enumerate the Blancan migrants from South America. Stirton (1936, p. 172) whose correlation of North American Pliocene faunas was the consensus for the time, lists an unidentified ground sloth and a glyptodon (Glyptotherium) in the Blancan. The only other South American mammals included by Stirton in upper Pliocene are the cricetine Sigmodon, a genus then regarded as of North American origin, and the dubious "Eligmodontla" (= Bensonomys) of supposed South American origin (see p. 737). Regarding the Blancan ground sloth, Simpson (supra cit., p. 694, footnote 12) suggests it was "an- cestral to or belonged to Megalonyx, a genus that arose in North America from primitive South American ancestors of Miocene or early Pliocene type." The Blancan glyptodont, so far as known, is also North American and therefore may have evolved from an earlier South American migrant. The Benson and Curtis Ranch Sigmodon remains as the only undoubted South American immigrant found in the North American late Pliocene. It is intimately related to the Chapadmalalan Reithrodon. The seven Chapadmalalan families treated by Simpson as North Amer- ican may have originated in North America or Eurasia. Only the history or place of origin of their South American representatives is relevant. These families and five Pampean (Pleistocene) families as listed by Simpson (1940, pp. 678-680), are enumerated as follows and then discussed sepa- rately. Cricetidae (mice) Procyonidae (raccoons, coatis, etc.) Ursidae (bears) Felidae (cats) Equidae (horses Tayassuidae (peccaries) Cervidae (deer) The following are listed as Pampean or Pleistocene. Canidae (dogs) Mustelidae (weasels, otters, etc.) Tapiridae (tapirs) Camelidae (llamas, vicugnas) Gomphotheriidae (mastodons) The only Cricetidae, more precisely, Cricetinae, known from the Chapadmalalan is Reithrodon, an autochthonous mouse of the sigmodont group (see above p. 736). Cricetines ancestral to South American forms diverged from North American stock, possibly in the Oligocene. The initial invasion of South America was probably over water, and judged by the diversity and complexity of subsequent radiations and the pervasiveness of the dispersal may have occurred during early Pliocene or Miocene. 742 ECTOPARASITES OF PANAMA According to Reig (1958, p. 252), the Ursidae and Felidae are not Chapadmalalan. So far as known, South American bears and cats date from the Pleistocene and were subsequently so listed by Simpson (1945, pp. Ill, 118). Two extinct genera of Procyonidae found in the Chapadmalalan are known since Mesopotamian or early Pliocene. This old American family of carnivores may have originated and radiated in Middle America. In any case a distinct species of the nominate genus Procyon evolved in South America and two highly specialized and monotypic genera, the prehensile tailed kinkajou (Potos) and the olingo (Bassaricyon) are certainly tropi- cal American if not specifically South American in origin. Fossil remains of these strictly arboreal animals are unknown. Simpson had no knowledge of the occurrence of mustelids in the Chapadmalalan but Reig (1958, p. 252) records a species of hog-nosed skunk, Conepatus.6 The Mustelidae are cosmopolitan and well represented in South America. The respective histories of such recent South American mustelids as the giant Amazonian otter (Pteronura) , the huron (Lyncodon) and the grisonella (Grisonella) as well as the skunk (Conepatus) must reach far back. Mustela frenata and Lutra canadensis (= annectens) are the only certain land bridge migrants from North America corresponding to cricetine stratum V. Whether the tayra (Eira barbara) and the grison (Galictis vittatus) spread from South to Middle America (stratum IV) or vice versa (stratum V), is moot. In any case, the dispersal of mustelids. particularly of otters, could not have been significantly controlled by minor water gaps. Absence of canids from the Chapadmalalan or earlier formations may be misleading. The South American radiation of the Canidae culminated in several genera and more living species than evolved on any other con- tinent. Such canids as Chrysocyon, Atelocynus and Speothos are most highly specialized for particular habitats in South America. The evolu- tionary history of these dogs is undoubtedly spread over a long span of time but the remains of their forest dwelling ancestors may be lost forever in middle Tertiary formations of tropical America. Among living canids, 6 There is some confusion attending the statigraphic position of Conepatus altiramus Reig. It was first (1952, p. 45) said to be from lower Chapadmalalan in the Barranca de los Lobos region. Later Reig (1957 , p. 39, footnote 3) reassigned it to the Barranca de los Lobos horizon which is lower Pleistocene. He then changed his mind again (1958, p. 252, footnote) and returned it to the Chapadmalalan on the basis of a personal in- spection of the actual site of discovery shown hkn by the collector of the type specimen. Now, Churcher and Van Zyll de Jong (1965, p. 3) compound the confusion by omitting the distinction between Barranca de los Lobos as a geographic region and as a geological horizon and, without consulting the pertinent literature (cited above, see also Kraglievich, 1952) imply that Conepatus altiramus is of lower Pleistocene age. Apart from all this, there are grounds for questioning the occurrence of Conepatus in the upper Pliocene of Argentina. The genus was abundant in the Pleistocene and is common throughout South and Middle America today. Scores of "species" have been described but all appear to belong to the same species complex for which the oldest name available is Conepatus chinga Molina, 1782. HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 743 only the gray fox (Urocyon) is an undoubted late or land bridge migrant from Middle America and almost certainly belongs to the cricetine stratum V. The dire wolf Aenocyon Merriam, first known from the upper Pleistocene of North America, also appears in the upper Pleistocene of Talara, north- western Peru (Lesson and Churcher, 1961, pp. 425-426) . Like many others of the same f aunal stratum it became extinct with the change from a moist to dry climate. Peccaries are retained by Reig (op. cit., p. 246) in upper Pliocene but the form known to Simpson (Antaodon chapalmalensis Ameghino) is ex- cluded. Horses and deer are also excluded. Later, Simpson (1945, pp. 137, 154) recorded the Equidae and Cervidae from the Pleistocene but not the Pliocene of South America. In any case, there is reason to believe that deer, at least, invaded South America during late Tertiary. The history of the Cervidae parallels that of the Cricetinae. The forest- dwelling spike-antlered pudus and brockets are probably old over-water invaders ; the more specialized woodland and savanna branch-antlered deer arrived later. The white-tailed deer, Odocoileus virginianus, with a con- tinuous range from southern Canada to the northern half of South America, may have spread from Middle to South America over the completed land bridge (stratum V) . One species of brocket (Mazama americana) returned over the same route to Middle America (stratum IV).7 Three species of tapirs occur in tropical America. The Andean Tapirus pinchaque and the lowland T. terrestris are confined to forested South America and certainly could have reached that continent by the over water route. Tapit*us bairdi with a continuous distribution in Middle America and northwestern South America is evidently a late overland arrival and corresponds to cricetine stratum V. In spite of its vagility and aquatic proclivities, Baird's tapir seems to have crossed the Rios Atrato-San Juan "gap" comparatively recently. So far as known, it has yet to reach the next important water gap, the Rio Cauca-Magdalena, only some 100 kilometers distant across the lowlands (Hershkovitz, 1954). From the foregoing it appears that only one genus of undoubted South American origin, the cricetine cotton rat Sigmodon, occurred in the upper Pliocene of North America. Of seven families of North American (and or Eurasian) origin treated by Simpson as Chapadmalalan in the upper Pli- ocene of South America, only three belong with certainty to that formation. Among their genera, the taxonomic units used here for correlations, only a peccary (Platygonus, Tayassuidae) may be of North American origin. All other undoubted Chapadmalalan genera originated in South America. 7 The Yucatan Peninsula brocket is a red brocket and should be known as Mazama americana pandora. Its generally brownish color (but not its color pattern), back- wardly directed nuchal hairs, and small size misled authors, including myself, into regarding pandora as a race of the brown brocket, Mazama gouazoubira. Pale brown or pandora-like red brockets are common enough as individual color variants or as local constants throughout the range of the species. With few exceptions, the white belly sharply denned from chest and flanks distinguishes Mazama americana from M. gouazoubira. 744 ECTOPARASITES OF PANAMA Cotton rats and peccaries are among the most viable and vagile of New World mammals. The presence of only one migrant genus (or species) in the upper Pliocene of each continent is not "a whole immigrant fauna" (Patterson, 1937, p. 379) or convincing evidence of an "extensive faunal interchange" (Simpson, 1940, p. 694, footnote 13) after closure of the Isthmian gap. Interchange there was, but since the Cretaceous, with the volume and variety directly proportional to the diversity of the faunas, and the population pressures at the intercontinental boundary whether or not a land bridge was present. The notion that first representatives of most families of northern North American origin crossed a land bridge into South America and radiated there in the Quaternary only, is not compatible with the facts. The tendency has been, however, to reject or ignore the facts incompatible with the notion. Patterson (in Stirton and Gidley, 1949, p. 175) thought it likely that a Megatherium from the Bone Valley Middle Pliocene of Florida was an acci- dental inclusion from overlaying Pleistocene because "the Americas were not united until late Pliocene." In the same work Stirton (loc. cit., footnote) points out that "both megalonychid and mylodontid ground sloths did get through from South America in Hemphillian, Middle Pliocene time." Stir- ton adds that "no megatheres are recorded in those faunas," but this qualifi- cation does not validate Patterson's objection. Thinking along the same line, Patterson (1957, p. 45) regarded the differentiation of a gyropid louse species during Pleistocene as remarkable and the evolution of an anopluran louse genus in the same period as "unlikely" because, "sucking lice [on their mammalian hosts] did not reach South America until the Pleistocene faunal interchange." Hooper (1949, p. 23) was perhaps the first to recognize the existence of two or more faunal strata among South American immigrants of north- ern North American origin. His observation that the "Oryzomys-like and Akodon-\ike forms" were of an older stratum than that containing Reithrodontomys (included Aporodon), anticipated his classification of complex-penis-type and simple-penis-type cricetines and my present ar- rangement of them in South American strata II and V, respectively. Nevertheless, in conforming to the Simpson-Patterson time table, Hooper telescoped the history of these strata into the Pliocene-Pleistocene transition. His conclusion that the differentiation of cricetines in South America was of "a low taxonomic level," somewhat follows Simpson's (1945, p. 207) miscalculated appraisal of South American cricetine evolution. Hooper's estimates have since undergone a radical change (cf. Hooper and Musser, 1964a, p. 55, and above, p. 734). My.previous (1962, p. 18) hypothesis of the time and sequence of the cricetine invasions of South America was also adjusted to the Simpson-Patterson chronology. It now appears unrealistic and must be discarded. CONCLUSIONS AND SUMMARY The evolution of the mammalian fauna of South America began in early Paleocene or late Cretaceous with migrants from Middle America. The first known immigrants were marsupials, condylarths and edentates. Primates HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 745 and caviomorph rodents appeared in the Middle Tertiary. Evidently, Ter- tiary Middle America was a transition zone for mammals passing from North America to South America. More importantly, it was an evolutionary center for mammals of North American extraction and a staging area for successive invasions of northern North America and South America by some of the newly evolved forms. This Middle American fauna must have been rich and flourishing but there are no fossil records of the tropical forest species. The meager middle and late Tertiary records of mammals are of essentially temperate zone pastoral faunas (cf. Olson and McGrew, 1941 ; Stirton, 1954; Whitmore and Stewart, 1965) which parallel the pres- ent temperate zone pastoral fauna of the Mexican and Guatemalan high- lands. These faunas no more deny the occurrence of a dominant lowland tropical forest fauna throughout Middle America now than in the past. Cor- respondingly, the presence of a diversified tropical forest fauna in Tertiary South America demands the existence of an earlier and more primitive tropical forest fauna in Middle America. Intercontinental faunal exchange at first was a flow from Middle Amer- ica to South America. With increase in their numbers and diversity some of the newly evolved South American mammals trickled into Middle Amer- ica by middle Tertiary. By late Tertiary the interchange in kind may have been more or less equal, but on a much larger scale than heretofore. Finally, from late Pliocene onward, the faunal flow from South America to Middle America became the greater, to the extent that the latter is a zoogeographic province of the former (Hershkovitz, 1958, p. 806). The five periods of intercontinental connections, the first in Cretaceous, the last in late Miocene, described by Nygren (1950), may have facilitated the faunal flow for species already on or at the bridge. The intercontinental separations during the intervening periods may have acted as faunal filters but they could never have been barriers comparable in effectiveness to climatic barriers. Virtually all South American mammals, past or present, crossed or could be passed over water gaps such as those which separated the continents during the Tertiary. The crossings must have been continu- ous with different individuals of the same species crossing at different times and different species of the same genera or families crossing at the same or different times. The volume and rate of mammalian faunal flow at the Isthmus must have always been directly proportional to the diversity of the faunas and their population pressures at the points of interchange. The last separation between North and South America is the Bolivar geosyncline. This depression formerly extended as a seaway from the Golfo de Uraba in the Caribbean to the Golfo de Guayaquil in the Pacific. Uplift of the Cordillera Occidental and increasing sedimentation reduced the water gap to the Rios Atrato and San Juan in western Colombia. The Bolivar Trough, nevertheless, still marks the boundary separating many mammalian species, genera, and one family (Geomyidae) of the Middle American Province from the rest of the Neotropical Region. The late Pliocene-early Pleistocene date generally accepted by zooge- ographers for completion of the intercontinental bridge and the beginning of wholesale faunal exchange is not in harmony with the fossil evidence. 746 ECTOPARASITES OF PANAMA Completion of the bridge at whatever date, did not mark the beginning of a rapid and large scale exchange of highly diversified mammalian faunas between Middle and South America. There is slight likelihood that the species and genera of cricetines, canids, procyonids, mustelids, deer, and others could have become peculiarly specialized for isolated niches in South America since the Pleistocene only. The probability is even less that these species and genera invaded South America preadapted for distant and un- tried habitats and predestined to reach them after trials through hostile environments. The zoological evidence suggests that representatives of all families of northern North American (or Eurasian) origin except those of horses, mastodons, camels, spiny mice, rabbits, shrews and, perhaps, some carnivores (bears, cats), were in South America before the end of the Pli- ocene. The invasions or, more realistically, the two way traffic, or continen- tal interchange, could have been operative since late Miocene. South American cricetine rodents are examples of the interchange. Their grade of evolution and the pervasiveness of their adaptive radiation point to long isolation from their North or Middle American ancestors. The history of tropical American cricetines appears in five faunal strata. Stratum I includes the ancestral North and Middle American complex penis stock. Over water migrants from stratum I evolved into the genera and generic groups or tribes of South American stratum II. Some divergent elements of stratum II returned via water routes to Middle and North Amer- ica and the West Indies. They became the genera, subgenera and species of stratum III. With completion of the land bridge many South American species of stratum II spread overland into Middle America to form stratum IV while two species of a purely North American tribe of simple penis type cricetines invaded South America and became stratum V. There has been little or no change in the taxonomic grade of strata IV and V. Preliminary analysis of the distribution patterns of a number of other tropical American families with intercontinental range reveals the same or parallel faunal stratifications. As in the case of cricetines, these strata indicate faunal interchange before, during and after completion of the inter- continental land bridge. NOMENCLATORIAL NOTE The Family Group Names of New World Cricetine Rodents The term Hesperomyini for New World cricetines is the tribal form proposed by Simpson (1945, p. 83) from the family group name Hesperomyinae Murray (1866, p. 358). Neither the family nor tribal name is tenable on nomenclatorial or zoological grounds. Murray's con- cept of Hesperomys, which is the type of his Hesperomyinae and Simpson's Hesperomyini, is based on twenty North American species, most of them referrable to Peromyscus and none identical or congeneric with the South American Hesperomys bimaculatus Waterhouse, type of the original Hesperomys Waterhouse. Hesperomys Murray, 1866, therefore, is an in- valid homonym of Hesperomys Waterhouse, 1839, and unavailable. Hesperomys Waterhouse, in turn, has been shown (Hershkovitz, 1962, p. HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 747 129) to be a junior synonym of Calomys Waterhouse, 1837. Finally, Calomys has been treated (op. cit.) as a member of the phyllotine group. The oldest valid tribal name for New World cricetines is Sigmodontes Wagner, 1843 (p. 509). As originally erected, it comprised the genera Hesperomys (= Calomys) , Reithrodon, Holochilus, Sigmodon and Neotoma. These taxons were employed in the broadest sense to include all New World species then known. Sigmodon J5ay and Ord, 1825, the type of Sigmodontes, is the oldest generic name for a New World cricetine. Its type, S. hispidus Gay and Ord, is cited and diagnosed by Wagner (1843, p. 556) . Sigmodontes was adopted by a number of authors including Baird (1857, pp. xxvii, 445) , Coues (1877, p. 7), Thomas (1897, p. 1019 [Sigmodontinae] ) , and Gyldenstolpe (1932, p. 1 [sigmodont] ) . Sigmodontinae as the subfamily name and Sigmodontini as a tribal name are available for New World cricetines and supercede Hesperomyinae and Hesperomyini, respectively. The supergeneric names based on Sigmodon which is a complex penis-type- cricetine, apply primarily to sigmodonts in particular and South American cricetines in general. Should North American simple-penis-type cricetines be regarded as tribally distinct, the name Peromyscini is available. More study of New and Old World cricetines, living and extinct, is needed, how- ever, before a relatively stable and generally acceptable classification can be reached. For present purposes, current supergeneric group names used informally are adequate and realistic. References ARATA, ANDREW A. 1964. The anatomy and taxonomic significance of the male accessory reproductive glands of muroid rodents. Bull. Fla. State Mus., 9, (1). 42 pp., 9 figs. BAIRD, SPENCER FULLER-TON 1857. Mammals of North America. The description of species based chiefly on the collection in the museum of the Smithsonian Institution. Part 1. General report on mammals of the several Pacific railroad routes. Washington D. 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Family Muridae, British Museum (Natural History), xii+690 pp., 50 figs. FISCHER, STANLEY P., AND PESSAGNO, JR., EMILE A. 1965. Upper Cretaceous strata of northwestern Panama. Bull. Amer. Assoc. Petrol. Geol., 49, (4), pp. 433-444, 4 figs. GAZIN, C. LEWIS 1942. The late Cenozoic vertebrate faunas from the San Pedro Valley, Ariz. Proc. U. S. Nat. Mus., 92:475-518, figs. 39-47, pis. 42, 43. 748 HERSHKOVITZ : LAND BRIDGES, FAUNAL INTERCHANGE 749 GUT, JAMES H., AND RAY, CLAYTON E. 1963. The Pleistocene vertebrate fauna of Reddick, Florida. Quart. Jour. Florida Acad. Sci., 26, (4), pp. 315-328. GYLDENSTOLPE, NILS 1932. A manuel of neotropical sigmodont rodents. Kungl. Svensk. Vet. Akad. HandL. 11, (3), pp. 1-164, 18 pis. HALL, E. RAYMOND 1955. Handbook of mammals of Kansas. Misc. Publ. Univ. Kans. Mus. Nat. Hist., no. 7. 303 pp., 99 figs., 1 cover pi. HALL, E. RAYMOND, AND KELSON, KEITH R. 1959. The mammals of North America. Ronald Press, N. Y. 1083+79 index pp., 553 skull figs, numbered, 500 map figs, numbered, many animal figs, unnumbered. HERNANDEZ CAMACHO, JORGE (in BORRERO, J. I. AND J. HERNANDEZ C.) 1957. Inf orme preliminar sobre aves y mamiferos de Santander. Colombia. Anal. Soc. Biol. Bogota, 7, (5), pp. 197-230. HERNANDEZ CAMACHO, JORGE, AND DE PORTA, JAIME 1960. Un nuevo bovido pleistocenico de Colombia: Colombibos atactodontus. Bol. Geol. Univ. Industr. Santander, no. 5, pp. 41-52, 1 pi. HERSHKOVITZ, PHILIP 1944. A systematic review of the neotropical water rats of the genus Nectomys (Cricetinae). Misc. Publ. Univ. Mich. Mus. Zool., no. 58. 88 pp., 4 pis., 5 figs., 2 maps. 1954. Mammals of northern Colombia, preliminary report no. 7: Tapirs (genus Tapirus), with a systematic review of American species. Proc. U. S. Nat. Mus., 103:465-496, figs. 58-62. 1955. South American marsh rats, genus Holochilus, with a summary of sigmodont rodents. Fieldiana: Zool., 37:639-673, figs. 139-144, pis. 17-29. 1958. 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Checklist of the Mammals of Panama CHARLES O. HANDLEY, JR.1 Since Goldman (1920) published his Mammals of Panama, and par- ticularly in recent years, much additional information has been gathered on the mammals of the Republic. Now there is need for an up-to-date check- list to supplement the still basic Goldman volume. Chief among recent contributors to the knowledge of Panamanian mam- mals have been Robert K. Enders and Oliver P. Pearson of Swarthmore College; Carl M. Johnson, Pedro Galindo, and Eustorgio Mendez of the Gorgas Memorial Laboratory, Panama ; several officers of the United States Army Preventive Medicine Survey Detachment in the Canal Zone : Colonel Franklin S. Blanton, Major Gordon Field, Major Robert Altman, Major Vernon J. Tipton, and Charles M. Keenan; Conrad E. Yunker, Middle America Research Unit; Nathan Gale, Canal Zone Veterinary Office; E. L. Tyson, Florida State University ; the former resident naturalist of the Canal Zone Biological Area, Carl B. Koford; and Alexander Wetmore, former Secretary of the Smithsonian Institution. Since 1952, seven field parties of the Smithsonian Institution have visited Panama to study mammals. These parties have enjoyed the support and cooperation of the Gorgas Memorial Laboratory, the United States Army, the Canal Zone Biological Area, the Middle America Research Unit, and the Chiriqui Land Company (through the Director of its Almirante Hospital, Dr. Gustav Engler). Part of the funds for field work and the preparation of this manuscript have been supplied by the National Science Foundation (grant no. G19415). The checklist follows the order of Simpson's classification of mammals (Simpson, 1945). As far as possible, the nomenclature adopted in the checklist reflects my study of specimens. Where subspecies are not men- tioned in a species account, it may be presumed that the species is thought to be monotypic. Since Goldman (1920) and Hall and Kelson (1959) are 1 Smithsonian Institution, Washington, D.C. 753 754 ECTOPARASITES OF PANAMA the references most likely to be available to persons using this checklist, departures from the nomenclature of these authors are indicated in syn- onymies. Other synonyms have not been mentioned, except in a few in- stances in the text. Full synonymies are readily accessible in Miller and Kellogg (1955), Hall and Kelson (1959), and Cabrera (1958 and 1961). Species that were not known to Goldman (1920) as part of the fauna of Panama are marked with an asterisk ( * ) . Those that were not known within the area covered by North American lists or had not been described when the most recent lists were prepared (Miller and Kellogg, 1955, and Hall and Kelson, 1959) are marked with a dagger (f). In instances where I was not the initial collector of a species here reported in Panama for the first time, I have noted this fact in the account of the species. Geographic distribution has been determined from specimens and from literature (citations omitted for brevity). Most of the specimens of Panamanian mammals are in the Academy of Natural Sciences of Phila- delphia, the American Museum of Natural History, the British Museum (Natural History), the Museum of Comparative Zoology (Harvard Uni- versity), and the United States National Museum, or in collections in Pan- ama. The brief notes on habits and habitat are based entirely on Pana- manian observations. Order MARSUPIALIA Family Didelphidae Caluromys derbianus Waterhouse = Philander laniger Goldman Common at lower elevations (sea level up to at least 3200 feet) , probably throughout Panama. Evergreen and deciduous forest ; arboreal. Several Panamanian subspecies: (1) C. d. derbianus Waterhouse. Darien west to Canal Zone and La Chorrera (Panama) ; similar specimens from Guanico (Los Santos), Santiago (Veraguas), and Boqueron and Isla Parida (Chiri- qui). (2) C. d. fervidus Thomas (=C. d. centrcdis Hollister). Caribbean coast of western Panama (mainland, Isla Bastimentos, and Cayo Agua, Bocas del Toro) and Santa Fe (Veraguas). (3) C. d. nautlcus Thomas. Islands off the Pacific coast of western Panama (Isla Brava, Chiriqui; Isla Gobernadora and Isla Cebaco, Veraguas) ; similar specimens from Parita (Herrera), and Panama Viejo (Panama). (4) C. d. pallidus Thomas. Pa- cific coast of extreme western Panama (Divala and Bugaba, Chiriqui) . Monodelphis adusta Thomas = Peramys melanops Goldman = Monodelphis melanops Hall and Kelson Rare at medium elevations in extreme eastern Panama. Specimens from Cana, 2000 feet, Guayabo, and Tacarcuna Village, 1950 feet (Darien) . Trapped among boulders on a dry, partially open gravel bar (Tacarcuna Village) and among rocks on a riverbank in heavy forest (Cana). Mono- delphis adusta occurs from eastern Panama through Colombia and Ecuador HANDLEY : CHECKLIST OF MAMMALS 755 to central Peru without exhibiting geographic variation. Peramys melanops Goldman and Peramys peruvianus Osgood are synonyms of adusta. Their supposed distinctive characteristics are merely individual variations. On the other hand, Monodelphis osgoodi Doutt of western Bolivia and southern Peru is probably a distinct species. It has been regarded as a subspecies of M. adusta. * f Marmosa impavida Tschudi Rare. At high elevations in extreme eastern Panama. 0. P. Pearson collected specimens on Loma Cana, 4900 feet, and Cerro Pirre, 5300 feet (Darien). The Panamanian subspecies is M. i. caucae Thomas. Marmosa invicta Goldman Rare. At medium elevations (1500-4000 feet), possibly throughout Panama, in suitable habitats. It has been trapped on the ground, and on or under logs in moist fog forest. Specimens from Cana, Tacarcuna Casita, and Tacarcuna Laguna (Darien) ; Cerro Azul (Panama) ; and Cylindro (Bocas del Toro). Marmosa mexicana Merriam = Marmosa mexicana Goldman (part.) Uncommon. In the lowland savannas of western Panama. The sub- species in Panama is M. m. savannarum Goldman. The relationship of M. mexicana to M. robinsoni is not clear. * f Marmosa species A species resembling Marmosa phaea Thomas has been taken at Tacar- cuna Casita, 1500 feet (Darien) ; Armila (San Bias) ; Salamanca Hydro- graphic Station (Canal Zone) ; and Almirante (Bocas del Toro). Ever- green forest; arboreal. Marmosa robinsoni Bangs = Marmosa fulviventer Goldman = Marmosa mexicana Goldman (part.) = Marmosa mitis Hall and Kelson Abundant throughout a large part of Panama from sea level to ele- vations of 4000-6000 feet. Semi-arboreal, but often trapped on the ground. Two subspecies have been reported in Panama : (1) M. r. fulviventer Bangs, Isla Saboga and Isla del Rey ; (2) M. r. isthmica Goldman, mainland. Prac- tically ubiquitous. For use of the name Marmosa robinsoni Bangs in place of M. mitis Bangs, see Cabrera (1958, p. 24). Philander opossum Linnaeus = Metachirus opossum Goldman Locally abundant at lower elevations (up to 2000 feet) , possibly through- out Panama. Mostly terrestrial, but also climbs. Evergreen and deciduous forest. The subspecies in Panama is P. o. fuscogriseus J. A. Allen. 756 ECTOPARASITES OF PANAMA Metachirus nudicaudatus E. Geoff roy St.-Hilaire Locally common at lower elevations (up to 2000 feet) , possibly through- out Panama. Terrestrial. Evergreen and deciduous forest. The subspecies in Panama is M. n. dentaneus Goldman. Where Metachirus nudicaudatus and Philander opossum occur together one is usually much more abundant than the other. Didelphis marsupialis Linnaeus Abundant throughout Panama from sea level up to 5000 feet. Ubiqui- tous. Terrestrial, but also climbs. Three subspecies are thought to occur in Panama: (1) D. m. battyi Thomas, Isla Coiba; (2) D. m. caucae J. A. Allen (=Z). m. etensis J. A. Allen) , mainland ; (3) D. m. particeps Goldman, Isla del Rey. Chironectes minimus Zimmermann = Chironectes panamensis Goldman = Chironectes panamensis Hall and Kelson Locally common to abundant throughout, from sea level to 4000 feet, in suitable streams. Aquatic, possibly restricted to streams with large populations of shrimp. Specimens from Boca de Rio Paya, Cana, 2000 feet, Guayabo, Rio Jaque, and Tacarcuna Village, 1950 feet (Darien) ; Armila and Mandinga (San Bias) ; Cerro Azul, 2000 feet (Panama) ; Fort Sherman, New San Juan, and Pedro Miguel (Canal Zone) ; Cerro Hoya, 1450 feet and Guanico (Los Santos) ; Rio Bubi and Rio Viejo, near Sona (Veraguas) ; Rio Colorado, 4000 feet (Chiriqui) ; and 7 km. SSW. of Changuinola and Sibube (Bocas del Toro) . Study of specimens from Bra- zil, British Guiana, Venezuela, and Central America shows that only one species can be distinguished. Variations in coloration and perhaps in cranial characters as well suggest that the species is polytypic. The Pana- manian subspecies is C. m. panamensis Goldman. Order INSECTIVORA Family Soricidae * Cryptotis endersi Setzer Rare. Known by one specimen from Cylindro (Bocas del Toro). Fog forest. Terrestrial. Cryptotis nigrescens J. A. Allen = Cryptotis merus Goldman = Cryptotis mera Hall and Kelson = Cryptotis tersus Hall and Kelson = Cryptotis zeteki Hall and Kelson Rare. At high elevations in evergreen forest. Terrestrial. Subspecies in Panama: (1) C. n. mera Goldman, Cerro Pirre, 4500 feet (Darien) ; (2) C. n. zeteki Setzer (= C. tersus Goodwin), Cerro Punta, 6500 feet, and Santa Clara, 4200 feet (Chiriqui). These subspecies are poorly differenti- ated from typical C. nigrescens of Costa Rica. HANDLEY : CHECKLIST OF MAMMALS 757 Order CHIROPTERA Family Emballonuridae Rhynchonycteris naso Wied-Neuwied = Rhynchiscus naso Goldman Abundant near sea level, probably throughout Panama. Flies over or near forest-bordered streams or lakes; roosts on rocks, logs, or trunks of trees over water. Specimens from Boca de Rio Paya, Paya Village, and Rio Jesucito (Darien) ; Armila and Mandinga (San Bias) ; Barro Colorado Island, Corozal, Fort Gulick, Gamboa, Lagartera, 2 mi. NW. of Pedro Miguel, Rio Mandinga, and Summit (Canal Zone) ; Almirante, 7 km. SSW. of Changuinola, and Sibube (Bocas del Toro) ; and San Felix (Chiriqui). For use of the generic name Rhynchonycteris Peters in place of Rhynchisciis Miller, see Husson (1962, p. 35). Saccopteryx bilineata Temminck Abundant near sea level, probably throughout Panama; occasionally found to 2000 feet. Flies over or near forest-bordered streams, lakes, or beaches; roosts on trunks of trees, on logs and rocks, and in caves and houses. Specimens from Cituro, Cana, and Cerro Sapo (Darien) ; Armila (San Bias) ; Alhajuela, Ancon, Balboa, Barro Colorado Island, Culebra, Fort Kobbe, Fort San Lorenzo, Fort Sherman, 3 mi. SE. of Gamboa, Gatun, Las Cascadas, Madden Dam, Orchid Island, Rio Puente, Summit, Tabernilla, Venado Beach (Canal Zone) ; Cerro Azul, Chilibre, Chilibrillo Caves, Isla Chapera, Isla San Jose, La Chorrera, Panama Vie jo, and Isla Taboga (Pa- nama) ; Guanico (Los Santos) ; and Almirante (Bocas del Toro). Saccopteryx leptura Schreber Uncommon. Near sea level in the Canal Zone and eastern Panama. Flies over or near forest-bordered streams and lakes. Specimens from Boca de Rio Paya and Paya Village (Darien) ; Pacora (Panama) ; and Alha- juela, Barro Colorado Island, Fort Randolph, and Gamboa (Canal Zone) . * Cormura brevirostris Wagner Uncommon. Lowlands of eastern Panama and on the Caribbean coast of western Panama. Evergreen forest. Roosts under logs and rocks. Spec- imens from Capeti (Darien) ; Armila (San Bias) ; and Almirante (Bocas del Toro) . *Peropteryx kappleri Peters Rare. Lowlands of eastern Panama and on the Caribbean coast of western Panama. Roosting in a cave at Rio Puente and netted over a stream flowing through a pasture at Almirante. Specimens from Yavisa (Darien) ; Rio Puente (Panama); and Almirante (Bocas del Toro). The Pana- manian subspecies is P. k. kappleri Peters. 758 ECTOPARASITES OF PANAMA Peropteryx macrotis Wagner = Peropteryx canina Goldman Uncommon. Possibly widespread in the lowlands, but presently re- corded only in and near the Canal Zone. Roosts in caves and rock crevices near water. Specimens from Buena Vista (Colon) ; Balboa, Gatun, and Santa Rosa (Canal Zone) ; and Rio Puente (Panama). The Panamanian subspecies is P. m. macrotis Wagner. Centronycteris maximiliani Fischer = Centronycteris centralis Goldman Rare. Lowlands of the Canal Zone and the Pacific coast of western Panama. Specimens from Cerro Mali (Darien) , Barro Colorado Island and Fort Clayton (Canal Zone) , and Bugaba (Chiriqui) . The Canal Zone speci- mens were taken in clearings. The Panamanian subspecies is C. m. cen- tralis Thomas. Diclidurus virgo Thomas Rare. Canal Zone and Pacific coast of western Panama at low and medium elevations. Specimens from Albrook Field, Fort Gulick, and Gam- boa (Canal Zone), and Boquete and Pueblo Nuevo (Chiriqui). Family Noctilionidae Noctilio labialis Kerr = Dirias albiventer Goldman Common at lower elevations, probably throughout Panama. Flies over streams and lakes ; roosts in hollow trees and houses. Specimens from Boca de Rio Paya (Darien) ; Armila (San Bias) ; Pacora (Panama) ; Nuevo Limon (Colon) ; Barro Colorado Island, Empire, Fort Gulick, Galeta Island, Gamboa, Juan Mina, and Puma Island (Canal Zone) ; Guanico (Los Santos) ; and Almirante, 7 km. SSW. of Changuinola, and Sibube (Bocas del Toro). The Panamanian subspecies is N. I. labialis Kerr (= Noctilio minor Osgood. See Cabrera, 1958, p. 55) . * Noctilio leporinus Linnaeus Common at lower elevations, probably throughout Panama. Flies over streams, lakes, and sea. Roosts in hollow trees. Specimens from Boca de Rio Paya (Darien) ; Armila (San Bias) ; Fort Sherman, Rio Chagres, and Rio Mandinga (Canal Zone) ; and Almirante and 7 km. SSW. of Changuinola (Bocas del Toro) . The Panamanian subspecies is 2V. I. mexicanus Goldman. Family Phyllostomidae Pteronotus parnellii Gray = Chilonycteris rubiginosa Goldman zz Chilonycteris parnellii Hall and Kelson Abundant throughout the lowlands of Panama and locally up to 4800 HANDLEY: CHECKLIST OF MAMMALS 759 feet. Evergreen and deciduous forests and fruit groves. Roosts in caves, tunnels, and probably in hollow trees. Specimens from Rio Chucunaque (Darien) ; Armila (San Bias) ; Cerro Azul, 2000 feet, and Chilibrillo Caves (Panama) ; Buena Vista (Colon) ; Bas Obispo, Galeta Island, Paraiso, 2 mi. N. of Summit and Vijia (Canal Zone) ; Cerro Hoya, 1500-2900 feet, and Guanico (Los Santos) ; Penonome (Code) ; and Almirante, 7 km. SSW. of Changuinola, head of Rio Changena, 4800 feet, and Sibube (Bocas del Toro). I am in full agreement with Burt and Stirton (1961, p. 24) on the synonymy of the generic names Pteronotus Gray and Chilonycteris Gray. At the species level, my own study of the problem shows that Koopman (1955, p. Ill) was right in his contention that P. parnellii Gray and P. rubiginosus Wagner are conspecific. According to the 1961 International Code of Zoological Nomenclature (Art. 21bii, p. 19) P. parnellii must be considered to be the prior name. The Panamanian subspecies is P. p. fuscus J. A. Allen. * Pteronotus psilotis Dobson = Chilonycteris psilotis Hall and Kelson In Panama, known only from the caves at Penonome (Cocle), where it is numerous, and at Armila (San Bias), where a few have been caught in mist nets set over forest streams. This species occurs without significant variation from Mexico to Trinidad. It may be conspecific with Chilonycteris personata Wagner, but specimens that have been positively identified as the latter species have not been available for comparison. The description and measurements of a specimen of C. personata from Mato Grosso (Bur- meister, 1854, p. 76, and Wagner, 1855, p. 680) agree with those of speci- mens of C. psilotis from Central America and northern South America. * Pteronotus suapurensis J. A. Allen Locally abundant in lowlands and up to 5300 feet, probably throughout Panama. Roosts in caves. Specimens from Tacarcuna Village, 1950 feet (Darien) ; Armila (San Bias) ; Chilibrillo Caves (Panama) ; Madden Dam (Canal Zone) ; Penonome (Cocle) ; and Cerro Punta, 5300 feet (Chiriqui). * Micronycteris brachyotis Dobson = Micronycteris platyceps Hall and Kelson Rare. Evergreen and deciduous forest. Roosts in caves and hollow trees. Specimens from Cerro Azul, 2000 feet (Panama) ; Fort San Lorenzo (Canal Zone) ; and Guanico (Los Santos). For the use of M. brachyotis in place of M. platyceps, see Goodwin and Greenhall (1961, p. 231). * Micronycteris hirsuta Peters Rare. Possibly throughout the lowlands of Panama and locally up to 4800 feet elevation. Evergreen and deciduous forests. Roosts in tree holes and buildings. Specimens from Cerro Azul, 2000 feet (Panama) ; Orchid Island (Canal Zone) ; Guanico (Los Santos) ; and upper Rio Changena, 4800 feet (Bocas del Toro). 760 ECTOPARASITES OF PANAMA Micronycteris megalotis Gray = Micronycteris microtis Goldman Locally common. Probably throughout the lowlands of Panama and up to 5300 feet elevation. Evergreen and deciduous forest. Roosts in houses and culverts and in hollow trees, logs, and stumps. Specimens from Pinogana and Rio Jesucito (Darien) ; Cerro Azul, 2000 feet, Isla San Jose, Pacora, Isla Taboga, and Isla Taboguilla (Panama) ; Barro Colorado Island, Cocoli, Fort Clayton, Fort Kobbe, Fort Randolph, and Orchid Island (Canal Zone) ; Guanico (Los Santos) ; Boquete and Cerro Punta, 5300 feet (Chiriqui) ; and Almirante and Punta de Pena (Bocas del Toro). The Panamanian subspecies is M. m. microtis Miller. * f Micronycteris minuta Gervais Rare. Taken in mist nets over streams in evergreen forest. Specimens from Boca de Rio Paya (Darien) and Cerro Hoya, 1800 feet (Los Santos). First taken in Panama by Rudolpho Hinds. * f Micronycteris nicef ori Sanborn Rare. Caribbean coast of eastern Panama. Taken in mist nets over stream in evergreen forest and from roost in concrete building. Specimens from Armila (San Bias) and Fort Gulick (Canal Zone). First taken in Panama by C. M. Keenan. * Micronycteris schmidtorum Sanborn Rare. Deciduous forest. Known in Panama only by specimens taken by F. M. Greenwell at Guanico (Los Santos). * Micronycteris sylvestris Thomas Rare. Taken in mist net over stream in evergreen forest at Armila (San Bias). Lonchorhina aurita Tomes = Lonchorina aurita Goldman Uncommon, but fairly widespread in Panama. Evergreen and de- ciduous forest. Roosts in caves and mine tunnels. Specimens from Chili- brillo Caves (Panama) ; 2 mi. N. of Summit and Vijia (Canal Zone) ; and 14 km. SSW. of Changuinola and upper Rio Changena, 4800 feet (Bocas del Toro) . The Panamanian subspecies is L. a. aurita Tomes. Macrophyllum macrophyllum Schinz Uncommon, but probably occurs throughout the lowlands of Panama. Flies over forest-bordered streams. Roosts in caves and masonry ruins (Panama Viejo) . Specimens from Jaque and Boca de Rio Paya (Darien) ; Armila and Puerto Obaldia (San Bias) ; Chepo, Pacora, Cerro Azul, 1200 feet, and Panama Viejo (Panama) ; Madden Dam, Fort Gulick, Fort Davis, and Salamanca Hydrographic Station (Canal Zone) ; Cerro Hoya, 1800 feet HANDLEY : CHECKLIST OF MAMMALS 761 (Los Santos) ; and Almirante and 12 km. SSW. of Changuinola (Bocas del Toro). * Tonatia bidens Spix Rare. Evergreen forest. Specimens from Cerro Azul, 2000 feet (Pana- ma) ; Madden Dam (Canal Zone) ; and Almirante (Bocas del Toro). * Tonatia minuta Goodwin = Tonatia nicaraguae Hall and Kelson Uncommon, but rather widespread in Panama, possibly throughout the lowlands. Evergreen and deciduous forests and fruit groves. Specimens from Armila, Mandinga, and Puerto Obaldia (San Bias) ; Cerro Azul, 2000 feet (Panama) ; Fort Sherman (Canal Zone) ; Guanico and Cerro Hoya, 1800 feet (Los Santos) ; 7 km. SSW. of Changuinola and Sibube (Bocas del Toro). Tonatia nicaraguae Goodwin is a synonym. Tonatia silvicola D'Orbigny = Tonatia amblyotis Goldman = Tonatia sylvicola Hall and Kelson Uncommon, but rather widespread in Panama, possibly throughout the lowlands. Evergreen and deciduous forest. Roosts in hollow termite nests. Specimens from Boca de Rio Paya, Rio Esnape, and Tacarcuna Village, 1950 feet (Darien) ; Armila and Puerto Obaldia (San Bias) ; Cerro Azul, 2000 feet (Panama) ; Barro Colorado Island, Corozal, and Rodman Naval Station (Canal Zone) ; Guanico (Los Santos) ; and Bugaba (Chiri- qui). The Panamanian subspecies is T. s silvicola D'Orbigny. For use of the name T. silvicola see Husson (1962, p. 87) . *Mimon cozumelae Goldman Rare. Evergreen forest. Known in Panama only by specimens found roosting in a small cave on the banks of the Rio Changuinola, 20 km. SSW. of Changuinola (Bocas del Toro) . *f Mimon crenulatum E. Geoffrey St.-Hilaire Rare. Found roosting in a hollow, rotting tree stump, and mist-netted over a forest stream and in a cacao grove. Evergreen forest. Specimens from Fort Gulick (Canal Zone) and Almirante and Sibube (Bocas del Toro) . The Panamanian subspecies is M. c. keenani Handley. * Phyllostomus discolor Wagner Common at lower elevations, probably throughout Panama. Evergreen forest and fruit groves. Specimens from Boca de Rio Paya, Paya Village, and Tacarcuna Village, 1950 feet (Darien) ; Armila, Mandinga, and Puerto Obaldia (San Bias) ; Cerro Azul, 2000 feet, and Chilibrillo Caves (Panama) ; Barro Colorado Island (Canal Zone) ; Guanico (Los Santos) ; Almirante, 7 and 14 km. SSW. of Changuinola, and Sibube (Bocas del Toro). The Pan- amanian subspecies is P. d. discolor Wagner. 762 ECTOPARASITES OF PANAMA Phyllostomus hastatus Pallas Common in lowlands, up to 2000 feet, throughout Panama. Forest and fruit groves. Roosts in hollow trees, caves, and houses. Specimens from Boca de Cupe, Boca de Rio Paya, Boca de Rio Punusa, Capeti, El Real, and Jaque (Darien) ; Armila and Puerto Obaldia (San Bias) ; Cabima, Cerro Azul, 2000 feet, Chepo, Chilibrillo Caves, Pacora, and Panama (Panama) ; Balboa, Barro Colorado Island, Fort Gulick, Juan Mina, Madden Dam, Paraiso, and Summit (Canal Zone) ; Guanico (Los Santos) ; Boqueron (Chiriqui) ; and Almirante, Boca del Drago, Cayo Agua, Isla Bastimentos, and Sibube (Bocas del Toro). The Panamanian subspecies is P. h. pana- mensis J. A. Allen. * Phylloderma stenops Peters •=. Phylloderma septentrionalis Hall & Kelson Rare. Evergreen forest. Specimens from Armila (San Bias). The Panamanian specimens combine the characters of the nominal species P. stenops Peters and P. septentrionalis Goodwin, but are nearer the former. Thus, the Panamanian subspecies is P. s. stenops Peters. Trachops cirrhosus Spix Common in lowlands and up to 2800 feet, probably throughout Panama. Flies over forest-bordered streams. Roosts in caves and houses. Specimens from Rio Jesucito and Tacarcuna Village, 1950 feet (Darien) ; Armila, Mandinga, and Puerto Obaldia (San Bias) ; Cerro Azul, 2000 feet, Chepo, Chilibrillo Caves, and Pacora (Panama) ; Fort Sherman (Canal Zone) ; Guanico and Cerro Hoya, 2800 feet (Los Santos) ; and 7-20 km. SSW. of Changuinola, Isla Bastimentos, and Sibube (Bocas del Toro). The Pan- amanian subspecies is T. c. cirrhosus Spix. * Chrotopterus auritus Peters Rare. Evergreen forest. Specimens from Tacarcuna Casita, 1500 feet (Darien), Armila (San Bias), and Cerro Azul, 2000 feet (Panama). I doubt that subspecies are recognizable in C. auritus. Vampyrum spectrum Linnaeus = Vampyrus spectrum Goldman Rare. Netted over forest-bordered streams and in fruit groves. Speci- mens from Tacarcuna Village, 1950 feet (Darien) ; Armila (San Bias) ; Boqueron (Chiriqui) ; and upper Rio Changena, 2400 feet, and Sibube (Bo- cas del Toro) . The subspecies V. s. nelsoni Goldman does not appear to be recognizable. I have compared the type with other specimens from Panama (7), Venezuela (4), Trinidad (10), Ecuador (1), and Peru (1). Although the Mexican and Central American specimens average smaller than those from South America, the difference is not great. The dental and cranial characteristics that Goldman supposed would distinguish nelsoni all prove to be individual variations. HANDLEY : CHECKLIST OF MAMMALS 763 * f Glossophaga commissarisi Gardner Common in the lowlands of eastern Panama, on the Caribbean coast of western Panama, and at medium elevations, possibly throughout Panama. Evergreen forest. Specimens from Boca de Cupe, Boca de Rio Paya, Paya Village, and Tacarcuna Village, 1950 feet (Darien) ; Armila and Mandinga (San Bias) ; Candelaria Hydrographic Station and Cerro Azul, 2000 feet (Panama) ; and Almirante, Boca del Drago, Cayo Agua, and Sibube (Bocas del Toro). Glossophaga soricina Pallas Abundant in the lowlands in central Panama and on the Pacific coast of western Panama and locally common on the Caribbean coast. Deciduous and evergreen forest. Roosts in caves, culverts, houses, hollow trees, and hollow logs. Specimens from Mandinga (San Bias) ; Colon and Portobelo (Colon) ; Isla Saboga, La Chorrera, Nueva Gorgona, Pacora, Panama, and Panama Viejo (Panama) ; many localities throughout the Canal Zone; El Cope, Rio Hato, and Santa Clara (Code) ; Guanico (Los Santos) ; Isla Canal de Afuera (Veraguas) ; and Almirante, Boca del Drago, 7 km. SSW. of Changuinola, Isla Bastimentos, Isla Colon, Isla Escudo de Veraguas, and Sibube (Bocas del Toro) . The Panamanian subspecies is G. s. leachii Gray. Lonchophylla mordax Thomas = Lonchophylla concava Goldman = Lonchophylla concava Hall and Kelson Rare. Evergreen forest. Roosts in caves. Specimens from Cana, 2000 feet, and Tacarcuna Village, 1950 feet (Darien) ; and Armila and Puerto Obaldia (San Bias) . The Panamanian subspecies is L. m. concava Goldman, which is only slightly differentiated from typical L. mordax. Lonchophylla robusta Miller Common. Eastern Panama and Caribbean lowlands of western Pan- ama. Evergreen forest. Roosts in caves. Specimens from Cana, 2000 feet, Tacarcuna Casita, 2600 feet, and Tacarcuna Village, 1950 feet (Darien) ; Armila and Puerto Obaldia (San Bias) ; Cerro Azul, 2000 feet, Chilibrillo Caves (Panama) ; Buena Vista (Colon) ; Barro Colorado Island, Salamanca Hydrographic Station and Summit (Canal Zone) ; and Almirante and Isla Bastimentos (Bocas del Toro). * f Lonchophylla thomasi J. A. Allen Rare. Known in Panama by a single specimen netted in a riverside clearing at Boca de Rio Paya (Darien) . * f Lionycteris spurrelli Thomas Rare. Evergreen forest. Specimens from Cana, 2000 feet (Darien) and Armila (San Bias) . First taken in Panama by O. P. Pearson. 764 ECTOPARASITES OF PANAMA * f Anoura cultrata Handley Uncommon. Evergreen forest. Specimens from Tacarcuna Village, 1950 feet (Darien) ; Cerro Punta, 5300 feet (Chiriqui) ; and upper Rio Changena, 2400 feet (Bocas del Toro). * Anoura geoffroyi Gray Uncommon. Evergreen forest. Specimens from Cerro Hoya, 3000 feet (Los Santos) ; Cerro Punta, 5300 feet (Chiriqui) ; and upper Rio Changena, 4800 feet (Bocas del Toro). The Panamanian subspecies is A. g. lasiopyga Peters. * Hylonycteris underwood! Thomas Rare. Known in Panama only by a specimen taken at Santa Clara, 4200 feet (Chiriqui) by F. A. Hartman. * Lichonycteris obscura Thomas Rare. Evergreen forest and fruit groves. Specimens from Tacarcuna Village, 3200 feet (Darien) ; Armila (San Bias) ; and Almirante and upper Rio Changena, 2400 feet (Bocas del Toro). Carollia castanea H. Allen = Hemiderma castaneum Goldman = Carollia castanea Hall and Kelson (part.) Common at lower elevations throughout Panama. Forest and fruit groves. Apparently most abundant in evergreen forest. Found roosting under overhanging stream bank and in mine. Specimens from Boca de Rio Paya, Cana, 2000 feet, Paya Village, Rio Seteganti, Tacarcuna Casita, 2600 feet, and Tacarcuna Village, 3200 feet (Darien) ; Armila, Mandinga, and Puerto Obaldia (San Bias) ; Cerro Azul, 2000 feet (Panama) ; Frijolito (Colon) ; Barro Colorado Island, Chiva Chiva, and Fort Kobbe (Canal Zone) ; Cerro Hoya, 3000 feet (Los Santos) ; and Almirante, Boca del Drago, 7 km. SSW. of Changuinola, and Sibube (Bocas del Toro). Carollia castanea is monotypic. It has been confused with Carollia subrufa Hahn, but it can be easily distinguished from that species by its smaller size (forearm usually less than 38 mm.) ; duller, less sharply de- fined banding of dorsal hairs ; occlusal outline of crown of inner lower in- cisor usually ovoid (wider than long) rather than triangular (longer than wide) ; first three lower post-canine teeth graded (increase in size from the first) rather than subequal ; Mt almost 'flat-crowned (all cusps obsolete) , sig- nificantly lower than adjacent teeth ; second upper premolar without postero- external cusp. Carollia castanea ranges from Costa Rica through Panama to northern and western Colombia. Carollia perspicillata Linnaeus = Hemiderma perspicillatum Goldman Abundant throughout the lowlands of Panama ; uncommon to rare above HANDLEY : CHECKLIST OF MAMMALS 765 3000 feet. Ubiquitous. The Panamanian subspecies is C. p. azteca Saus- sure. * Carollia subrufa Hahn = Carollia castanea Hall and Kelson (part.) Locally abundant throughout eastern Panama and at high elevations and on the Caribbean coast of western Panama. Usually the most abundant species of Carollia above 3000 feet. Evergreen forest. Roosts in caves and in hollow trees. Specimens from Paya Village, Rio Chucunaque, Rio Seteganti, Tacarcuna Casita, 1500 feet, and Tacarcuna Village, 1950 feet (Darien) ; Armila, Mandinga, and Puerto Obaldia (San Bias) ; Cerro Azul (Panama) ; Cerro Punta, 5300 feet (Chiriqui) ; and Almirante, Boca del Drago, Cayo Agua, 7 km. SSW. of Changuinola, Isla Bastimentos, Isla Colon, upper Rio Changena, 2400-4800 feet, and Sibube (Bocas del Toro) . Carollia subrufa has been confused with Carollia castanea, but actually it is most like and is more closely related to Carollia perspicillata. From C. perspicillata, C. subrufa differs in average smaller size (forearm 38-42 mm. vs. 41-45 mm.) ; average hairier forearm, tibia, and foot; average longer, silkier fur ; less crowded and posteriorly more divergent tooth rows ; reduced cusps of cheek teeth (especially protocone of M1) ; outer incisors not reduced ; highest point of P4 well ahead of the mid-length of the tooth. Carollia subrufa seems to be monotypic. It ranges from western Mexico to Colombia and Venezuela. Sturnira lilium E. Geoffrey St.-Hilaire Abundant at lower elevations (sea level to about 4000 feet) throughout Panama. Forests and fruit groves. Specimens from Boca de Cupe, Boca de Rio Paya, Paya Village, and Tacarcuna Village, 1950 feet (Darien) ; Armila, Mandinga, and Puerto Obaldia (San Bias) ; Cerro Azul, 1800-2000 feet (Panama) ; Quebrada Bonita (Colon) ; Cerro Hoya and Guanico (Los Santos) ; Volcan de Chiriqui (Chiriqui) ; and Almirante and Isla Bastimen- tos (Bocas del Toro). The Panamanian subspecies is S. L parvidens Gold- man. Specimens from Tacarcuna Village are larger than those from lower elevations in Darien and elsewhere in Panama and may represent an unde- scribed subspecies of