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University of Southern California Dissertations and Theses
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Systematics and host relationships of the mites of the family Spinturnicidae In Costa Rica (Acarina: Spinturnicidae)
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Systematics and host relationships of the mites of the family Spinturnicidae In Costa Rica (Acarina: Spinturnicidae)
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SYSTEMATICS AMD HOST KELATIOMSHIFS OF THE MITES OF THE FAMILY SPINTURNICIDAE IN COSTA RICA (ACARINA: SPINTURNICIDAE) by Richard Stratton Casebeer A Dissertation Presented to the FACULTY OF THE GRADUATE SCHOOL UNIVERSITY OF SOUTHERN CALIFORNIA In Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY (Biology) June 1966 U N IV E R SIT Y O F S O U T H E R N C A L IF O R N IA T H E G R A D U A T E S C H O O L U N IV E R S IT Y PA R K L O S A N G E L E S , C A L IF O R N IA 9 0 0 0 7 This dissertation, -written by ...... under the direction of h.%$...Dissertation Com mittee, and approved by all its members, has been presented to and accepted by the Graduate School, in partial fulfillment of requirements for the degree of D O C T O R OF P H I L O S O P H Y ..... D ate .June.1.966............ DISSERTATION COMMITTEE Chairman TABLE OF CONTENTS Page LIST OF MAPS....................................... v LIST OF TABLES..................................... vi LIST OF FIGURES......................................vii Chapter I. INTRODUCTION................................ 1 Acknowledgments Methods II. CHIROPTERA..................... 10 III. COLLECTION LOCALITIES...................... 14 Tropical Belt Subtropical Belt Lower Montane Belt Montane Belt Subalplne Belt IV. SYSTEMATICS............................... 24 Historical Review Systematic Arrangement of Species Keys to the Genera and Species of Splntumlcldae of Costa Rica V. SPINTURNICIDAE.............................. 41 Genus Perlallschrus Genus Cameronleta Genu8 Snlnturnlx Genus Parasplnturnlx ill iv Chapter Page VI. DISCUSSION............................. 105 Biology Life Cycle Biogeography Host Specificity VII. SUMMARY AND CONCLUSIONS...................... 126 LITERATURE CITED.......................................130 LIST OF MAPS Map Page 1. Costa Rican Forest Formations ......... 23 2. Collection localities of Periglischrus calisus..........• • • • ......... 50 3. Collection localities of Periglischrus ihertngi................................... 57 4. Collection localities of Periglischrus vargasi • • • • .........• • • • .......... 62 5. Collection localities of Periglischrus acutistemus............................... 66 6. Collection localities of Periglischrus olastii................................... 71 7• Collection localities of Periglischrus parvus. ................................... 74 8. Collection localities of Periglischrus torrealbai................................. 77 9. Collection localities of Periglischrus herrerai................................... 82 10. Collection localities of Cameronieta thomasi......................... 89 11. ' Collection localities of Cameronieta elongatus • • • • ......................... 94 12. Collection localities of Splntumix americanus................................. 99 13. Collection localities of Spinturaix subacuminatus •••••••••••••.•• 102 v LIST OF TABLES Table Page 1. Distribution of Genera of Spintumicidae on Families of Chiroptera................. 115 2. Distribution of Species of Periglischrus on Species of Chiroptera in Costa Rica, . . • 117 3. Distribution of Species of Cameronieta and Snintumix on Species of Chiroptera in Costa Rica.................................121 vi LIST OF FIGURES Figure Page 1. Typical Periglischrus (dorsal view) with structures labeled ..................... 33 2. Typical Periglischrus (ventral view) with structures labeled................ 34 vii CHAPTER I INTRODUCTION The spinturnicid mites are exclusively parasitic on bats (Chiroptera) in all stages of their life cycle. They seem to be well adapted for their precarious life on such active hosts. Their flattened bodies and thick, strong legs with large claws enable them to cling effectively to the wing and tail membranes of their hosts. The abbrevi ated life cycle in which the egg and larval stage are passed completely within the body of the female undoubtedly enhances their chances for survival. They are able to feed on the blood of their hosts in all independent stages of their life cycle. Because of this close association with their hosts, the spinturnicid mites are of interest for two reasons: (1) They are prime suspects as possible transmitters of disease although, as yet, they have not been incriminated as vectors of any diseases. Many species of bats are now 1 2 i known to be vectors, reservoirs, or both of a number of significant diseases affecting man and his domestic ani mals . Perhaps the most important of these diseases is rabies, which annually produces economic losses of many millions of dollars. Studies in Trinidad and Brazil (Pa- wan, 1936; Gilyard, 1945) demonstrated conclusively that rabies was transmitted to cattle, other domestic animals and to man by the bite cf the vampire bat, Desmodus rotun- dus. More recent studies have indicated that many species of insectivorous and frugivorous bats are also capable of carrying rabies and of transmitting it to other animals in cluding man by means of their bite (Deane, Maddy, Cockrum and Crecelius, 1960; Kent and Finegold, 1960; Condit and Humphrey, 1961). Constantine, (1962) offered evidence that bats may perhaps also transmit rabies by other means than their bite. He indicated that transmission by an airborne medium, such as an aerosol, was probable but did not rule out transmission by blood-sucking arthropods. The only re cord of the examination of bat mites for rabies was the re port by Irons (1955) of the inoculation of Bdellonvssus robustlpes (Ichoronvssus robustipes) into mice with nega tive results. There are no reports of the examination of spinturnicid mites for the rabies virus. The fungus, His tool asma capgtil atnm. has been re covered from bat guano and bat liver and spleen tissues in Panama (Schacklette, Diercks and Gale, 1962). The role of bats in the dissemination of this fungus in nature remains unknown. Likewise, the exact relationship between bats and encephalitis (Corristan, La Motte and Smith, 1956) and den gue (O'Conner, Rowan and Lawrence, 1955) is still largely unknown. Studies of this nature have stimulated greater in terest in bat ectoparasites, since knowledge of the system- atics and biology of these blood-sucking forms is essential to complete understanding of potential disease-vector rela tionships • (2) In addition to their potential epidemiological significance, the spinturnicid mites are also of interest because of the host specificity exhibited by many members of the family. The interpretation of this specificity is that the association between parasite and host goes far back into geological history so that the evolution of the parasite is closely correlated with that of the host. Thus, the phylogenetic relationships of the parasites may be indicative of the {foylogeny of the hosts. Studies of this nature, however, are dependent upon sound systematlcs land reliable host records. This study of the spinturnicid mites found on bats in Costa Rica has two specific goals. They are (1) to as certain what species of spinturnicid mites are found in Costa Rica and to elucidate their systematic relationships, thus making this information available for future epidemi ological studies, and (2) to analyze the associations of r* these mites with their hosts in respect to host specificity and phylogeny. Since certain aspects of host specificity are influenced by ecologic factors, some consideration of the environment of both the host and the parasite is given. The material utilized in this study was obtained in the course of a program designed to sample intensively a restricted area (Costa Rica) with the goal of collecting and examining for ectoparasites as many forms of mammalian hosts as possible, in as wide a variety of ecologic situa tions as possible. The author spent 14 months in Costa Rica, from July, 1962, through August, 1963, working with field teams during the summers of these years and alone during the intervening period. Other collecting teams were in the field from April through November, 1964. Thus the collectors were able to visit a large number of localities during various seasons. Although efforts were made to collect all forms of mammals and all ectoparasltlc groups, this paper deals only with bats and mites of the family Spinturnlcidae. Acknowledgments The author is grateful to Fred S. Truxal and Charles A. McLaughlin for allowing this work to be done while the author was employed as principal professional as sistant on their grant, "A Study of the Mammalian Ectopara sites and Their Hosts in Costa Rica," United States Army Medical Research and Development Command, Office of the Surgeon General, Grant Numbers DA-MD-49-193-62-G54 and DA- MD-49-193-63-G94. The author is also grateful to Andrew Starrett for funds from a Bache Fund Grant (National Acad emy of Sciences) which allowed a preliminary collecting trip to Costa Rica In 1961. * The advice and assistance of Andrew Starrett and Charles A. McLaughlin in making the host identifications is gratefully acknowledged, but the author accepts responsi bility for the accuracy of the host names reported here. 6 The discussions and advice of Charles L. Hogue and Andrew Starrett were both stimulating and helpful. The author also wishes to express his gratitude to John S. Garth, Chairman, and to the other members of the Doctoral Committee, Robert M. Chew, John L. Mohr, John W. Reith and Fred S. Truxal for critical review of the manuscript and helpful suggestions. Appreciation is expressed to Edward W. Baker of the United States National Museum and to Rupert L. Wenzel of the Chicago Natural History Museum for permission to ex amine specimens housed in their institutions. The author also wishes to thank his many friends in the Republic of Costa Rica for making the field work in that country not only possible but enjoyable. Methods The collecting of bats requires the use of many specialized techniques not normally utilized by mammalo- gists. One of the most successful methods involves the use of fine nylon "mist" nets. These nets are placed in clear ings or aeros8 natural flyways such as trails and streams. Such nets proved to be quite successful in ensnaring ! frugivorous, sanguivorous and pollen and nectar-feeding bats; however, the Insectivorous bats -with their more re fined sonar systems seemed able to detect and avoid the nets so that they were only occasionally taken In this manner. These forms were most often collected by shooting them with dust-shot as they hawked Insects near lights. A great many bats were collected by diligently searching out their daytime roosting places such as caves, culverts, attics and hollow trees and either shooting or hand col lecting them. A very limited number of bats was purchased from local collectors. As soon as they were collected, the bats were I placed in Individual cloth bags until examined. In some cases, where a large series of a single species was col lected, more than one individual was placed in a bag. The ectoparasites from such series were later pooled also. Be fore examination, the unopened bags and their contents were placed In a chloroform chamber to kill the bats and stupefy or kill the parasites. The hosts and the bags were then carefully examined under a dissection microscope and the ectoparasites collected. The bags were laundered after each use to prevent contamination with ectoparasites from a 8 i previous host. The ectoparasites from each host and in some cases from specific locations on the host (e.g., wing, nose leaf, ear or tragus) were placed in 70 per cent ethyl ! alcohol in individual vials which were labeled with the host18 field number and parasite location. In the laboratory the various groups of ectopara sites were segregated and prepared for further study. The spinturnicid mites were bleached in 10 per cent K0H in a warming oven, rinsed, neutralized in acid alcohol and mounted in Hoyer's medium. After drying in a warming oven (37 to 40 C) for several days the cover slips were ringed with King*s solution. The high-dry objective of a compound microscope was most frequently used for the study and identification of the spinturnicid mites. The use of phase-contrast objec tives was often necessary for the examination of delicately sclerotized structures. All information concerning the collecting locality, host and ectoparasites was recorded on "Unisort" punch cards in order to facilitate retrieval. Throughout the entire procedure of collecting, ex amining, sorting, mounting and labeling, extreme care was 9 taken to prevent contamination or confusion of ectopara sites from one host with another. In addition to the specimens collected during the grant project, a number of specimens collected by the au thor and Andrew Starrett in Costa Rica during the summer of 1961 was also utilized. For comparative purposes, a lim ited number of specimens from other countries was obtained by examination of dead bats preserved in alcohol in the collections of the Los Angeles County Museum. Only those mites still attached to their preserved hosts in their normal locations were considered to be associated with that host. Trips were made to the Chicago Natural History Mu seum and the United States National Museum in order to ex amine type specimens housed in these Institutions. CHAPTER II CHIROPTERA As the result of approximately twenty-two months of Intensive field collecting, 3,603 mammals were collected and examined for ectoparasites. Of these, 2,864 were bats representing eight families: Emballonuridae, Noctilionidae, Phyllostomatidae, Desmodontidae, Natalidae, Thyropteridae, Vespertilionidae and Molossldae. Although there are two records of spinturaicids from members of the family Noctil- lonidae (Furman, 1966; Machado-Allison, 1965b) and one re cord from a member of the family Emballonuridae (Turk, 1950), these two families and the Thyropteridae and Molos- sidae do not appear to be normal hosts for spinturnicid mites and the records probably represent errors of collec tion or identification or cases of "adventitious" or acci dental parasitism. The genera and species of bats in the remaining 10 four families which were collected in Costa Rica are pre sented in the following list (based on Hall and Kelson, 1959): i Family Phyllostomatidae Coues and Yarrow, 1875 Subfamily Chilonycterinae Flower and Lydekker, 1891 Chilonvcteris parnellii Rehn, 1904 Chilonvcteris psllotis Dobson, 1878 Pteronotus dawi Gray, 1838 Pteronotus suapurensis (J. A. Allen, 1904) Subfamily Phyllostomatinae Flower and Lydekker, 1891 Micronvcteris megalotis Miller, 1898 Micronvcteris schmidtorum Sanborn, 1935 Micronvcteris hirsute (Peters, 1869) Lonchorhina aurlta Tomes, 1863 Macrophvllum macrophvllum (Schinz, 1821) Phvllostomus hastatus (Pallas, 1767) Phvllostomus discolor (Wagner, 1843) Trachops cirrhosus (Splx, 1823) Chrotopterus auritus (Peters, 1856) Vampvrum spectrum (Linnaeus, 1758) Subfamily Glossophaglnae Gill, 1872 Glossophaga soricina (Pallas, 1766) Subfamily Glossophaginae Gill, 1872— Continued QtaWPhflgfl commlesarisl Gardner, 1962 Lonchophvlla concava Goldman, 1914 Lonchonhvlla robusta Miller, 1912 Anoura geoffrovl Gray, 1838 Anoura n. sp. ChoeronlBcus godmani (Thomas, 1903) Hvlonvcterls underwood! Thomas, 1903 Llchonvcterls obscura Thomas, 1895 Subfamily Carolliinae Miller, 1924 Carollia perspicillata (Linnaeus, 1758) CaroIlia castanea H. Allen, 1890 Carollia subrufa (Hahn, 1905) Subfamily Stumirinae Miller, 1907 Stumira lillum (Geoffrey, 1810) Sturnira ludovici Anthony, 1924 Stumira mordax (Goodwin, 1938) Subfamily Stenoderminae Gill, 1872 Uroderma bilobatum Peters, 1866 Vampvrops heller! Peters, 1866 Vampvrops vittatus (Peters, 1859) Vampvrodes major G. M. Allen, 1908 Subfamily Stenoderminae Gill, 1872— Continued Vampvressa Pusllla (Wagner, 1843) Chiroderma vlllosum Peters, 1860 Chiroderma salvlnl Dobson, 1878 Ectophvlla alba H. Allen, 1892 Artlbeus lamalcensls Leach, 1821 Artlbeus llturatua (Olfers, 1818) Artlbeus dnereus (Gervals, 1855) Artlbeus toltecua (Saussure, 1860) Centurlo senex Gray, 1842 Family Desmodontldae Gill, 1884 Desmodus rotundus (Geoffrey, 1810) Family Natalldae Miller, 1899 Natalus stramlneus Gray, 1838 Family Vespertllionidae Gray, 1821 Myotls nigricans (Schlnz, 1821) Myotls chlloensls (Waterhouse, 1838) Enteslcus fuscus (Beauvols, 1796) Epteslcus gaumerl (J. A. Allen, 1897) Epteslcus andinus J. A. Allen, 1914 Pa8ypterus ega (Gervals, 1855) Rhogeessa tumlda H. Allen, 1866 CHAPTER III COLLECTION LOCALITIES The Republic of Costa Rica occupies a territory of about 19,238 square miles. Within the boundaries of this relatively small region are found an extremely rich and varied flora and fauna. A number of factors account for this great biotic wealth. The area lies wholly within the tropics, a region which characteristically has a more com plex biota than that of temperate regions. Also, Costa Rica, as well as much of the rest of Central America, rep resents a meeting ground for many of the North and South American forms of plants and animals. Moreover, the varia tions in topography and wide range of altitudes, annual precipitation and edaphic factors found in Costa Rica pro duce pronounced differences in the natural vegetation with in relatively short distances. In most cases these dif ferences are also reflected in the composition of the 14 15 i [ fauna. The varied environments present In Costa Rica* com bined with the periods of Isolation from parts of North and South America during Its geologic history! have also re sulted In the evolution of many endemic forms (Lloyd, 1963; Maldonado-Koerdell, 1964,. In order to understand the relationships of the various vegetation associations, some system of classifica tion must be utilized as a guide for comparison and orien tation. A method of proven value in Costa Rica is the Holdridge System (Holdridge, 1947, 1953, 1964; Tosi, 1960, 1964; Slud, 1964) which has permitted mapping of plant-for mations or life-zones on the basis of climate. Five alti- tudlnal temperature belts are recognized in Costa Rica: the Tropical, the Subtropical, the Lower Montane, the Montane and the Subalpine. Each of these belts is subdivided into plant-formations on the basis of annual precipitation. Tropical Belt In Costa Rica the Tropical Belt extends, with local modifications, to an elevation of about 700 meters on the Pacific slope but only reaches 500 to 600 meters on the Caribbean slope due to the cooling effect of greater 16 cloudiness and heavier precipitation. Within this belt three plant-formations are recognized in Costa Rica. They are the Tropical Dry Forest, the Tropical Moist Forest and the Tropical Wet Forest. Tropical Drv Forest The Tropical Dry Forest formation is characteristic of much of Guanacaste Province in northwestern Costa Rica. It has a dry season of approximately five months. This formation originally supported a tall, deciduous forest but because of the valuable timber species and the ease of maintaining grasslands by the use of fire, man has modified considerably the original vegetation. Much of the forest has been burned and replaced by open savannas used for raising beef cattle. Tropical Moist Forest The Tropical Moist Forest formation is extensive in Costa Rica, covering much of the Caribbean lowlands and much of the southern part of the Pacific coastal plain. This formation is composed of tall, predominantly evergreen forest with three or four stories. Many of the trees have straight trunks free of branches for almost three-quarters of their total height. The main canopy Is quite dense and permits only a rather open understory of low, herbaceous undergrowth except along streams and the edges of clearings where more light Is admitted to the forest floor. There, impenetrable tangles of shrubby growth and vines may develop. Tropical Wet Forest The Tropical Wet Forest formation occurs in the northeast and In the Golfo Dulce lowlands in the southwest. This formation, like the preceding one, is composed of pre dominantly evergreen forest but is taller and more dense. There are many trees with buttressed trunks and stilt- rooted palms are common, especially in the more moist areas. Subtropical Belt The Subtropical Belt extends about 1000 meters above the Tropical Belt to an altitude of 1400 or 1500 meters. In Costa Rica two pi ant-format ions, the Subtrop ical Moist and the Subtropical Wet, are generally recog nized. Subtropical Moist Forest The Subtropical Moist Forest formation forms a nar row belt along the northern portion of the continental di vide. It occurs in the general region of the city of Car- tago and again in the south in the area around San Vito. Much of this formation is now utilized for coffee planta tions. The original forest is composed of relatively tall* evergreen trees in the few areas where it still exists. Subtropical Wet Forest The Subtropical Wet Forest formation covers large areas on both the Atlantic and Pacific slopes. This forma tion is composed of tall, evergreen forest with four stories and large, emergent trees. Buttressed trunks and stilt roots are relatively uncommon in this formation. In undisturbed areas the understory is quite open. Lower Montane Belt The Lower Montane Belt occupies an area of approxi mately 1000 meters of altitudlnal range above the Subtrop ical Belt. In Costa Rica it is found only on the Cordil lera Central and Cordillera Talamanca. Three plant-formations within this belt are found in Costa Rica. They are the Lower Montane Moist* Wet and Rain Forest for- i nations. Lower Montane Moist Forest The Lower Montane Moist Forest formation Is found only in a very restricted area near Irazu Volcano. The forest is composed of medium sized trees forming two stories. These trees are not particularly dense and allow sufficient light to the forest floor to support an abundant growth of tall shrubs and ferns. Epiphytes are present but not abundant. Lower Montane Wet Forest The Lower Montane Wet Forest formation is wide spread on the mountain slopes of central and southern Costa Rica. This formation is composed of tall trees forming an upper story and relatively small trees forming two lower stories. The understory Is not dense but at the ground level is found a dense carpet of ferns* mosses and lichens. Epiphytes are abundant* especially on the larger trees forming the upper story. Dense thickets form where 20 landslides and washouts expose the ground to sufficient light. Lower Montane Rain Forest The Lower Montane Rain Forest formation has been found only on the northeastern slopes of the Cordillera Central in Costa Rica. The forest is composed of smaller trees reaching a height of only 20 to 25 meters. The dense cloud cover which is present over this formation for most of the year supports a dense growth of epiphytes. Mosses, lichens and ferns cover the trunks and branches of the trees and bromeliads and orchids are abundant. The under story is dense and contains many tree ferns. Montane Belt The Montane Belt extends approximately 1000 meters above the Lower Montane Belt. Thus, in Costa Rica it is found only on the higher peaks of the Cordillera Central and the upper slopes of the Cordillera Talamanca. Within this belt only one plant-formation, the Montane Rain Forest, is clearly recognized in Costa Rica. Montane Rain Forest The Montane Rain Forest formation Is composed of a dense forest with a very dense understory of herbs and shrubs. Dense thickets of Chusquea sp., a spindly bamboo, fill almost every opening in the forest. Fog is frequent and in some places almost eternal. Accordingly, the forest is perpetually damp and supports an exceedingly lush epi phytic growth. Not only the vegetation but the ground as well is covered with dense growths of mosses, ferns and lichens. Subalpine Belt In Costa Rica only one formation, the Subalpine Wet Paramo, is found within this belt and this formation is found only in very restricted areas on the highest peaks of the Cordillera Talamanca. This formation is above the tim ber line and consists of thickets of spiny bamboo and scat tered low shrubs and herbs. The ground is densely covered with mosses and lichens. Floristically this formation is closely related to the paramos of the Andes. Many plants characteristic of the Andes find their northern limits here. 22 These plant-formations are delineated on Map 1. It must be noted that these formations are generally not sharply defined and that transitional zones usually exist between them. In addition, local variations In edaphlc or atmospheric conditions may produce many distinct associa tions within a given plant-formation. On the maps associated with each mite species, these plant formations are outlined and the localities where mites were collected are indicated. C O STA RICAN FOREST FORM ATIONS TROPICAL DRY TROPICAL MOIST TROPICAL WET SUBTROPICAL MOIST SUBTROPICAL WET LOW ER M O N TA N E MOIST LOWER M O NTANE WET LOWER M O NTANE RAIN M O NTANE RAIN SUBALPINE W ET PARAM O Mllil Map 1.— Costa Rican Forest Formations 23 ! CHAPTER IV SYSTEMATICS Historical Review The recent revision of the family Spintumlcidae by Rudnick (1960) included a detailed account of the history of the group from the time of Linnaeus. Considerable con fusion existed in the literature for some time as the re sult of the application of the specific name vespertilionis to both an insect (Nvcteribia) and a mite (Spinturnlx). Collins (1931) presented a detailed analysis of the nomen clature! problems involved. This problem was resolved by Opinion 128 of the International Commission on Zoological Nomenclature (1936) which invalidated the use of the spe cific name vespertilionis for the type of Spinturnix. From this confused beginning the history of the systematics of the group continued in a similar manner. Many of the early workers described or figured new species 24 25 very inadequately and did not designate type specimens so that it was difficult to identify them objectively. Thus, of the twenty-five species and seven genera described by Kolenati (1856 and 1857) in his two monographic works on the family, only six species and three genera were recog nizable by Rudnick (1960). The group was first named as a subfamily (Spintur- nlcinae) of the family Farasltidae by Oudemans in 1902. It is generally recognized as a family today (Vitzthum, 1940- 1943; Baker and Wharton, 1952), although a few recent workers have retained the subfamily Splntumicinae, placing it in the Parasitidae (Keegan, 1943; Hoffmann, 1944a, 1944b) or in the Laelaptldae (Evans, 1957; Evans, Sheals and Macfarlane, 1961). Rudnick (1960) recognized thirty-seven species dis tributed as indicated in the following seven genera: Ancvs- troous Kolenati, 1856, 5 species; Meristaspis Kolenati, 1857, 6 species; Evndhovenia Rudnick, 1960, 1 species; Paraperiglischrus Rudnick, 1960, 1 species; Perialischrus Kolenati, 1857, 4 species; Splnturnix von Heyden, 1826, 19 species; Paraspintumlx Rudnick, 1960, 1 species. He ap parently overlooked the paper by Till (1958) in which she described three additional species of Periglischrus. These three species undoubtedly belong in the genus Paraneriglia- chrus Rudnick, 1960, as evidenced by the extremely short dorsal peritremes, weakly sclerotized dorsal plates, deeply concave posterior coxal margins and host affinities. Benoit <1961) described a species of Periglischrus which also belongs in the genus Paraneriglischrus Rudnick, I960. Rudnick (1960) indicated that Ancvstronus kanheri Hiregaudar and Bal, 1955, was a rather aberrant member of the genus as Indicated by the large, recurved, hook-like processes on tarsus I and coxa I. Delfinado and Baker (1963), upon obtaining additional specimens of this spe cies, placed it in the new genus Oncoscelus. thus bringing the number of genera in the Splntumicidae to eight. They also described two new species of Ancvstronus. one new spe cies of Meristasnis and one new species of Snintumix. In 1964 Baker and Delfinado described still another new genus and species, Perigllschrodes gressittl Baker and Delfinado, 1964. In this same paper they also described five other new species in existing genera. In the New World, Machado-Allison (1964) described four new species of Periglischrus. In a succeeding paper 27 (Machado-All Ison, 1965a), he described the new genus and ispecles Cameronleta thomasl and in still another paper i (Machado-Allison, 1965b) he described four additional spe cies of Periglischrus. Furman (1966) In his excellent paper on the spin- turnlcids of Panama describes two additional new species of PerIgl 1 schrus. one new species of Snlnturnlx and synony- mizes one species of Snlnturnlx. He also redescrlbes Peri glischrus callgus Kolenati. o Analysis of these descriptions and comparisons of Furman's type specimens and Costa Rican specimens Indicates that some revisions are In order. Two of Machado-Allison's species of Periglischrus are synonyms of existing species. One of Furman's species of Periglischrus appears to belong In the genus as does one older species of Periglischrus. Thus the genus Periglischrus Kolenati, 1857, now contains ten species and the genus At-a Machado-AllIson, 1965, contains three species. The details of these synonymies and recombinations are presented In the discussions of the genera and Individual species. The systematic arrangement of the current genera and species of the family Splntumlcldae Is as follows: 28 I Systematic Arrangement of Species Family Spinturnicldae Oudemans, 1902^ Genus Ancvstronus Kolenati, 1856 Ancvstronus zeleboril Kolenati, 1856 Ancvstronus aethionicus Hirst, 1923 Ancvstronus indicus Hiregaudar and Bal, 1955 Ancvstronus tanrobanlus (Turk, 1950) Ancvstronus nalawanensls Delfinado and Baker, 1963 Ancvstronus eonvcteris Delfinado and Baker, 1963 Ancvstronus rudnicki Baker and Delfinado, 1964 Genus Oncoscelus Delfinado and Baker, 1963 Oncoscelus kanheri (Hiregaudar and Bal, 1956) Genus Heristasnis Kolenati, 1857 Meristasnis lateralis (Kolenati, 1856) Meristasnis calcaratus (Hirst, 1923) Meristasnis -tofdawl (Radford, 1947) Meristasnis kenvaensis (Radford, 1947) Meristaspls lavellanus (Radford, 1951) Meristasnis macroglossi (Hirst, 1923) Meristasnis mlndanaoensis Delfinado and Baker, 1963 ^Forms marked by an asterisk represent species re ported for the first time in Costa Rica. Genus Meristasnis Kolenati, 1857--Continued Meristaspls dusbabekl Baker and Delfinado, 1964 Genus Evndhovenia Rudnick, 1960 Evndhovenla eurvalls (Canestrlnl, 1884) Kvndhovenla ahl Baker and Delfinado, 1964 Genus Paraneriglischrus Rudnick, 1960 Paraperlsllschrus rhinoloohlnus (C. L. Koch, 1841) Paraperlsllschrus zuluensis (Till, 1958) Paraperlsllschrus moucheti (Till, 1958) Paraperlsllschrus nycterls (Till, 1958) Paraperlsllschrus trlaenopsls (Benoit, 1961) Paraperlsllschrus strandtmanni Baker and Delfinado, 1964 Paraperlsllschrus hlpposlderos Baker and Delfinado, 1964 Genus Perisllschrodes Baker and Delfinado, 1964 Perlsllschrodes sressittl Baker and Delfinado, 1964 Genus Perlsllschrua Kolenati, 1857 ♦Perlsllschrus callsus Kolenati, 1857 ♦Perlsllschrus lherlnsl Oudemans, 1902 ♦Perlsllschrus varsasl Hoffmann, 1944 ♦Perlsllschrus acutlsternus Machado-A111son, 1964 ♦Perlsllschrus olastil Machado-Allison, 1964 Genus Periglischrus Kolenati, 1857— Continued ♦Periglischrus parvus Machado-Allison, 1964 ♦Periglischrus torrealbai Machado-Allison, 1965 Periglischrus honkinsi Machado-Allison, 1965 ♦Periglischrus herrerai Machado-Allison, 1965 Periglischrus natali Furman, 1966 Genus fla— mn< efa Machado -Allison, 1965 ♦Cameronleta thomasi Machado-Allison, 1965 Cameronieta strandtmanni (Tibbetts, 1957) ♦Cameronleta elongatug (Furman, 1966) Genus Spinturnix von Heyden, 1826 Spintumix mvoti (Kolenati, 1856) Spinturnix mvstacinus (Kolenati, 1857) Spinturnix plecotinus (C. L. Koch, 1839) Spintumix multisetosus Rudnick, 1960 Spinturnix lawrencei Zumpt, 1951 Spintumix £si (Kolenati, 1856) Spintumix semilunaris De Meillon and Lavoipierre, 1944 Spintumix acuminatus (C. L. Koch, 1836) Spintumix abyssinicus Hirst, 1927 Spintumix scotophill Zumpt and Till, 1954 Genus Snlnturnlx von Heyden, 1826— Continued gpitttttniX valkerae Zumpt and Till, 1954 *§ginfofflais (Banks, 1902) gpintfflmix banks! Rudnick, 1960 gRtatarata kolenatll Oudemans, 1910 SRjUtiamta orlentalls Turk, 1950 Snlnturnlx bakerl Rudnick, 1960 Snlnturnlx orrl Rudnick, 1960 Spinturnix mexlcanus Rudnick, 1960 SJBta&mlK verutus Delfinado and Baker, 1963 Snlnturnlx naraeuminattia Baker and Delfinado, 1964 *5Pln&mvix subacumlnatus Furman, 1966 Genus Parasnlnturaix Rudnick, 1960 Parasnlntumlx globosus Rudnick, 1960 Keys to the Genera and Species of Snlntumlcldae of Costa Rica The following keys Include not only the forms col lected In Costa Rica and reported in this paper but also those forms expected to be found eventually in Costa Rica as indicated by their known distribution and host affini ties. Those forms marked by an asterisk represent forms not yet reported for Costa Rica. Important morphological structures are indicated in Figures 1 and 2. The terminol ogy follows that of Baker and Wharton (1952) and Rudnick ! 1(1960). Key to the Genera of Sninturnicidae of Costa Rica 1. Single dorsal shield; tritostemum present........ 2 Two dorsal shields, sometimes very delicately sclerotlzed; tritostemum absent.............. 3 2. Peritremes bend ventrad............ . Spintumix Peritremes completely dorsal ........ Parasplnturaix* 3. Unarmed ventral cutlcula of idlosoma with numerous minute, thorn-like mammilla- tlons; female sternal plate with trans verse striation8 ...................... Cameronleta Unarmed ventral cutlcula of idlosoma lack ing maomillations; female sternal plate longer or approximately as long as wide; male sternal plate without transverse 8trlatlons ............... Periglischrus Key to the Species of Periglischrus , of Costa Rica Females 1. Perltreme of normal size over coxa III, narrow and thread-like from coxa 111 to I; from Natalus mexicanus............natall Furman* Perltreme of normal size throughout, not constricted anteriorly ............ 2 Gnathosom a 'Dorsal Shiold Podosomo Fig. I.— 1 Typical Periglischrus (dorsal view) with structures labeled. T onus — Tibia — Patella — Femur---- Trochanter Palpus Coxa Epigynial Plate Claw Caruncle ' m t Fig. 2.— Typical Periglischrus (ventral view) with structures labeled. 34 I ; Females--Continued 35 2. 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 spp..............torrealbal Machado -A11 i son Ventral body setae simple, not with grossly expanded bases; first pair dorsal pro podosomal reduced or unreduced; second pair not reduced................................ 3 3. Femur, patella and tibia of legs III and IV each with an Inflated, straight, blade-like postero-ventral seta; leg IV lacking three aplcally recurved postero- ventral setae ........................ , Femur, patella and tibia of legs III and IV lacking Inflated, straight, blade- like postero-ventral seta; leg IV with three aplcally recurved postero-ventral setae ............................... 4. 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; usually from genera of Stenoderminae....................iherinai Oudemans First pair of dorsal propodosomal setae well developed, inserted on unarmed cuti- cula; ratio of distance between first pair of propodosomal setae to that be tween first and second pairs greater than 4:1; usual hosts Sturnira spp. . . . . . . ‘ ..................... last 11 Machado-Allison Females--Continued 36 5. Dorsal prodosomal setae relatively long, the longest measuring over 6Q*; tibia and tarsus of legs I and II lacking In flated, recurved postero-ventral setae.......... 6 Dorsal prodosomal setae relatively short, the longest measuring not over 5Q*,; tibia and tarsus of leg I and patella and tibia of II each with an Inflated, recurved, postero-ventral seta, superficially ap pearing blunt ........... 7 6. Femur II with only one of dorsal setae minute; ratio of distance between first pair of propodosomal setae to that be tween first and second pairs greater than 4:1; common on Desmodus . . .herrarai Machado-Allison Femur II with two of dorsal setae minute; ratio of distance between first pair of propodosomal setae to that between first and second pairs less than 3:1................. 8 7. Sternal plate broadly jug-shaped with short, narrow neck; anterior dorsal plate longer than broad (320* by 281^,); coxa III with relatively large posterior seta; common on Phvllostomus. Trachons .............. acutlsternus Machado-Allison Sternal plate narrowly pear-shaped with eroded margins and a long neck; anterior dorsal plate about as broad as long meas ured on the mid-line (243»); coxa III with very small posterior seta; common on Mlcronvcteris..............parvus Machado-Allison 8. Palpal tibia with strong medio-distal lobe; leg IV with large, broadly inflated, scimitar-like postero-ventral setae; dor sal opisthosoma bearing four small setae; from Glossophaga................. calieus Kolenati 37 Females— Continued Palpal tibia lacking medio-distal lobe; leg IV with elongate, setaceous, curved postero-ventral setae; dorsal oplsthosoma bearing six medium to large setae; hosts ■Anoura. Lentonvcterls ............ vargas1 Hoffmann Males 1. Perltreme constricted and thread-like anterior to mid-level of coxa III ...... natall Furman* Perltreme of normal size throughout.............. 2 2. Legs I and II with several blunt, fusiform setae; some ventral setae between coxae IV Inflated............ torrealbal Machado-Alllson Legs I and II without blunt, fusiform setae; ventral setae between coxae IV not Inflated . . . 3 3. Sternal plate setae short; anterior pair extending about one-half distance to base of second pair ......................... Sternal plate setae relatively long; anterior pair extending about four-fifths or more of distance to base of second pair .......... 4. Size small, Idlosoma less than 40Q**. long; posterior seta of coxa II about 5QA*long; dorsal propodosomal setae short, not over 4$«, l o n g ...........................callgus Kolenati Size large, Idlosoma over 50Q*~long; posterior seta of coxa II about 13Q«. long; dorsal propodosomal setae long, some over 6 l o n g vargaai Hoffmann 38 Males — Continued 5• 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 .............. acutlstemus Machado-Allison Tarsi III and IV superficially nude; ratio of length of posterior seta to that of anterior seta of coxa II more than 2 : 1 ........ 6 6• Anterior dorsal propodosomal setae rela tively close together; ratio of distance between bases of first pair of setae to that between bases of first and second pairs about 2:1................ iherinei Oudemans Anterior dorsal propodosomal setae rela tively far apart; ratio of distance be tween bases of first pair of setae to that between bases of first and second pairs about 4:1 or more........................ 7 7. Three pairs setae on unarmed dorsal oplstho- soma; ventral pair setae behind sternal plate about three-fourths length of pos terior setae of sternal plate ...... herrerai Machado-Allison One pair setae on unarmed dorsal opisthosoma; ventral pair setae behind sternal plate about one-fourth length of posterior setae of sternal plate .............................. 8 8. Relatively large mite with anterior legs, exclusive of ambulacra* over 50Qt*> long; spermatophoral process less than lOQ*^ long* shaped as a shepherd's crook . . . .......................... olastil Machado-Aliison 39 Males--Contlnued Relatively small mite with anterior legs, exclusive of ambulacra, about 400^ long; spermatophoral process over 15Q^ long and extensively recurved . . parvus Machado-AllIson Key to the Species of Cameronleta of Costa Rica Females 1. Medial margins of first pair of coxae heavily sclerotlzed and articulating with sternal plate; camerostome sur rounding the gnat ho soma present . . . ........................... thomasl Machado-AllIson Medial margins of first pair of coxae not heavily sclerotlzed and not articu lating with sternal plate; camerostome surrounding the gnathosoma absent.............. 2 2. Anterior margin of sternal plate with broad, anteromedian projection; opisthosoma with four pairs prominently plumose ventral setae in addition to normal setae; all legs with broad, flat, prominently fringed ventral setae . . . ..................................elongatus (Furman) Anterior margin of sternal plate slightly concave; opisthosoma lacking prominently plumose ventral setae; all legs with ventral setae of normal shape and at most narrowly fringed ........ strandtmannl (Tibbetts)* Males 1. Coxa II with posterior seta much larger than anterior seta; distal seta of coxa I expanded, blade-like .............. strandtmanni (Tibbetts)* Males— Continued 40 Coxa II with two long, subequal setae; distal seta of coxa I simple, setiform........ 2 2. Coxa I with two subequal, setiform setae .......... elonaatus (Furman) Coxa I with distal setae approximately twice as long as proximal setae . . . ......... thomasl Machado-Allison Kev to the Species of Snlnturnlx of Costa Rica Females and Males 1. Lateroventral setae of legs mostly short; pair of proximal dorsal setae of femora I and II minute; proximal postero-dorsal seta of femora III and IV minute . amerlcanus (Banks) Lateroventral setae of legs mostly long; pair of proximal dorsal setae of femora I and II long; proximal postero-dorsal seta of femora III and IV long . atibactml natus Furman CHAPTER V S PINTURNICIDAE Genus Periglischrus Kolenati, 1857 Periglischrus Kolenati, 1857, Wien. Ent. Monatschr., 1(2):60; Rudnick, 1960, Univ. Calif. Publ. Ent., 17(2):195 (complete bibliographical synonymy to 1960); Machado-Alli- son, 1964, Rev. Soc. Mexicana Hist. Nat., 25:193; Machado- Allison, 1965, Acta Biol. Venez., 4(10):243; Machado-Alli- son, 1965, Acta Biol. Venez., 4(11):259; Furman, 1966, In Ectoparasites of Panama, In press. Tvne:species Periglischrus caligus Kolenati, 1857, by subsequent designation (Oudemans, 1903, Tljdschr. Ent., 45:135). Diagnosis Two dorsal shields. Tritostemum absent. Peri- treme completely dorsal, usually extending from level of 41 42 I coxa IV to level of coxa I. Abdomen of non-teneral females usually greatly expanded, fan-shaped. Dorsum:— Two dorsal shields closely approximated, occupying greater part of podosoma in female and most of idlosoma in male. Five pairs of setae bordering anterior dorsal shield anterior to stigmata. One pair of setae slightly posteromedial to stigmata. Four or six pairs of dorsal opisthosomal setae in female. Venter:— Lacking tritosternum. Female sternal plate as long as or longer than wide with three pairs of marginal setae, which may be set off plate. Male sternal plate with five pairs setae. Epigynial plate reduced to narrow solerotization with pair of small genital setae close to or on lateral margins of plate near posterior tip. Opisthosoma of female greatly expanded to broad, flat, fan shaped sppearance with characteristically shaped areas of heavy sclerotization. Anal plate of female small, subter minal with pair adanal setae and dorsal postanal seta. Op isthosoma of male reduced, projecting only slightly poste rior to level of coxa IV. Anal plate of male large, oc cupying much of area between coxa IV, with several pairs of setae and minute dorsoterminal postanal seta. 43 Legs:— Large caruncles and claws on all legs. Dor sal and lateral setae short to long. Ventral setae usually i short. Tarsus 1 of males with two long, bluntly tipped idorsal sensory setae located respectively at basal one- third and apical positions. I The genus Periglischrus Kolenati, 1857, contained a number of varied forms from both the New and Old World un til the revision of Rudnick (1960) established the genus Paraoeriglischrus to include the Old World forms character ized by reduced sclerotization of the dorsal shields and very short, completely dorsal peritremes. Paraperipj lia- chrus has been reported only from bats of the family Rhi no lophidae. Rudnick (1960) restricted the genus Periglischrus to New World forms from bats of the family Phyllostomatidae. He included the type-species, Periglischrus calieus Kole nati, 1857, although the original description and figures were Inadequate for a species diagnosis and the type spec imens could not be located. Furman (1966) redescribes P. caligus from a specimen Identified as this species by Kole nati. Rudnick (1960) recognized three additional species, P* iherin^i Oudemans, 1902, P. vargasi Hoffmann, 1944, and 44 P. atrandtmannl Tibbetts, 1957. P. strandtmamU is as signed to the genus Cameronleta. Since Rudnlck's revision seven valid new species have been described, bringing the total number to ten. Considerable confusion exists at the present time in the nomenclature of many of the species of Perialischrus Much of this confusion arises from the almost simultaneous publications of descriptions of numerous new species from the Neotroplcs by two different authors (Machado-Allison, 1964 and 1965b; Furman, 1966). Unfortunately, each was un aware of the other's work and was thus describing many of the same species. According to the Law of Priority, many of these species must fall into synonymy. Thus, five of the seven new species of Perialischrus described by Furman become synonyms of species described by Machado-Allison, whose papers were published first. This condition is further confused by the fact that two of Machado-Allison's species appear to be synonyms of previously described species, although in one case he was not aware of the forthcoming redescription of P. call gum by Furman. The present author has had the opportunity of ex amining all of Furman's type specimens and his manuscript 45 j on the spinturnidd mites of Panama and of comparing them ! with specimens from Costa Rica. Analysis of these speci mens and the descriptions by Machado-Allison makes possible the synonymies listed under the discussions of the Individ ual species. Periglischrus calieus Kolenati Periglischrus caligus Kolenati, 1857, Wien. Ent. Monatschr., 1(2):60; Rudnlck, 1960, Univ. Calif. Publ. Ent. 17(2):196; Machado-Allison, 1965, Acta Biol. Venez., 4(11): 259; Furman, 1966, In Ectoparasites of Panama, In press. Periglischrus setosus Machado-Aliison, 1964, Rev. Soc. Mexicans Hist. Nat., 25:193; Machado-Allison, 1965, Acta Biol. Venez., 4(11):271. (New synonymy). This species was described by Kolenati (1857) from Glossonhaga soricina from Brazil and Surinam. Although the original description and figures did not permit species diagnosis, Rudnlck (1960) retained it as a valid species in the hope that the type or new specimens from the type host might be found. Furman (1966) obtained a female specimen from the Museum National d'Histoire Naturelle, Paris, iden tified as this species by Kolenati. Upon comparison this i specimen proved to be identical with numerous specimens from Glossophase sorlcina from Panama and Trinidad. Fur- man, therefore, redescribes and Illustrates the female and describes the male for the first time from these specimens. He deposited the plesiotype female and male In the United States National Museum. The present author examined these specimens and was thus able to identify positively the specimens from Costa Rica. Machado-Allison (1964) described the female of Periglischrus setosus collected from Glossoohaga. In a succeeding paper (Machado-Allison, 1965b), he described the male of this species and indicated that, on the basis of host affinities, P. setosus may be a synonym of P. caligus but is retained as a valid species until the type of P. caligus can be compared. He was, of course, unaware of the forthcoming description of P. caligus by Furman (1966). Comparison of Costa Rican and Brazilian specimens of P. caligus in the collections of the Los Angeles County Museum with the descriptions of P. setosus (Machado-Allisoxv 1964) clearly indicates that the latter species is a syno nym of P. caligus. Females of this species may be recognised by the relatively long dorsal propodosomal setae, large medio-dis tal lobe of palpal tibia, characteristically shaped sternal plate and pattern of dorsal shield. The males are distinc tive in having very short setae on the sternal plate and by their small size. Recorded Hosts and Distribution Brazil and Surinam:— This species has been re ported from GlosBpphaga soricina in Brazil and Surinam (Kolenati, 1857). Venezuela:— Specimens are recorded from Glos- sophaga soricina. G. longirrostris [sic], Anoura caudifera and A. cultrata in Venezuela (Machado- Allison, 1964, 1965b). Machado-Allison (1965b) also indicated in his host distribution chart (cuadro No. 11) one specimen taken from Noctilio lablalis but makes no mention of this in his col lection records. Since this chart contains nu merous errors, it is not inconceivable that this record might also be in error. Panama: — Furman (1966) reports this species from Gloaaophapa soricina. Trinidad: — The records from Trinidad are also from Glossophaga soricina (Furman, 1966). Costa Rican Hosts and Distribution Periglischrus caligus is recorded for the first time in Costa Rica from the following hosts and localities: 48 Glossophaga soricina: — Rfo Vlrllla Caves, Here dia Prov., July 7, 1962 and June 12, 1963; Playa del Coco, Guanacaste Prov., July 20, 1962; Boca de Barranca, Puntarenas Prov., July 31, 1962 and July 28# 1964; San Jos£, San Jos£ Prov., Oct. 16, 1962; Finca La Lola, Lim&n Prov., Sept. 26, 1962; Rio Oro, one mile west of Santa Ana, San Jos£ Prov., Oct. 8, 1962; Caves, two miles west of Turrlalba, Cartago Prov., Dec. 4, 1962; Turrlalba, Cartago Prov., Dec. 4, 1962; 57.5 miles south of San Isidro on Pan American Highway, Puntarenas Prov., Aug. 22, 1963; San Isidro, San Jose Prov., June 23, 1964; Rio Con- vento, Puntarenas Prov., June 24, 1964; Estrellas Oeste, Puntarenas Prov., November 26, 1964. Glossoohaga conanissarlsl: — Rincdn. Osa Penin sula, Puntarenas Prov., June 29, 1963. Glossophaga sp. (probably ltmglroatrial:— Plava del Coco, Guanacaste Prov., July 19, 21 and 22, 1962; San Josl, San Jos6 Prov., Sept. 10, 1962. Discussion This species of mite appears to be restricted to bat hosts of the genus Glossophaga in Costa Rica. Seven other species of bats in other genera of the subfamily Glossophaginae, as listed in the host list, lacked this mite. Only a small number of bats was infested. Even in cases where large samples of hosts were taken from a colo ny, the occurrence of infested individuals was only spo radic. Juvenile bats usually carried larger numbers of mites than did the adults. As many as fourteen mites were taken from one juvenile bat. 49 In addition to the usual location# on the wing and i tail membranes of the hosts# a number of mites mas found in the ears attached to either the pinna or the tragus. Glos- sophfga bats also have a shallow fossa immediately below i Ithe eye that frequently contained mites. These mites were identical morphologically with those found on the wings or ears. Rudnlck (1960) states that "all (mite) species with t the exception of those in the genera Ancvstronus. Mar is- tasois and Parasointurnlx are found exclusively on the wing and tail membranes•*' This is not the case with many spe cies of Periglischrus. Periglischrus caligus was collected from all forma tions occurring in the Tropical and Subtropical Belts in Costa Rica (Map 2). Periglischrus iheringi Oudemans Periglischrus iheringi Oudemans# 1902# Ent. Ber.# Amsterdam# 1(6):38; Rudnlck# 1960# Univ. Calif. Publ. Ent. 17(2): 197; Machado-Allison# 1964# Rev. Soc. Mexicana Hist. Nat.# 25:194; Machado-Allison# 1965# Acta Biol. Venes.# 4 (11): 264; Furman# 1966# In Ectoparasites of Panama# In press. 50 This species was originally described by Ondemans !(1902) and later redescribed in more detail and illustrated i by Rudnick (1960), after examination of the type specimens. Characters distinguishing the females include the ; minute anterior pair of dorsal propodosomal setae inserted relatively close together usually on the anterior margin of the dorsal shield, the straight blade-like setae on the postero-ventral margins of legs III and IV, and the blade like posterior setae of coxa III. The males of J?. iheringi have long, dorsal propodosomal setae, the anterior pair of which are situated relatively close together. Males pos sess only one small antero-basal seta on the dorsal surface of femur II. Recorded Hosts and Distribution This widespread species of mite has been recorded from a number of genera and species of bats of the subfam ily Stenoderminae (Fhyllostomatidae) from many parts of the New World. Brasil: — The type host, Vjmpymps lineatus. is reported as a host in Brasil (Oudemans, 1902). Rudnlck (1960) also reported specimens from Artib- eus lituratus in Brasil. Paraguay:— Specimens are reported from Vampyr- ops sp. (Rudnick, I960). 52 Colombia; --Records are from A. lituratus In Colombia (Rudnick, 1960). Venezuela:--Rudnick (1960) reported specimens from Artibeus lamaicensis. In addition to these records Machado-Allison (1965b) reported specimens from A. lituratus. A. lamaicensis. A. cinereus. A. concolor. Vampvrops dorsalis. V. vittatus. Chiro- derma sp., Enchisthenes hartii [sic] and Uroderma bilobatum. He also lists one specimen from Stur- nira ludovici but does not include this in his host distribution chart. Trinidad:— Rudnick (1960) reported specimens from A. lituratus. Cuba. Puerto Rico and Virgin Islands; — Records are from A. lamaicensis (Rudnick, 1960). Panama:— Furman (1966) reports the following bats as hosts in Panama: A. literatus. A. lamai censis. A. to It ecus. A. cinereus. A. ax¥ecus. Uroderma blloh*tiim. V-ampyr-npg vittatus. V. hellerl. Vampvrodes malor. Vampvressa pusllla. Chlroderma salvlni. Enchis thenes harti and Desmodus rotundus. Honduras: — Specimens recorded are from A. lituratus (Rudnick, 1960)• Guatemala: — Records are from A. lituratus and Uroderma bilobatum (Rudnick, 1960) • Mexico:— Rudnick (1960) reported specimens from lamaicensis and A. cinereus in Mexico. Only two records are from bats other than members of the subfamily Stenoderminae. Machado-Aliison (1965b) listed a specimen from Stumlra ludovici. a member of the subfamily Sturairinae, and Furman (1966) lists specimens 53 from Dewodus rotundus. a member o£ another family, the Desmodontidae• Costa Rican Hosts and Distribution Periglischrus iheringi is reported for the first time in Costa Rica from nine species of bats representing five genera in the subfamily Stenoderminae. The records are as follow: Uroderma bijyobatum:--Playa del Coco, Guanacaste Prov., July 19 and 25, 1962; two miles south of Tilardn, Guanacaste Prov., July 28, 1962; Boca de Barranca, Puntarenas Prov., August 1 and 2, 1962; Flnca La Lola, Limdn Prov., September 30, 1962, July 23 and 24, 1963; Dominical, Puntarenas Prov., November 22 and 23, 1962. Vampvrons hellerl:— Gromaco Foundation, Punt* arenas Prov., August 24 and 25, 1962; Hacienda Moravia, Moravia de Chirripd, Cartago Prov., Octo ber 21, 22 and 24, 1962; San Isidro de General, San Jos6 Prov., November 20, 1962; Turrlalba, Car tago Prov., December 5 and 6, 1962; Rio Damitas, nine miles north of Quepos, San Jos£ Prov., Jan uary 11, 1963; Boca de Barranca, Puntarenas Prov., June 20 and 21, 1963; Rinc6n, Osa Peninsula, Punt arenas Prov., July 1, 1963; three miles north Villa Nelly, Pan American Highway, Puntarenas Prov., August 10, 1963; Bambu, Rio Sixaola, Limdn Prov., November 20, 1964. Vampvrops vittatus: — Flnca La Hondura, San Jos6 Prov., September 6, 1962; Hacienda Moravia, Moravia de Chirrip6, Cartago Prov., October 21, 1962; Finca Las Brumes, four and one half miles southwest of Villa Quesada, Alajuela Prov., April 26, 1963; Tapantl, Cartago Prov., July 30, 1963; three miles 54 south of Culdad Quesada, Alajuela Prov., April 16, 17 and 18, 1964, Monteverde, Ri6 Guacimal, Punt arenas Prov., Nay 15, 1964* Vamnvrodes major:— Los Diamantes, Limdn Prov., April 11, 1964. Chiroderma salvini: — One mile southwest of Canas Gordas, Puntarenas Prov., June 6, 1964. Selected records of Artibeus lamaicensis are as follow: Artibeus lamaicensis:— San Jos£, San Josl Prov., July 7, 1962, October 30, 1962, November 3, 1962, December 28, 1962, March 29, 1963; Ri6 Colorado, five miles north of Liberia, Guanacaste Prov., July 15, 1962; Playa del Coco, Guanacaste Prov., July 20, 1962, April 4, 1963; Boca de Bar ranca, Puntarenas Prov., July 30, 1962, June 20 and 21, 1963; Rincon, Osa Peninsula, Puntarenas Prov., August 17, 1962, June 29, 1963, July 4, 1963; Gromaco Foundation, Puntarenas Prov., August 25, 1962; Finca La Selva, Heredia Prov., September 4, 1962; Finca La Lola, Lim6n Prov., September 24, 1962, July 23, 1963; Hacienda Moravia, Moravia de Chirrip6, Cartago Prov., October 21, 1962; Domin ical, Puntarenas Prov., November 22, 1962; Tur rlalba, Cartago Prov., December 5, 1962; Rio Coronado, thirteen miles northeast of Puerto Cor tes, Puntarenas Prov., March 7, 1963; Quebrada del Zapatero, one mile south of Puerto Cortes, Puntarenas Prov., March 8, 1963; Isla de los Pi- jaros, one mile north of Fortete, Lim6n Prov., May 8, 1963; Las Cuevas, one mile northwest of Lim$n, Llm6n Prov., May 8, 1963; three miles east of Golfito, Puntarenas Prov., March 29, 1964; Los Diamantes, Lim£n Prov., April 8, 1964; Estrellas Oeste, ten miles southeast of Jac6, Puntarenas Prov., November 27, 1964. Selected records of Artibeus lituratus are as follow: 55 Artibeus lituratus:— San Jos£, Sen Jos£ Prov., July 6, 1962, October 30, 1962, November 29, 1962, December 29, 1962; Boca de Barranca, Puntarenas Prov., July 31, 1962; Rio Colorado, five miles north of Liberia, Guanacaste Prov., July 15, 1962; Playa del Coco, Guanacaste Prov., July 19, 1962; Rincdn, Osa Peninsula, Puntarenas Prov., August 16, 1962, June 28, 1963; Finca La Selva, Heredia Prov., September 4, 1962; Flnca La Lola, Lim6n Prov., September 29, 1962; Hacienda Moravia, Moravia de Chlrrip6, Cartago Prov., October 21, 1962; Domini cal, Puntarenas Prov., November 22, 1962; Turrlalba, Cartago Prov., December 7, 1962; three miles north of Villa Nelly, Pan American Highway, Puntarenas Prov., August 9, 1963; Tortuguero, Llm£n Prov., October 25, 1964; Estrellas Oeste, ten miles south east of Jac6, Puntarenas Prov., November 26, 1964. Selected records of Artibeus cinereus are as fol low: Artibeus cinereus:— Playa del Coco, Guanacaste Prov., July 19, 1962; Turrlalba, Cartago Prov., August 9, 1962; Rinc6n, Osa Peninsula, Puntarenas Prov., August 16 and 20, 1962, July 3, 1963; Gro- maco Foundation, Puntarenas Prov., August 24, 1962; Finca La Selva, Heredia Prov., September 4, 1962; Moravia, Moravia de Chirrlpd, Cartago Prov., Octo ber 21, 1962; San Isidro de General, San Jos6 Prov., November 20, 1962; Boca de Barranca, Punta renas Prov., June 21, 1963; Flnca La Lola, LlmSn Prov., July 23, 1963; three miles north of Villa Nelly, Pan American Highway, Puntarenas Prov., August 11, 1963. Artibeus toItecus:— Monteverde, r£o Guacimal, Puntarenas Prov., May 16, 1964. Discussion As indicated by the extensive host lists from both Costa Rica and elsewhere, this species of mite is not only 56 i |widespread in its geographic distribution but also rather I ' ! :catholic in its host a££inities. It does not seem to ex hibit any preference for a particular species of host with in the subfamily Stenoderminae • It seems to occur through out the geographic range of its various host species in Costa Rica, being found in all of the formations of the Tropical and Subtropical Belts and in the Lower Montane Wet and Lower Montane Rain Formations (Map 3) • These mites occur in considerable numbers on their hosts. They are usually found on the wing and tail mem branes but occasionally occur in the ears. Periglischrus vargaai Hoffmann Periglischrus vargaai Hoffmann, 1944, Rev. Inst. Salub. y Enferm. Trop., Mexico, 5(2):91, figs. A, B; Rud nick, 1960, Univ. Calif. Publ. Ent. 17(2):199; Machado- Allison, 1965, Acta Biol. Venez., 4(11):270; Furman, 1966, In Ectoparasites of Panama, In press. Periglischrus soua- mosus Machado-Allison, 1965, Acta Biol. Venez., 4(11):279. (New synonymy). The distinguishing characteristics of this species as described by Hoffmann (1944a) were the presence of \ 10° I Mill! ae Map 3•’ —Collection localities of Periglischrus iheringi. 57 58 i chltinous formations in the opisthosomal region and the shape of the genital plate. These chltinous formations are! now known to be present in varying degrees in all species of the genus Periglischrus. and the genital plate is known to be quite variable within a species. Her description was based only on female specimens. Rudnick (1960) described and illustrated the male and also described and illustrated the female in more detail. He neglected to mention the scale-like pattern usually visible on the dorsal shield of the female. Machado-Allison (1965b) described Periglis chrus scuamoaus as a distinct species mainly on the basis of this pattern on the dorsal shield. All of the other characteristics and the photomicrographs accompanying his description agreed perfectly with the excellent descrip tions and figures of P. vargaai by Rudnick. Examination of Costa Rican specimens of P. vargaai under the phase micro scope revealed that this questionable pattern on the dorsal plate is present but varies with the degree of clearing of the specimens during preparation of the slide mount. Since P. squamosus Machado-Allison is distinguished only by this variable character it is here considered to be a synonym of P. vargaai Hoffmann. 59 The females of P. vargasl can be recognized by the stout* posteroventral setae with attenuated* recurved tips present on femur* genu and tibia IV; by the presence of six pairs of dorsal opisthosomal setae and by the absence of a medio-distal lobe on the palpal tibia* The males can be distinguished from similar species by the presence of two minute dorsal setae on femur II; by the relatively short sternal plate setae and large size. Recorded Hosts and Distribution Mexico: — The original description of this spe cies was based on specimens taken from Leptonvcteris nivalis from Mexico (Hoffmann* 1944a) • She also recorded specimens from Anoura geoffrovi, Stumira lilium and Macrotus californlcus from Mexico. Rud nick (1960) reported specimens from Leptonvcteris nivalis from Mexico. He also indicated that the specimens reported by Hoffmann from Sturnira lilium and Macrotus californlcus appear to represent new species and not P. vargaai. Guatemala: — Specimens are recorded from Anoura geoffrovi and Anoura sp. from Guatemala (Rudnlck* 1960). United States: — Rudnlck reported specimens from Leptonvcteris nivalis from Texas. Venezuela; — Machado-Allison (1965b) extended the range to Venezuela with records from Anoura geoffroyi. A. caudifera and A. cult rat a. He also* again* Indicated in his host distribution chart (cuadro Mo. 11) a single specimen from NOctillo lablalis but did not cite this record in text. 60 Panama:— Furman (1966) records specimens from Anoura geoffrovi and A. cultrata from Panama. He j also records one specimen from Trachons cirrhoius from Panama. Trinidad: — Anoura geoffrovi and A. cultrata are reported as hosts from Trinidad "(Furman, 1966). Costa Rican Hosts and Distribution The collections of Periglischrus vargasl from two specimens of Anoura geoffrovi are new records for Costa Rica although this species of mite has been reported from this host in both Panama and Guatemala. The records are as follow: eleven females, one male and one protonymph from one specimen of Anoura geoffrovi collected eight miles north of San Isidro on Pan American Highway, San Jos£ Prov., March 27, 1964; two females and one male from the same host species collected at Turrlalba, Cartago Prov., May 7, 1964. These mites were all collected from the wing membranes. Only three other specimens of A. geoffrovi and two of Anoura n. sp. were taken in Costa Rica. They were not parasitized with this species of mite nor were any of the other seven species of Glossophaginae collected. Discussion P. vargasl seems to be quite host specific on two 61 j ! genera of bats, Llchonvcteris and Anoura. Furman's (1966) record of a specimen from Trachons cirrhosus is the only well-authenticated record of a bat host belonging to the subfamily Fhyllostominae. As both Trachons and Anoura are primarily cave-dwelling forms, there may be occasional op portunity for accidental transfer of ectoparasites from one host genus to another. The two records of this species from Costa Rica were from Subtropical Wet and Lower Montane Wet Formations (Map 4) • Periglischrus acutisternus Machado-Allison Periglischrus acutisternus Machado-Allison, 1964, Rev. Soc. Mexicans Hist. Nat., 15:200; Machado-Allison, 1965, Acta Biol. Venez., 4(11):262. Periglischrus tintoni Furman, 1966, In Ectopara sites of Panama, In press• (New synonymy)• Machado-Allison (1964) described this species of mite from bats of the genus Phvllostomus. He further des cribed and illustrated it in a succeeding publication (Ma chado-Allison, 1965b). The photomicrographs in this pub lication agree quite closely with specimens from Costa Rica which, after comparison with the Furman type material, had 63 | been Identified as Periglischms tintoni Furman, 1965. Thus, P. tintoni becomes a synonym of P. acutistemus . The following characters suffice to diagnose Peri- glischrus acutisternus females: broadly jug-shaped sternal |plate; five small dorsal propodosomal setae, the first pair I of which are widely spaced; very pronounced apical median lobe on palpal tibia; well-developed posterior seta on coxa III. Males possess flattened peg-like setae ventrally on tarsi III and IV and coarsely barbed dorsal setae on tarsi III and IV. They lack inflated setae ventrally. Recorded Hosts and Distribution Venezuela:— This species of mite is reported from Venezuela (Machado-Allison, 1965b) from Fhyl- lostomu8 elongate [sic], P. concolor [sic?] and P. hastatus. Panama:— Furman (1966) reports specimens from P. discolor. P. hastatus and Trachops cirrhosus from Panama. He also reports a single collection of one female and one protonymph from Mvotis chiloensis. Trinidad: — Specimens were from P. hastatus and T. cirrhosus (Furman, 1966). Colombia: — Specimens are reported from P. elonaatus and P. hastatus (Furman, 1966) • Costa Rican Hosts and Distribution The following collections of Periallschrus 64 acutisternus represent new records for Costa Rica: Phvllostomus hastatus: — Rinc6n, Osa Peninsula, Puntarenas Prov., June 28, 1963; Finca Coyolar, three miles north of Liberia, Guanacaste Prov., August 5, 1964; Finca Don Nicholas, two miles north of Tambor, Puntarenas Prov., November 13, 1964. Phvllostomus discolor:— Playa del Coco, Guana caste Prov., July 20, 21 and 25, 1962; April 4, 1963; one mile west of Liberia, Guanacaste Prov., July 26, 1962; Boca de Barranca, Puntarenas Prov., August 1, 1962; Rincdn, Osa Peninsula, Puntarenas Prov., August 18, 1962, June 28, 1963; Finca La Lola, Limon Prov., September 24, 1962; Playa de Tamarlndo, Guanacaste Prov., January 27, 1963. Trachons cirrhosus: — Boca de Barranca, Punta renas Prov., August 1, 1962, June 20, 1963; Domin ical, Puntarenas Prov., November 23, 1962. Discussion This species of mite was not found on any of the other species of bats in the subfamily Fhyllostomatinae ex amined in this study. The mites were found in almost equal numbers on the ears and wings of Phvllostomus hastatus but were collected only from the wings of P. discolor and Trachops cirrhosus. Another species of spinturnicid mite, Periglischrus torrealbai. was frequently found on the same host specimens as P. acutistemus. These two species of mites were found in almost equal numbers on both the ears and wing membranes of Phvllostomus hastatus. On P. discolor, however, Perl* i ellschrus acutisternus was much less abundant than the other species of mite. This Is a most unusual biological situation In which two species of mites of the same genus appear to be successfully living simultaneously on the same host in approximately the same specific habitat on the host. It is hoped that even more detailed studies of these two species of mites will be made in the future. Periglischrus acutisternus. like the closely re lated specie8, P. torrealbai. was collected from all three formations in the Tropical Belt (Map 5)• Periglischrus olastil Machado-Aliison Periglischrus olastii Machado-Allison, 1964, Rev. Soc. Mexicans Hist. Nat., 15:197; Machado-Allison, 1965, Acta Biol. Venez., 4(11):268. Periglischrus aitkeni Furman, 1966, In Ectopara sites of Panama, In press. (New synonymy). This species was described by Machado-Allison (1964) and further illustrated in a succeeding paper (Ma chado -Alii son, 1965b). The photomicrographs of P. olastii (Machado-Allison, 1965b) are sufficient to allow diagnosis 66 I 67 i of the species and comparison with the descriptions and illustrations of Periglischrus aitkeni by Furman (1966) • ;As Indicated by a number of characteristics, these two spe cies are identical • Periglischrus aitkeni Furman is, therefore, a synonym of Periglischrus olastii Machado-Alli son. P. olastli is closely related to P. iheringi but females may be distinguished from the latter in having the first pair of dorsal propodosomal setae well developed and inserted very near the bases of the second pair of dorsal propodosomal setae. The males also differ in having the first pair of dorsal propodosomal setae displaced far laterad. Recorded Hosts and Distribution Venezuela:— Machado-Aliison (1965b) recorded specimens from Venezuela from Stumira 1 * 1 • > * « " > and £>. ludovici. Furman (1966) also reports spec imens from S. 1 ilium from Venezuela. Panama; — From Panama, Furman (1966) records specimens from S. lilium. S. ludovici and an un usual record from Hoctilio leporinus. Trinidad: — Specimens are recorded only from S. lilium from Trinidad (Furman, 1966) • Rudnlck (1960) indicated that the identification of specimens from Stumira lilium from Mexico (Hoffman, 1944a) 68 as Periglischrus vargasl Is questionable and that these probably represent a new species. In view of the restrict ed host range of P. olast11. it seems likely that these specimens were P. olastli. Costa Rican Hosts and Distribution Periglischrus olastli is reported from Costa Rica for the first time from Stumira lilium. S. ludovici and S mordax. The records from £>. mordax also are new host re cords. The collections are as follow: Stumira lilium: — San Jos£, San Jos6 Prov., July 6 and 7, 1962, October 11, 1962, December 18 and 28, 1962, March 28, 1963; one mile south east of Playa del Coco, Guanacaste Prov., July 17, 1962; Playa del Coco, Guanacaste Prov., July 25, 1962, April 4, 1963; seven miles south of La Georgiana, Pan American Highway, San Jos6 Prov., November 20, 1962; Dominical, Puntarenas Prov., November 22, 1962; Boca de Barranca, Puntarenas Prov., June 22, 1963; five miles southeast of Palmar Norte, Pan American Highway, Puntarenas Prov., March 29, 1964; Estrellas Oeste, Punta renas Prov., November 26, 1964. Stumira ludovici: — Hacienda Moravia, Moravia de Chirripo7 Cartago Prov., October 21 and 25, 1962; two miles south of La Georgiana, Pan Ameri can Highway, San Jos£ Prov., November 20, 1962; San Jos£, San Jos6 Prov., December 20, 1962; Llano Grande, Cartago Prov., December 22, 1962; Monteverde, Rio Guacimal, Puntarenas Prov., April 27, 1963; eight miles south of La Georgiana, Pan American Highway, San Jos6 Prov., July 14, 1963, 69 8even miles south of Le Georgians, Pan American Highway, San Jos! Prov., July 15, 1963; La Georgians, Pan American Highway, San Jos! Prov., July 16, 1963; Tapantl, Cartago Prov., July 30, 1963; Finca El Helechales, nine miles east of Fotrero Grande, Puntarenas Prov., October 3, 1964. Stumira mordax: — Dominical. Puntarenas Prov., November 22, 1964; Rio Damitas, ten miles north of Quepos, San Jos! Prov., January 11, 1963; eight miles south of Le Georgians, Sfn Jos! Prov., July 15, 1963; one mile north of El Angel Falls, Heredia Prov., August 18, 1963. Discussion This species of mite was commonly found on the wing membranes of all three species of bat hosts. Only oc casionally were specimens found in the ears of their hosts. Since P. olaatii seems to be quite host specific to members of the genus Stumira. the record from Noctilio leporinus is surprising and may represent a collection error although the chance of fortuitous or "accidental" parasitism is possible. Perialischrus olastii appears to have the largest altltudinal range of the spinturaicids collected in Costa Rica. Specimens were collected from the Tropical Dry, Tropical Moist, Subtropical Moist, Subtropical Wet, Lower Montane Moist, Lower Montane Rain and Montane Rain 70 | Formations (Map 6) • i Pcrittliachrus parvus Machado-Allison i Periglischrus parvus Machado-Aliison, 1964, Rev. Soc. Mexicans Hist. Nat., 15:195; Machado-Allison, 1965, Acta Biol. Venez., 4(11):270. Periglischrus mlcronvcteridis Furman, 1966, In Ectoparasites of Panama, In press. (New synonymy). This species was described by Machado-Allison (1964) and later illustrated by means of photomicrographs (Machado-Allison, 1965b). Comparison of these photomicro graphs with Furman's (1966) descriptions of Periglischrus mlcronvcteridis indicates that these two species are iden tical. Periglischrus micronvcteridis Furman, therefore, becomes a synonym of Periglischrus parvus Machado-Allison. Females of this species may be recognized by their small size, relatively short dorsal propodosomal setae, en larged medial lobe of palpal tarsus and very small poste rior seta on coxa 111. The males are also small, lack coarsely barbed dorsal leg setae and have a pair of minute setae behind their sternal plate. Map 6.— Collection localities of Periglischrus olastil. 71 72 Recorded Hosts and Distribution 1 Venezuela: — This species is recorded from Venezuela on Micronvcteris hirsute (Machado-Alli son, 1965b)• Panama: — Furman (1966) reports specimens from Micronvcteris megalotis and M. minuta from Panama. Trinidad:--Specimens are reported from M. megalotis from Trinidad (Furman, 1966) • Costa Rican Hosts and Distribution Periglischrus parvus is reported for the first time in Costa Rica from three species of Micronvcteris. The following records from Micronvcteris schmidto-nm also rep resent new host records: fourteen females from one M. schmidto-nm collected at the Rio Colorado, six miles north of Liberia, Guanacaste Prov., July 15, 1962; seven females and four males from five specimens of M. schm*dtnrum col lected two miles south of Playa del Coco, Guanacaste Prov., July 22, 1962. Two female mites were taken from a pooled lot of three specimens of Micronvcteris hirsute collected at Finca La Lola, Lim6n Prov., July 23, 1963. Three males were taken from M. megalotis at Finca El Helechales, nine miles east of Potrero Grande, Puntarenas Prov., October 4, 1964. Discussion The host specificity of these mites to bats of the genus Micronvcteris is well demonstrated by both the Costa Rican records and the records from Venezuela, Panama and Trinidad. These mites were found only on the wing and tail membranes of their hosts in Costa Rica. The few Costa Rican records of Periglischrus parvus came from the Tropical Dry, Tropical Moist and Subtropical Wet Formations (Map 7) • Periglischrus torrealbai Machado-Allison Perigllschrus torrealbai Machado-Allison, 1965, Acta Biol. Venez., 4(11):276. Periglischrus inflatlseta Furman, 1966, In Ecto parasites of Panama, In press. (New synonymy). This distinctive species was first described by Machado-Allison (1965b). Examination of Furman's (1966) description of P. inflatlseta reveals that it is obviously the same species. Periglischrus inflatlseta Furman, there fore, is a synonym of Periglischrus torrealbai Machado- Allison. This species is distinct from all other members of the genus in that the females possess several pairs of [84®1 10° I M .Map 7*— Collection localities of Periglischrus parvus. ventral setae with grossly Inflated bases and acuminate !tips. The males also possess a pair of such setae between coxae IV. I Recorded Hosts and Distribution Venezuela:— Machado-Allison (1965b) reported this species from Phvllostomus hastatus and P. hastatus In his host distribution chart (cuadro NO. 11). Colombia: — Records from Colombia were from P. hastatus (Furman, 1966)• Panama: — Specimens were taken from P. hastatus in Panama (Furman, 1966) • ~ Trinidad: — Furman (1966) also reports specimens from Trinidad from P. hastatus. Costa Rfern Hosts and Distribution The following collections of Periglischrus torreal bai from two species of Phvllostoimis represent new records from Costa Rica. Phvllostomus hastatus; — Dominical. Puntarenas Prov., November 23, 1962; Rincdn, Osa Peninsula, Puntarenas Prov., June 28, 1963; Finca Coyolar, three miles north of Liberia, Guanacaste Prov., August 5, 1964; Finca Don Nicholas, two miles north of Tambor, Puntarenas Prov., November 13, 1964. Phvllostomus discolor:— Playa del Coco, Guana caste Prov., July 19 and 20, 1962, April 14, 1963; one mile west of Liberia, Guanacaste Prov., July 26, 1962; Boca de Barranca, Puntarenas Prov., 76 I July 31, 1962, August 1 and 2, 1962; Rlnc£n, Osa Peninsula, Puntarenas Prov., August 18, 1962, June 28, 1963; Finca La Lola, Lim6n Prov., Septem ber 24, 1962; one mile north of Palmar Norte, Pan American Highway, Puntarenas Prov., March 6, 1963. ' Discussion The host distribution of this species of mite in dicates that it is perhaps more restricted in its range of host species than is the closely related species, Periglis chrus acutisternus. Both of these species are reported from Fhvllostomus hastatus and from P. discolor, but only 1 Periglischrus acutisternus is found on Trachons cirrhosus. The two species of mites occurred in almost equal numbers on Phvllostomus hastatus. but Periglischrus torrealbai was more common on Phvllostomus discolor. Periglischrus torrealbai was found on both the ears and wing membranes of Phvllostomus hastatus. This species of mite was collected only from the three formations in the Tropical Belt, although its hosts have a wider range (Map 8). Periglischrus hopkinsi Machado-Allison Periglischrus hopkinsi Machado-Allison, 1965, Acta Biol. Venez., 4(11):275-276, figs. 3 and 20. Hap 8.— Collection localities of Periglischrns torrealbai. 77 78 The description of this species was based on a single female specimen collected from a single specimen of ; the rather rare bat Lionvcteris snurelli Thomas from Vene zuela. This bat is known only from Colombia, Venezuela and British Guiana (Cabrera, 1957) and was not collected in Costa Rica. Although Periglischrus hopkinsi appears to be related to P. caligus and P. vargasi. sufficient details are not given in the original description to characterize the species without examination of actual specimens • It is hoped that additional specimens of this species will be found and described in greater detail. Periglischrus hopkinsi was not collected in Costa Rica in this study and judging from its host's distribution probably does not occur in Costa Rica. It was, therefore, not included in the keys to the Splntumicldae of Costa Rica. c f Periglischrus herrerai Machado-Allison Periglischrus herrerai Machado-Allison, 1965, Acta Biol. Venes., 4(11):282. Periglischrus desmodi Furman, 1966, In Ectopara sites of Panama, In press. (New synonymy). 79 | ! This species was first described by Machado-Allison! ! (1965b) from the vampire bat, Pesmodus rotundas. Compari- ! son with Furman's (1966) description and illustrations of P. desmodi reveal that it is obviously the same species. I Periglischrus desmodi Furman, therefore, is a synonym of Periglischrus herrerai Machado-Allison. The female of this species can be recognized by the very widely spaced, long first pair of dorsal propodosomal setae, the characteristic shape of the sternal plate and presence of only one minute basal seta on the dorsum of femur II. The males are distinct from all other members of the genus in possessing three pairs of dorsal opisthosomal setae. Recorded Hosts and Distribution Venezuela;— This species is recorded from Pesmodus rotundus in Venezuela (Machado-Allison, 1965. Panama: — Furman (1966) reports specimens from Panama from D. rotundus. Trinidad: — Records from Trinidad are also from D. rotundus (Furman, 1966). Costa Rican Hosts and Distribution Periglischrus herrerai is reported for the first 80 i ; time from Costa Rica from Pesmodus rotundus* The records jare as follow: Three miles south of Playa del Coco, Guana caste Prov., July 23, 1962; Boca de Barranca, Puntarenas Prov., August 1, 1962; Finca La Lola, Lim£n Prov., September 26 and 30, 1962, July 25, 1963; Rio Oro (Uruca), one mile west of Santa Ana, San Jos£ Prov., October 8, 1962; Hacienda Moravia, Moravia de Chirrip6, Cartago Prov., October 23, 1962; Rlncdn, Osa Peninsula, Puntarenas Prov., June 29, 1963. Discussion The occurrence of this mite was only sporadic. In fested hosts, however, usually had a relatively high number of parasites on them. The mites were usually on the wing membranes although a few specimens were found in the ears and on the fleshy protuberances of the bat's face. This is the only species of spinturnicid mite that normally occurs on bats of the genus Pesmodus. Furman (1966) reports several specimens of Periglischrus iherlngj. a species usually found on stenodermlne bats, on Pesmodus rotundus in Panama. This may be an example of fortuitous distribution as the result of predation by the vampire bat, Pesmodus. on some of the smaller species of bats in the subfamily Stenoderminae. 81 j Periglischrus herrerai was collected from all for* nations found In the Tropical and Subtropical Belts in Costa Rica. The host, Pesmodus rotundus. is most common in - , . i these formations but also occurs in most of the other for mations up to the Subalpine Wet (Map 9) • Periglischrus natali Furman Periglischrus natali Furman, 1966, In Ectoparasites of Panama, In press. This somewhat atypical species differs from all other members of the genus in that the peritremes are sharply constricted at the level of coxae III and continue anteriorly as a thread-like structure in the females. In the males this constriction is even more pronounced, with only the posterior portion to the mid-level of coxa III of normal width. The constricted anterior portion is reduced to a vestigial state and almost invisible. Recorded Hosts and Distribution Panama:— This species of mite is recorded by Furman (1966) from Natalus from Panama. Trinidad: — Specimens are reported from Natalus tumidirostrls from Trinidad. 83* 10° >o * • Map 9.— Collection localities of Perialischrus herrerai. 82 Furman also states that "this Is the only spin- turnlcid collected from bats of the genus Natalus." How ever, Hoffmann (1944b) reported Sninturnix carloshpffmarm* (S. f - anus [Banks]) from Natalus mexicanua (N, stra- mlneus) In Mexico. Rudnlck (1960) also recorded j> • carlo- shoffmawnl (S. amerlcanus [Banks]) from Natalus meTlcanua (N. «tr«n<neus). This appears to be a most unusual situa tion in which species of two genera of Spintumicidae are found on the Sfme host species. Unfortunately the specific habitats of these species on the host are not stated, nor are there any records of individuals of these two species of mites actually occurring on the same host simultaneous ly. Costa Rican Hosts and Distribution Only three specimens of Natalus stramineus were collected in Costa Rica. They did not have any spintumi- cid mites on them. Genus Cameronieta Machado-Allison, 1965 Cameronieta Machado-Allison, 1965, Acta Biol. Venes., 4(10):243-258. 84 i TvP€-8PeclC8 Cameronieta thomasl Machado-Allison, 1965. Diagnosis Two separate dorsal shields. Tritosternum absent. Perltreme usually completely dorsal, extending from level of coxa IV to level of coxa I • Female oplsthosoma not ex panded greatly. Unarmed ventral cutlcula of ldlosoma with numerous minute, thom-llke mammlllatlons. Dorsum:— Two separate dorsal shields, relatively small In female; occupying most of ldlosoma In male. Five pairs of setae bordering anterior dorsal shield anterior to stigmata. One pair of setae slightly posteromedial to stigmata. Seven pairs of dorsal oplsthosomal setae In fe male. Perltremes extending from level of coxa IV to level of coxa 1, bending markedly laterad between coxa II and III of male. Unarmed cutlcula between dorsal plates and body margins of males with minute granulations. Venter:— Lacking tritosternum. Female sternal plate twice or more as wide as long with three pairs of setae located on plate. Male sternal plate with five pairs setae; surface with pattern of transverse strlatlons. 85 Epigynial plate narrow, weakly sclerotlzed with pair of genital setae at posterior tip. Female anal plate reduced to narrow sclerotlzed structure with lightly plumose pair of adanal setae located at posterior tip. Male anal plate small with ill-defined margins. Fodosomal cutlcula In re gion of female sternal plate furnished with numerous, mi nute thorn-like mammlllations. Entire ventral cutlcula of ldlosoma of male with similar mammlllations. Legs:— Large caruncles and claws on all legs. Dor sal and lateral setae short to long. Many ventral setae sparsely bipectinate to heavily fringed and palmate. This genus was erected by Machado-Allison (1965a) to contain the species C. thomasl. All of the diagnostic characters given for the genus, with the exception of the heavily sclerotlzed camerostome and the articulation of the sternal plate with the posterior borders of the first pair of coxae, apply equally as well to the two closely related species Periglischrus strandtmanni Tibbetts and P. elon- gatus Furman. As indicated by the wide sternal plate, relatively unexpended female opisthosoma, cuticular mammil- lations, plumose ventral setae and other characteristics as given in the above diagnosis, as well as by their host 86 affinities* these two species are undoubtedly more closely related to C . thomasl Machado - ‘ Allison than to any members of the genus Periglischrus and are* therefore* assigned to the genus Cameronieta Machado-Allison. Cameronieta thomasi Machado-Allison Cameronieta thomasi Machado-Allison* 1965* Acta Biol. Venez., 4(10):243-258* figs. 1-15. Machado-Allison (1965a) described this species from Chilonvcteris rubieinosa (C. pamellii). Furman (1966)* in his discussion of Periglischrus elongatus (Cameronieta elongatus)* mentions two abnormal female specimens from Chilonvcteris pamellii from Panama. He indicates that these specimens may represent genetic freaks or may repre sent a new species. In his discussion he mentions the pre sence of a heavily sclerotlzed camerostome; thus* these specimens were probably examples of Cameronieta thomasi Ma chado -Allison. As Indicated by the key to the species of Camer onieta of Costa Rica* the females of this species may be recognized by the heavily sclerotlzed camerostome formed by the articulation of the sternal plate with the posterior 87 1 ! borders of the first pair of coxae. The males are very isimilar to C. elongatus but may be distinguished by their larger size and by the long distal setae on coxa I. Recorded Hosts and Distribution Venezuela:— This species is reported by Machado- Allison (1965a) in Venezuela from Chilonvcteris rublglnosa (C. parnellii)• Panama?: — The specimens mentioned by Furman (1966) may be examples of this species. Costa Rican Hosts and Distribution Cameronieta thomasl is reported for the first time i in Costa Rica from Chilonvcteris pamellii from the follow ing localities: Cameronieta thomasi:— Las Grutas del Venado, thirteen miles northeast of Arenal, Alajuela Prov., April 9, 1963; two miles west of La Fortuna, Ala juela Prov., April 21, 1964; Turrialba, Cartago Prov., May 6, 1964. Additional specimens, also from Chilonvcteris pamellii. were collected by the present author and Dr. Andrew Starrett at Finca La Lola, Limdn Prov., July 2, 1961. Discussion This appears to be a relatively rare mite, occur ring on only a few specimens of Chilonvcteris pamellii out of several hundred which were examined. They occurred only 88 i on the wing membranes of their host. On several occasions another species of spinturnicld mite, Cameronieta elon gatus. was also found on the wings of the same host. In these cases the latter species was usually more numerous • £• thomasi was not found on any of the other species of the subfamily Chilonycterinae named in the host list. Cameronieta thomasi was collected only from the [ Tropical Moist and Subtropical Wet Formations (Map 10) • Cameronieta strandtmanni (Tibbetts) New Combination Periglischrus strandtmanni Tibbetts, 1957, J. Kan sas Ent. Soc., 30(1):14; Rudnick, 1960, Univ. Calif. Publ. Ent., 17(2):199; Machado-Allison, 1965, Acta Biol. Venez., 4(11):259; Furman, 1966, In Ectoparasites of Panama, In press. This species is closely related to C. elongatus (Furman) from which it differs in that the anterior margin of the female sternal plate is slightly concave rather than with an anterior projection as in C. elongatus. The fe males of both species possess tri-polnted ventral setae on the opisthosoma but C, strandtmanni lacks the prominently plumose setae of C. elongatus. Many of the ventral leg thomasi. 89 90 setae of the females are sparsely blpectinate, thus showing the beginning of a trend towards the prominently fringed, flat setae of C. elongatus. This trend reaches its culmin ation in the many large, flat, very heavily fringed setae of C. thomasi Machado-Allison. The males of C. strandtman ni may be recognized by the expanded blade-like distal seta of coxa I. The description of this species was based on one male and one female collected from Mormoops megalophyla senicula from Frio Cave, Texas. It has been known only ifrom the type specimens and type locality until Furman (1966) reported one additional female specimen collected from Mormoops meealophvla tumidlceps from Tamana Cave, Trinidad. Examination of all of the alcoholic specimens of Mormoops in the Los Angeles County Museum collections, by the present author, revealed only one additional female specimen. This specimen was removed from the wing membrane of Mormoops megalophvla (LACM 18168) collected ten miles west of Progresso, Yucatan, Mexico, by Rook and Petite, June 1, 1961. This appears to be the first record of this species of spintumicld mite from Mexico. 91 i In view of the rarity but apparently widespread j geographic distribution of this mite, it seems reasonable to assume that it will be found ultimately in Costa Rica although it was not collected in the present study. r.fywm-raw-r »«•* elongatus (Furman) New Combination Periglischrus elongatus Furman, 1966, In Ectopara sites of Panama, In press. This species is intermediate in many of its charac teristics between Cameronieta thomasi and C. strandtmanni. It lacks the heavily sclerotlzed camerostome of C. thomasi but is very similar in possessing prominently fringed, flat setae on the ventral surface of the legs. Females may be distinguished from C. strandtmanni by the broad antero median projection of the sternal plate. The males are sim ilar to C. strandtmanni but may be distinguished by the setiform distal setae of coxa 1. Recorded Hosts and Distribution Panama:— Furman (1966) reports this species from Chilonvcteris pamellii and Ptcronotus suapurensia from Panama. Trinidad: — Specimens are reported from Chi lonvcteris pamellii from Trinidad (Furman, 1966). 92 Costa Rican Hosts and Distribution Cameronieta elongatus (Furman) is reported for the first time from Costa Rica from three species of bats of the subfamily Chllonycterlnae. The following records from Pteronotus dawl are also new host records: Pteronotus dawl: — Playa del Coco, Guanacaste Prov., July 25, 1962; five miles west of Atenaa, Alajuela Prov., August 13, 1963; five miles north of Liberia, Guanacaste Prov., August 3, 1964. Pteronotus suanurensls: — Three miles north of Villa Nelly, Pan American Highway, Puntarenas Prov., August 9, 1963; Rio Celba, thirty-five miles south of San Isidro de General, Puntarenas Prov., August 21, 1963. Chilonvcteris narnellll:— Rio Colorado, five miles north of Liberia, Guanacaste Prov., July 14, 1962, January 22, 1963, April 6, 1963; Curlol de Santa Rosa, Guanacaste Prov., January 24, 1963; Playa del Coco, Guanacaste Prov., July 17, 1962; Hacienda Moravia, Moravia de Chlrrlp6, Cartago Prov., October 23, 1962; Rio La Vieja, Pan Ameri can Highway, Puntarenas Prov., March 11, 1963; Las Grutas del Venado, thirteen miles northeast of Arenal, Alajuela Prov., April 9, 1963; two miles west of La Fortune, Alajuela Prov., April 21, 1964; Turrlalba, Cartago Prov., May 6, 1964. Discussion These mites were moderately common on the hosts ex amined In Costa Rica. They occurred primarily on the wing membranes of their host, although a few specimens were also 93 found in the ears* Males and immature forms seemed to be slightly more abundant than females. Cameronieta elongatus was collected from the Tropi cal Dry, Tropical Moist, Tropical Wet and Subtropical Wet Formations (Map 11) • Genus Spinturnix von Heyden, 1826 Snintumix von Heyden, 1826, Isis (Oken), 18(6): 612; Rudnick, 1960, Univ. Calif. Publ. Ent., 17(2):200; Evans, Sheals and Macfarlane, 1961, Terrestrial Acari of the British Isles, p. 140; Delfinado and Baker, 1963, Pa cific Insects, 5(4):905; Baker and Delfinado, 1964, Pacific Insects, 6(4):571; Machado-Allison, 1965, Acta Biol. Venez., 4(10):243; Machado-Allison, 1965, Acta Biol. Venez., 4(11):260; Furman, 1966, In Ectoparasites of Pan ama, In press. Tvne-species Pteroptus mvoti Kolenati, 1856, designated by Opin ion 128 of the International Commission on Zoological No menclature (1936). 85* 83° Mill! * 0 Map 11.— Collection localities of Cameronieta eloneatus. Diagnosis Single dorsal shield. Peritremes short, dorsal over coxa III, anterior end bending ventrad, usually reach ing ventral surface between coxae II and III. Tritostemum small to moderately large, reduced, or lacking. Dorsum:— Single large dorsal shield with several pairs of pores on surface. Three to many pairs of propo- dosomal setae, usually five. One pair metapodosomal setae near stigmata. Few to many opisthosomal setae. Venter:— Sternal plate with three pairs of setae and two pairs of pores in female; three to five pairs of setae and two pairs of pores in male. Epigynial plate small, usually lightly sclerotized, with pair of small gen ital setae on or near posterior tip. Anal plate small, ventroterminal, with pair of adanal setae and post-anal seta. Legs:— Legs stout, all with large claws and car uncles. Dorsal setae mostly long to very long. Ventral setae mostly short, except for lateroventral setae which may be long. Spintumix americanus (Banks) Pteroptus americanus Banks, 1902, Canad. Ent. 34(7):173. Spinturnlx americanus Banks, 1915, Rept. U. S. Dept. Agrlc., No. 108, P. 72; Rudnlck, 1960, Univ. Call£. Publ. Ent. 17(2):218, Furman, 1966, In Ectoparasites of Panama, In press• Spinturnlx carloshoffmannl Hoffmann, 1944, Ann. Inst. Biol. Univ. Nac. Mexico, 15(1):185; Furman, 1966, In Ectoparasites of Panama, In press. This widespread species exhibits little variation In most of Its characters. There Is some variation In size which seems to be correlated with the size of the host spe cies. It also exhibits some variation In the number of dorsal oplsthosomal setae and In the size of the postero lateral seta on tibia III and IV. These variable charac ters used for distinction of these two species, Furman (1966) considers J5. carloshoffmannl to be a synonym of . americanus. The females of jS. americanus may be recognized by the following combination of characters: Oplsthosoma with 10-25 moderate to long setae, those near posterior body margin longer than others; legs with lateroventral setae mostly short; femora I and II with pair of proximal dorsal 97 setae minute. The males have two pairs of long opisthoso- mal setae on unarmed cuticula near posterior apex of dorsal shield and a pair of minute proximal dorsal setae on femora I and 11. Recorded Hosts and Distribution Canada: — Specimens are recorded from Myotis lucifucus in Canada (Rudnick, 1960)• United States:--This species of mite is re corded from many widespread localities throughout the United States, including Alaska, from nine species of bats in the genus Myotis (Rudnick, 1960; Bradshaw and Ross, 1961). Mexico: — Rudnick (1960) reported specimens from Myotis velifer. M. thvsanodes and Natalus mexicanus (N. stramineus) from Mexico. Panama:— Recorded hosts in Panama are Mvotis nigricans. M. chiloensis. M. simus and M. albescens (Furman, 1966)• Venezuela:— Specimens are reported from Myotis nigricans in Venezuela (Rudnick, 1960)• Costa Rican Hosts and Distribution Snintumix americanus is reported for the first time in Costa Rica from the following hosts: Myotis chiloensis:— Four miles south of La Georgiana, Pan American Highway, San Jos£ Prov., July 16, 1963. % 98 Mvotis nigricans;— San Josi, San Josl Prov., July 20, 1963; four miles north of Vpra Blanca, Alajuela Prov., August 15, 1963; El Angel Falls, Alajuela Prov., August 15, 1963; Monteverde, Rio Guacimal, Funtarenas Prov., Kay 16 and 17, 1964; Playuelas, Rio Frio, Alajuela Prov., May 23, 1964; Rio Claro, sixteen miles northeast of San Josl, San Josl Prov., June 30, 1964; thirteen miles south of Liberia, Pan American Highway, Guanacaste Prov., August 3, 1964. Discussion The occurrence of bats infested with this species of mite was sporadic, and infested individuals harbored only a few mites. The mites were found only on the wing membranes of their host. Spintumix americanus was collected from all forma tions found in the Tropical and Subtropical Belts in Costa Rica as well as from the Lower Montane Wet Formation (Hap 12) . Spintumix subacuminatus Furman Spintumix subacuminatus Furman, 1966, In Ectopara sites of Panama, In press. This species resembles Spintumix orri from Antro- zous pallidus from the United States and Mexico but may be distinguished by the absence of a minute dorsal seta on Map 12.--Collection localities of Spintumix 99 100 femora I and II. Females of S.. subacuminatus are characterized by possessing 28-33 dorsal oplsthosomal setae, of which the posterior 6-9 are the longest. The males possess only one pair of dorsal oplsthosomal setae. Recorded Hosts and Distribution Panama:--Furman (1966) reports this species from Rhogeessa tumlda from two localities In Panama. Costa Rican Hosts and Distribution Spintumix subacuminatus is reported from Costa Rica for the first time from Rhogeessa tumlda from the fol lowing localities: Boca de Barranca, Puntarenas Prov., June 22, 1963; Samara, Nicoya Peninsula, Guanacaste Prov., Au gust 22, 1964. A single male specimen taken from Eptesicus an- dinus. from San Isidro de General, San Jos£ Prov., March 27, 1964, is tentatively identified as S. acuminatus. al though it varies from normal specimens in several charac ters . Discussion Only four specimens of Rhogeessa tumida were 101 collected in Costa Rica in the course of this project. Of these, three were infested with this species of mite. The mites occurred only on the wing membranes and in small num bers. Identification of the single specimen from Entesicus andinus is questionable and awaits the collection of fur ther specimens to verify its taxonomic position. The specimens of S. subacuminatus from Costa Rica were all taken in the Tropical Dry Formation (Map 13). Genus Paras pintumix Rudnick, 1960 Parasnintumix Rudnick, 1960, Univ. Calif. Publ. Ent., 17(2):231-232. Type:species Paraspintumix globosus Rudnick, 1960. Diagnosis Female idiosoma greatly distended and globose in engorged and in gravid specimens, normally shaped in fresh ly emerged specimens; single dorsal shield; five pairs of dorsal propodosomal setae anterior to peritremes; peri- trernes short, completely dorsal. Map 13.— Collection localities of Spintumix 102 103 Dorsum:--Single dorsal shield with several pairs o£ pores on surface; five pairs of dorsal propodosomal setae and one pair of long dorsal metapodosomal setae arising at level of stigmata; peritremes short, completely dorsal, overlying coxae III. Venter: — Tri to sternum small; three pairs of margin al setae on female sternal plate; one pair of metastemal setae; anal shield with pair of adanal setae and minute postanal seta. Lees:— Claws and caruncles large; ventral setae short; lateral setae long; dorsal surfaces with many long setae. This genus was erected by Rudnick (1960) for a sin gle species collected only from the anal orifices of vari ous species of Myotis. Paraspintumix globosus Rudnick Par as pintumix globosus Rudnick, 1960, Univ. Calif. Publ. Ent., 17(2):231-232, pi. 48, figs. 1, 2; Machado- Allison, 1965, Acta Biol. Venez., 4(10):243; Furman, 1966, In Ectoparasites of Panama, In press. 104 This species was described by Rudnick (1960) from the anal orifice of Myotis sodalis. The male is unknown. It is reported from several states in the United States from Myotis sodalis. M. grisescens and M. velifer (Rudnick, 1960)• Furman (1966) reports a collection of three females from Finca Lara (Chiriqui.) , Panama, from a bat identified as either Mvotis n. nigricans or Myotis chiloensis. He does not mention the location of the parasite on the host. Numerous specimens of Spintumix americanus were collected from the same host. Paraspintumix globosus probably occurs in Costa Rica, although it was not found on thirty-three specimens of Mvotis nigricans or on the one specimen of M. chiloensis examined in this study. CHAPTER VI DISCUSSION Biology Very little Information has been recorded concern ing the biology of the Spintumicidae. Rudnick (1960) briefly summarized the comments of earlier workers concern ing the ability of these mites to cling very firmly to the wing membranes and to run rapidly on the membranes when disturbed. Among the Costa Rican forms, both the males and females of the genus Spintumix were active and moved rapidly on their hosts. However, in the genera Perfalls- chrus and Cameronieta the females were found tightly at tached to their hosts. Frequently the enlarged, flattened opisthosoma adhered to the surface of the wings or ears by means of a mucilaginous substance which made removal of the mites difficult. Machado-Aliison (1965a) indicated that the females of Cameronieta caused an excavated lesion at their point of attachment. The males and immature forms of 105 106 both Perigllschrus and Cameronieta were not attached In this manner and moved very rapidly when disturbed with a needle or brush. The ability of the females to adhere tightly to the host may explain the occurrence of specimens of Perlglls- chrus and Cameronieta In the ears and on the face of their hosts while Spinturnlx appears to be restricted to the wing and tall membranes. With the exception of females of Perigllschrus and Cameronieta. In which the legs were always extended, two resting positions were noted for the mites. In one, the legs were spread out star-shaped. In the other position, the legs were bent in under the body. Vitzthum (1932) felt that these two positions allowed the mites to adjust to changes in the shape of the wing membranes to which they were attached. Members of the genus Ancvstropus are reported to be restricted to the eyelids and eye canthi, and Merlstaspis is reported also from the eyelids and eye canthi as well as the wing membranes. Members of the genus Paraspintumlx are reported only from the anal orifices of their hosts (Rudnick, 1960)• With these exceptions, and those reported 107 above for Perigllschrus and Cameronieta, the splnturnicid mites are found on the naked surfaces of the wing and tail membranes. Careful examination of the furred parts of the wings and bodies of their hosts revealed no spinturnicids. Juvenile bats usually carried larger numbers of mites than did the adults. This may be a reflection of the more sedentary habits of the juveniles or related to mor phological factors such as less fur or more tender skin or possibly to more obscure physiological factors. Life Cycle The life cycle of the mites of the family Spintum- icidae is spent entirely on the host. There are five stages in the life cycle: egg, larva, protonymph, male or female deutonymph and adult. This pattern is obscured by the fact that the egg and larval stages are passed within the body of the female. The egg or the hexapod larval stage may be seen in well-cleared specimens of pregnant fe males. Only one young at a time is produced. Although the exact relationship between the developing larva and the fe male is unknown, the black caecal configurations present in engorged adults are sometimes seen in well-developed 108 larvae. Indicating that perhaps some form of nourishment is supplied by the female. The female gives birth directly to the eight-legged protonymph which molts once to the deu- tonymph stage. There are male and female deutonymphs which can be associated morphologically with their respective adults. The deutonymphs molt once becoming sexually mature male or female adults. Copulation was not observed but sperm transfer is probably effected by means of the specialized spermato- phoral process on the male chelicera, as is the case in most of the other Mesostigmata. The mouth parts of the protonymph, deutonymphs and adults are of the sucking type. These mites are apparently able to feed on the blood of their hosts in all three of these stages as is evidenced by the presence of intact blood cells in the digestive tract of freshly-engorged specimens• The Spinturnicidae possess many of the modifica tions commonly exhibited by ectoparasitic forms. These in clude flattened bodies, thickened legs and enlarged claws. In addition to these obvious morphological adaptations, the specialized life cycle in which the independent egg and 109 larval stages are eliminated, undoubtedly increases the survival rate. Among the other major groups of ectopara sites, for which the life cycle Is known, this type of mod ification occurs only In the Puplpara of the order Dlptera. In this case the female retains the egg and larva within her body and gives birth to a fully-developed larva ready to pupate. The dlpteran families Streblldae and Nycter- ibildae, also found exclusively on bats, have this type of life cycle. The mature larvae of these flies are deposited on the substratum, usually close to the roosting place of the hosts. The larvae pupate Immediately. As soon as the adults emerge from the puparla, they seek out a host. Thus, there Is ample opportunity for transfer to another host species (Ryberg, 1947). Independent stages, off the host, also occur In the life cycle of the Siphonaptera. In the Mallophaga and Ano- plura the entire life cycle Is again normally passed on the host, with the exception of the human body louse which spends much of Its life on the host's clothing. The significance of these varied life cycles In terms of their influence on host-paraslte relationships is discussed in the section on host specificity. 110 Blogeographv The reported collections of the various species of mites within the plant formations of Costa Rica cannot be Interpreted quantitatively due to variations In sampling In different areas. Also, the presence and relative abundance of suitable host species may vary from one formation to an other. However, these records may be Indicative of certain distributional trends for some of the species where ade quate samples are available. Several species of Perigllschrus appear to be re stricted In their distribution to formations in the warmer Tropical and Subtropical Belts although their hosts occur in a wider range of formations. The closely related spe cies Perigllschrus torrealbai and P. acutisternus are re corded only from formations in the Tropical Belt, while their hosts, Phvllostomus hastatus and P. discolor, occur in both the Tropical and Subtropical Belts. Perigllschrus calistus was collected in formations in the Tropical and Subtropical Belts although its host, Glossonhaga. occurs in these belts and in the Lower Montane Belt. The vampire bat, Desmodus rotundus. is widespread in most of the forma tions in Costa Rica up to the Subalpine Wet, but Ill Perigllschrus herrerai. the mite usually associated with Deswodus. is found on its host only in the Tropical and Subtropical Belts. Some species of Pegjglischrus. however, appear to occur throughout the range of their hosts in Costa Rica. Thus, the very common species, Perigllschrus iheringi. is found in almost all of the plant formations of Costa Rica in which the hosts are known to occur. Perigllschrus ojas- tii. found on bats in the genus Stumira, has the largest altitudinal range of any of the species of Perigllschrus collected in Costa Rica. The host genus also has a wide altitudinal range, although certain of the species within the genus are somewhat restricted in their altitudinal dis tribution. Due to the relatively stable microenvironment in which the mites live, it is difficult to visualize what climatic factors associated with the various plant forma tions are responsible for these distributions. These fac tors probably affect the mites only in an indirect manner by influencing their hosts. Since the mites of the genus Perigllschrus are found only on groups of bats of Neotropical origin, it may 112 be assumed that the mites are also of Neotropical origin. It might thus be speculated that this restriction of many of the species of Perigllschrus to the warmer portions of the range of their hosts Is the result of adaptations, on the part of the mites, to the warmer regions of the Neo- tropics. Many parasites are more conservative in their evolution than are the hosts (Mayr, 1957). Thus, some of the species of Perigllschrus may still be restricted to warmer regions while their bat hosts have developed mech anisms enabling them to invade cooler regions. Although collection records of the genus Cameronieta are rather lim ited, on the basis of the known distribution and host af finities it would appear that this genus is also probably of Neotropical origin. In contrast to the genus Perigllschrus. the members of the genus Spintumix appear to be primarily temperate in their affinities. They are found on members of the cosmo politan family Vespertlllonidae and are widespread and com mon In the Holarctlc Region. The genus Mvotis Is the most widely distributed genus of bats (Darlington, 1957) and Is parasitized by a number of species of Spintumix in various parts of its range. Spintumix americanus is reported from 113 nine species of bats in the genus Mvotis in North America and appears to be primarily a temperate region species. In Costa Rica it occurred more often in the Lower Montane and Subtropical Belts but was also found in formations in the Tropical Belt. The known spintumicid fauna of Costa Rica thus ap- pears to reflect the origins and distributions of its chi- ropteran hosts. By far the vast majority of the bats of Costa Rica are of southern origins (Hershkovitz, 1958; Stuart, 1964). Likewise, of the twelve species of spin- tumicid mites recorded from Costa Rica, ten of these ap pear to be Neotropical regionalites. Indeed, even Spin tumix subacuminatus. found on bats of the genus Rhogeessa. may prove to be of southern origin. Although the Atlantic and Pacific lowlands of Costa Rica are at least partially divided by two central mountain ranges, the Cordillera Guanacaste and the Cordillera Tala- manca, this barrier has little influence on the distribu tion of the Chiroptera (Ryan, 1963). Likewise, where ade quate host collection records are available, there appears to be no difference between the Atlantic and Pacific low lands in terms of the occurrence of spintumicid mites. 114 Host Specificity Analysis of recent papers (Delflnado and Baker, 1963, Baker and Delflnado, 1964) reveals a close correla tion between the genera of Splnturalcldae and the families of bats. Rudnick (1960) pointed out that almost all excep tions to this host specificity were found among very old records from the Old World, the collection records of which are difficult to verify. Table 1 presents the distribution of the genera of Splnturnlcidae on the families of bats. This table does not Include a small number of records which probably represent cases of adventitious parasitism or col lection errors. There are three genera, Ancystropus, Oncoscelus and Merlstaspls. restricted to the fruit bats of the family Fteropldae. Although the mites In these three genera ex hibit a number of morphological specializations related to parasitism, they appear to be the most primitive of the splntumlclds In terms of many features of their basic mor phology Including the presence and shape of their trltos- temum, the presence of a single dorsal shield, the size and position of the perltremes and the dorsal setatlon. In these features they are distinct from the remaining genera. TABLE 1 1X5 DISTRIBUTION OF GENERA OF SPINTURNICIDAE ON FAMILIES OF CHIROPIERA Genera of Splnturnicldae Families of Chiroptera C O g s 4 - > a ► I Oncoscelus m • r l O n < d ■u « o •H U £ Evndhovenia Paranerislischrus Periglischrodes* Perielischrus Cameronieta Spintumix Parasnintumix Pteropidae X X X Rhinolophidae X X Hipposideridae X Phyllostomatidae Chilonycterinae X Phyllo stomatinae X Glossophaginae X Caro11inae Sturninae X Stenoderminae X Desmodontidae X Natalidae X X Vespertilionidae X X *Host unknown 116 Evidence of phylogeny among the other genera is largely ob scured by secondary acquisition of adaptive characters or reductions that tend to conceal basic affinities. Until more detailed comparative morphological studies of all stages in the life cycle are made, it is impossible to con struct a phylogenetic series for the Spintumicidae. It is tempting to speculate that the family originated when bats first became differentiated from a primitive ancestor and that the three genera now found on the Pteropidae were sep arated at the very early age when the Chiroptera split into two major phylogenetic lines, the Megachiroptera and the Microchiroptera. An analogous situation exists in the flea family Ischnopsyllidae, which contains one small subfamily, the Thaumapsyllinae, restricted to the Megachiroptera and a larger subfamily, the Ischnopsyllinae, restricted to the Microchiroptera (Holland, 1964). At the species level the Spintumicidae exhibit varying degrees of host specificity. Some species of mites appear to be restricted to a single host species, others to a single host genus and still others to related genera, frequently in the same subfamily. Tables 2 and 3 present the distribution of the spintumicid species on the 117 TABLE 2 DISTRIBUTION OF SPECIES OF PERIGLISCHRUS ON SPECIES OF CHIROPTERA IN COSTA RICA Species of Perigllschrus • r l •H rH Phy 1 lo s tomat idae « Phy 1 lo s tomat inae Micronvcteris megalotis X Micronvcteris schmidtorum X Micronvcteris hirsuta X Lonchorhina aurita Macro phy Hum macrophyllum Phvllostomus hastatus X X Phvllostomu6 discolor X X Trachops cirrhosus X Chrotopterus auritus Vampvrum spectrum Glo s sophaginae Glossophaga soricina X 118 TABLE 2--Continued Phy1lo stomatidae Species of Perigllschrus OB 1 I as t f l ► < 0 iH t o o 4J a > J3 •y as t o t o a Glossophaglnae Glossonhaga commissarisi X LonchoDhvlla concava Lonchophvlla robusta Anoura geoffrovl X Anoura n. so. Choeronlscus godmanl Hvlonvcterls underwood! Lichonvcteris obscura Carolliinae Carollla persolclllata CaroIlia castanea Carollla subrufa TABLE 2- ‘ -Continued 119 Species of Perjglischras Phyllostomatidae 09 C0 o I 09 * 0 W S 0 J A i-l e 0 ) b s ■a b « A 09 a o > 4 - 1 0 0 < r l 4J i ■M 0 9 <0 Sturairinae Stumira lilium X Stumira ludovici X Stumira mordax X Stenoderminae Uroderma bilobatum X VanrovroDs helleri X Vampvroos vittatus X Vamovrodes malor X Vampvressa pusilla Chiroderma villosum X Chiroderma salvini X Ectophylla alba TABLE 2--Continued 120 Species of Periglischrus Fhyllostomatidae a t 3 <8 a ( X e •H U a J3 0 0 ( 8 0 1 S > < 8 .O rl 0 8 a > 5 < 8 « U « 4S 09 I t t > 4J at • r l 1 0 0 C 8 S t enoderminae Artibeus iamaicensis X Artibeus lituratus X Artibeus cinereus X Artibeus toltecus X Centurio senex Desmodontidae Desmodus rotundus X TABLE 3 121 DISTRIBUTION OF SPECIES OF CAMERONIETA AND SPINTURNIX ON SPECIES OF CHIROPTERA IN COSTA RICA Genera and Species of Spinturnicidae Chiroptera C t f 4J 0 n § 0 ■U V CO 2 g o 1 “ I 0 B o CO 9 -u 5 04 CO o a § ! 'o m Phyllostomatidae Chilonycterinae Chilonvcteris narnellii X X Chilonvcterls psilotis Pteronotus dawi X Pteronotus suanurensis X Vespertilionidae Mvotis niericans X Mvotls chiloensis X Entesicus fuscus Eotesicus Raumeri EDtesicus andinus Dasvpterus esca Rhoeeessa tumida X 122 pertinent species of bats collected in Costa Rica. A high degree of specificity is exhibited by most of the forms. In general the known systematic relationships of the mites agree with the accepted systematic relationships of the Chiroptera. Thus* Periglischrus calieus and P. vargasi seem to be closely related, as indicated by a number of morphological characters, and are found on members of the same subfamily, the Glossophaginae. Periglischrus torreal- bai and P. acutistemus are also closely related and both are found on Phvllostomus. Exceptions to these phylogenetic relationships are also of interest. Periglischrus herrerai is found on bats of the family Desmodontidae, although the usual hosts of Periglischrus are phyllostomatid bats. It is generally recognized, however, that the Desmodontidae are closely re lated to the Fhyllostomatidae and are placed in a separate family only on the basis of certain specialized morpholog ical features related to their sanguivorous havits. The two families are placed in the same superfamily. Thus this exception to the usual phylogenetic relationship between host and parasite is more apparent than real. 123 The new species, Periglischrus natali. described by Furman (1966) from bats of the family Natalidae is unusual both in its occurrence on a family of bats distantly re lated to the Phyllostomatidae and in certain of its morpho logical characters. This species may be an example of a form that has effectively transferred to a new host group and developed some distinctive characters as the result of isolation and speciation on the new host. Examples of this type of transfer are usually more common in groups of ectoparasites that are less closely as sociated with their hosts than are the spinturaicids. Nu merous examples of this type of secondary adaptation are found among the fleas (Holland, 1964). The ecology and the life cycle of both the parasite and the host must be considered in attempting to understand the complex relationships that exist between these two groups. Adventitious or accidental parasitism is a fre quent phenomenon among various groups of parasites. In these cases, chance transfer to a new host group occurs, frequently between hosts occupying a similar environment. Usually the parasite is not able to live successfully on the new host. However, if it is able to adapt to the new 124 host, Isolation and speciation can take place. The many scattered records of mite species on other than their usual hosts are probably examples of adventitous parasitism, al though the exact relationship to the new host is uncertain. While many species of colonial bats appear to offer ample opportunity for parasite transfer, closer examination of their roosting sites in such places as caves and hollow trees reveals that they tend to segregate by species in their selection of specific landing places. Thus, while several species of bats may occupy the same cave, they are usually isolated from each other. Since the life cycle of the mite is spent entirely on the host, the chance for transfer is further limited. The chances for adventitious transfer to different hosts are much better in the case of the Siphonaptera and the dipterous families Streblidae and Nycteribiidae in which part of the life cycle is spent off of the host. The food habits of many predators are often re flected in the ectoparasite fauna found on them (Linsdale and Davis, 1956). The occurrence of Periglischrus iheringi on Desmodus rotundus may be an indication that this sangui vorous bat occasionally feeds on the smaller members of the 125 subfamily Stenoderminae, although other means for transfer are also possible. The transfer of a parasite species to a host al ready harboring a congeneric species of parasite results In segregation or competition and the ultimate elimination of the less successful species, unless there Is an unoccupied ecologic niche available on the host (Mayr, 1957).“ The oc currence of Periglischrus torrealbai and P. acutisternus on the same host appears to be in violation of this basic bio logical principle. Further study is necessary to find out the exact ecologic niche occupied by each of these two species• CHAPTER VII SUMMARY AND CONCLUSIONS 1. As a result of three years of field work in Costa Rica, 2,864 bats of eight families were collected and examined for ectoparasites. Spinturnicid mites were found on bats of the families Fhyllostomatidae, Desmodontidae and Vespertilionidae. 2. A historical review of the family Spinturn- icidae, with emphasis on publications since the revision of the family by Rudnick (1960), is presented. Sixty species of ten genera are recognized and arranged systematically. Twelve species of mites of three genera are reported for the first time from Costa Rica. 3. Keys to the genera and species of Spinturn icid ae known to occur in Costa Rica are given. Forms ex pected to be found in Costa Rica are also included in these keys. 126 127 4. Systematics, hosts, distribution and host-para site relationships of the Costa Rican species of the genera Periglischrus, Cameronieta. Spinturnix and Paraspintumix are discussed. 5. The following names are reduced to synonyms as indicated: Periglischrus setosus Machado-AllIson for Peri glischrus caligus Kolenati; P. squamosus Machado-Allison for P. vargas 1 Hoffmann; P. inflatiseta Furman for P. tor- realbai Machado-Allison; P. desmodi Furman for P. herrerai Machado-Allison; P. tlptoni Furman for P. acutisternus Machado-Allison; P. aitkeni Furman for P. ojastii Machado- Allison; P. micronycteridis Furman for P. parvus Machado- Allison* 6• Periglischrus strandtmanni Tibbetts and Peri glischrus elongatus Furman are placed in the genus Camer onieta Machado-Allison. 7. Mites in the genus Periglischrus occur prima rily on bats of the family Phyllostomatidae. Exceptions are Periglischrus herrerai which is found on the closely- related family Desmodontidae, and P. natali. found on bats of the family Natalidae. In general, the primary hosts of a given species of Periglischrus are members of a single 128 bat genus or of closely-related genera. P. iheringi Is ex ceptional In that It regularly occurs on seven genera of the subfamily Stenoderminae. 8. Members of the genus Cameronleta occur on three genera of the subfamily Chllonycterinae. Species in the genera Spinturnix and Parasplnturalx are found primarily on bats of the family Vespertilionldae. 9. Members of the genera Periglischrus and Camer- onieta occur on the face and ears of their hosts as well as on the wings. Spinturnix occurs only on the wings. 10. The life cycle of the Spintumlcidae is spent entirely on the host and consists of the following stages: egg, larva, protonymph, male or female deutonymph and adult. The egg and larval stages are passed within the body of the adult female. 11. Periglischrus caligus. P. torrealbai. P. acuti- sternus and P. herrerai are restricted to the warmer parts of the range of their hosts. P. iheringi and P. olastii occur throughout the range of their hosts in Costa Rica. 12. Collection records and host affinities indicate that the genera Periglischrus and Cameronieta probably originated in the Neotropical Region, while Spinturnix 129 amerlcanus Is of Holarctic origin. 13. Tables presenting the distribution of genera of Spinturnlcidae on families of Chiroptera and host distribu tions of Costa Rican species of Spinturnlcidae are given. 14. Host specificity of the Spinturnlcidae corrob orates the current concept of phylogeny of the Chiroptera in most cases. 15. Exceptions to expected phylogenetic relation ships are discussed in terms of host transfer and adventi tious parasitism. The ecology and life cycle of both the parasite and the host are important factors in determining the degree of isolation between populations existing on different hosts. 16. Multiple invasions of hosts and competition be tween congeneric species of mites are discussed. LITERATURE CITED LITERATURE CITED Baker, E. \ 1964 Baker, E. 1 1952 Banks, N. 1902 1915 Benoit, P. 1961 Bradshaw, 1961 Cabrera, A 1957 ., and D. Delfinado. Spinturnlcidae of South East Asia and the Pacific region. Pacific Insects 6(4):571-591. ., and 6. W. Wharton. An introduction to acarology. Macmillan Co., New York. 465p. New genera and species of acariens. Canadian Entomol. 34:171-176. The Acarlna or mites. A review of the group for the use of economic entomologists. Rep. U. S. Dept. Agr. 108. 153p. L. 6. Periglischrus triaenopsis n. sp. parasitic on Trlaenops afer Peters (Acarina: Mesostigmata: Spinturnlcidae). J. Parasitol. 47(3):397-398. V. R., and A. Ross. Ectoparasites of Arizona bats. J. Arizona Acad. Sci. 1(4):109-112. Cat&logo de los mamiferos de America del Sur. Rev. Museo Argentine Cien. Nat., Bernardino Rivadavia, Buenos Aires, Cien. Zool. 4(1):1- 307. 131 132 Collins* B. J. 1931. Confused nomenclature of Nvcterlbia Latreille* 1796 and Spinturnix Heyden 1826. Nat. Inst. Health Bull. 155:743-765. Condit, P. K.* and 6. L. Humphrey. 1961. Rabies in bats. California's Health* Calif. State Dept. Publ. Health 19(6):14-43. Constantine* D. G. 1962. Rabies transmission by nonbite route. Public Health Repts. 77(4):287-289. Corristan* E. C.* L. C. La Mbtte* Jr.* and D. 6. Smith. 1956. Susceptibility of bats to certain encephalitis viruses. Federation Proc. 15(1):1905. Darlington* P. J. 1957. Zoogeography: The geographical distribution of animals. John Wiley and Sons* Inc.* New York. 675p. Dean* W. D., K. T. Maddy* E. L. Cockrum and H. G. Crecelius. 1960. Rabies in insectivorous bats of Arizona. Arizona Med. 17(2):69-77. Delfinado* M. D.* and E. W. Baker. 1963. Mites of the family Spinturnlcidae from the Philippines. Pacific Insects 5(4):905-920. Evans* G. 0. 1957. An introduction to British Mesostigmata (Aca- rlna) with keys to families and genera. J. Linn. Soc. Lond.* Zool. 43(291):203-259. Evans, G. 0., J. G. Sheals and D. Macfarlane. 1961. The terrestrial Acarl of the British Isles. British Museum (Nat. Hist.)* London. 219p. Furman* D. P. 1966. The spintumicid mites of Panama (Acarina* Spinturnlcidae) • In Ectoparasites of Panama. R. L. Wenzel* editor* Chicago Nat. Hist. Mus. In press. 133 Gilyard, R. 1945. Hall, E. R., 1959. Hershkovitz, 1958. Hoffmann, M. 1944a. 1944b. Holdridge, L 1947. 1953. j t 1964. Holland, G. 1964. Bat transmitted paralytic rabies. Cornell Vet. 35:195-209. and K. R. Kelson. The mammals of North America. Vol. 1. Ronald Press, New York. 546p. P. A geographic classification of Neotropical animals. Fleldlana: Zool. 36(6):581-620. A. Periglischrus vargas1 n. sp. (Acarina:Parasi- tldae) . Rev. Inst. Salubrldad y Enfermedades. Trop. 5(2):91-96. Un nueve acaro par&sito de murclelagos. An. Inst. Biol., Univ. Nac. Mex. 15(1):185-189. . R. Determination of world plant formations from simple climatic data. Science, 105(2727): 367-368. La vegetacldn de Costa Rica, p. 32-33. In Atlas Estadlstlco de Costa Rica. Casa Graf- lea, San Jose. Life zone ecology. Tropical Science Center, San Jose. 117p. Evolution, classification, and host relation ships of Siphonaptera. An. Rev. Entomol. 9: 123-146. International Commission on Zoological Nomenclature. 1936. Opinions rendered by the International Commis sion on Zoological Nomenclature. Smithsonian 134 Misc. Collect. (Publ. 3395), 73(8):l-44. (Opinion 128). Irons, J. V. 1955. A study of bats as possible reservoirs of the rabies virus. First annual progress report, Nat. Inst. Health Research Grant E 758 (C) (Oct. 1, 1954-Sept. 30, 1955). 15p. Keegan, H. L. 1943. Some host records from the parasitological collection of the State University of Iowa. Bull. Brooklyn Entomol. Sec. 38:54-57. Kent, J. R., and S. M. F inego Id. 1960. Human rabies transmitted by the bite of a bat. New England J. Med. 263:1058. Kolenati, F. A. 1856. Die Parasiten der Chlroptern. Brunn: Rudolph Rohrers Erben. 51 (Republished in 1857 in Dresden by Rudolph Runtze)• 1857. Synopsis prodroma der Flughaut-Milben (Pter- optida) der Fledermause. Wien. Entomol. Mon- atschr. 9(2):59-61. Linsdale, J. M., and B. S. Davis. 1956. Taxonomic appraisal and occurrence of fleas at the Hastings Reservation in Central Calif. Univ. Calif. Publ. Zool. 54(5):293-370. Lloyd, J• J• 1963. Tectonic history of the South Central-American orogen. In Backbone of the Americas. Sym posia, Associated Internet. Petroleum Consult ants. Memoir 2:88-100. Machado-Allison, C. E. 1964. Notas sobre Mesostigmata Neotroplcales II. Cuatro nuevas especies del glnero Periglis chrus (Acarina, Mesostigmata, Spinturnlcidae). 135 Rev. Soc. Mex. Hist. Nat. 15:193*207. Machado-Allison, C. E. 1965a. Notas sobre Mesostigmata Neotropicales III. thomasl: nuevo g£nero y nueva especle parasita de Chiroptera (Acarina, Spin- tumicidae). Acta Biol. Venez. 4(10):243-258. 1965b. Las especies Venezolanas del g£nero Periglis chrus Kolenati 1857 (Acarina, Mesostigmata, Spinturnlcidae). Acta Biol. Venez. 4(11):259- 348. Maldonado-Koerdell, M. 1964. Geohistory and paleogeography of Middle Amer ica. In Handbook of Middle American Indians. Vol. I, Natural environment and early cul tures. R. C. West, Editor, Univ. of Texas Press, Austin. Mayr, E. 1957. Evolutionary aspects of host specificity among parasites of vertebrates, p. 7-13. In First symposium on host specificity among parasites of vertebrates. Univ. of Neuchatel, Switzer land. O'Connor, J. L«, L. C. Rowan and J. J. Lawrence. 1955. Relationships between the flying fox (genus Pteronus) and arthropod-borne fevers of North Queensland. Nature 176(1):472. Oudemans, A. C• 1902. Acarologische aanteekeningen. Entomol. Ber., Amsterdam 1(6):36-39• 1903. Notes on Acari. Fifth series. Tijdschr. En tomol. 45:123-150. Pawan, J. L. 1936. The transmission of paralytic rabies in Trini dad by the vampire bat (Desmodus rotundus 136 Rudnick, A. 1960. Ryan, R. M. 1963. Ryberg, 0. 1947. Shacklette, 1962. Slud, P. 1964. Stuart, L. 1964. Tibbetts, T 1957. Till, W. M. 1958. Wagner, 1840). Ann. Trop. Med. 30:101-130. A revision of the mites of the family Spin- turaicidae (Acarina). Univ. Calif. Publ. En tomol. 17(2):157-284. The biotic provinces of Central America as in dicated by mammalian distribution. Acta Zool. Mex. 6:1-55. Studies on bats and bat parasites. Univ. Lund and Zool. Lab., Agr., Dairy and Hort. Inst, of Alnarp. Stockholm, Sweden. 330p. M. H., F. H. Diercks and N. B. Gale. Hlstoplasma cansulatum recovered from bat tis sues. Science 135(3509):1135. Birds of Costa Rica. Bull. Am. Mus. Nat. Hist. 128:1-399. Fauna of Middle America. In Handbook of Middle American Indians. Vol. I, Natural environment and early cultures. R. C. West, Editor, Univ. of Texas Press, Austin. Description of a new Periglischrus from a bat, Mormoops megalophvlla senlcula Rhen, together with a key to the species of Periglischrus (Acarina, Spinturnlcidae). J. Kansas Entomol. Soc. 30:13-19. Five new species of mites (Acarina: Laelap- tidae and Spinturnlcidae) parasitic on bats in the Ethiopian region, with a key to the spe cies of the genus Periglischrus. Rev. Suisse de Zoologie 65:241-258. 137 Tosi, J. A., Jr. I960. Zonas de vida natural en el Peru, memoria ex- plicativa sobre el mapa ecologico de Peru. Bol. Tec. No. 5, Zona Andina# Proy. 39# Pro- grama de Coop. Teen.# Inst. Interam. Cien. Agr.# Lima# Peru. 271p. 1964. Climatic control of terrestrial ecosystems: a report on the Holdridge model. Econ. Geog. 40 (2):173-181. Turk, F. A. 1950. Studies of Acari. VI. Parasitic mites from mammalian hosts obtained in Ceylon. Para sitology 40:63-76. Vitzthum# H. G. 1932. Neue parasitische Fledermaus-Milben aus Vene zuela. Ztschr. Parasitenk., Berlin, 4:1-47. 1940-1943. 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Casebeer, Richard Stratton
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Systematics and host relationships of the mites of the family Spinturnicidae In Costa Rica (Acarina: Spinturnicidae)
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1966-06
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Garth, John S. (
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