SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Biology of Bats of the New World Family Phyllostomatidae. Part II Edited by Robert J. Baker, J. Knox Jones, Jr., and Dilford C. Carter .. No. 13 ... ..... .'If ,, June 1977 Robert D. Bradley SPECIAL PUBLICATIONS THE MUSEUM TEXAS TECH UNIVERSITY Biology of Bats of the New World Family Phyllostomatidae. Part II Edited by Robert J. Baker, J. Knox Jones, Jr., and Dilford C. Carter No. 13 June 1977 TEXAS TECH UNIVERSITY Cecil Mackey, President Glenn E. Barnett, Executive Vice President Regents .—Judson F. Williams (Chairman), J, Fred Bucy, Jr., Bill E. Collins, Clint Form- by, John J. Hinchey, A. J. Kemp. Jr., Robert L. Pfluger, Charles G. Scruggs, and Don R. Workman. Academic Publications Policy Committee. —^J. Knox Jones, Jr, (Chairman), Dilford C. Carter (Executive Director and Managing Editor), C. Leonard Ainsworth, Harold E. Dregne, Charles S. Hardwick, Richard W. Hemingway, Ray C. Janeway, S. M. Kennedy, Thomas A. Langford, George F. Meenaghan, Marion C. Michael, Grover E. Murray, Robert L, Packard, James V. Reese, Charles W. Sargent, and Henry A. Wright. The Museum Special Publications No. 13 364 pp. 17 June 1977 $16.00 Special Publications of The Museum are numbered separately and published on an irregular basis under the auspices of the Dean of the Graduate School and Director of Academic Pub¬ lications, and in cooperation with the International Center for Arid and Semi-Arid Land Studies. Copies may be obtained on an exchange basis from, or purchased through, the Ex- ' change Librarian, Texas Tech University, Lubbock, Texas 79409. Texas Tech Press, Lubbock, Texas 1977 CONTENTS Introduction . 5 Endoparasites . 7 John E. Ubelaker, Robert D. Specian, and Donald W, Duszynski, Department of Biology, Southern Methodist University; Department of Biology, Southern Methodist University, Dallas, Texas 75222; Department of Biology, The University of New Mexico, Albuquerque, 87131. Ectoparasites . 57 James P. Webb, Jr., and Richard B. Loomis, Department of Biology, California State University, Long Beach, 90840. Oral Biology . 121 Carleton J. Phillips, Gary W. Grimes, and G. Lawrence Forman, Department of Biology, Hofstra University; Department of Biology, Hofstra University, Hempstead, New York 1 1550; Department of Biology, Rockford College, Rockford, Illinois 61101. Echolocation and Communication . 247 Edwin Gould, School of Hygiene and Public Health, The Johns Hopkins University, Baltimore, Maryland 21205. Thermoregulation . 281 John J. McManus, Department of Biology, Farleigh Dickinson University, Madison, New Jersey 07940. Feeding Habits .293 Alfred L. Gardner, U.S. Fish and Wildlife Service, National Fish and Wildlife Laboratory, National Museum of Natural History, Washington, D.C. 20560. Movements and Behavior . 351 M. Brock Fenton and Thomas FI. Kunz, Department of Biology, Carleton University, Ottawa, Canada KIS 5B6 and Department of Mammalogy, Royal Ontario Museum; Department of Biology, Boston University, Boston, Massachusetts 02215. INTRODUCTION Because of their adaptive diversity and, in many instances, unique morpholog¬ ical attributes, bats of the family Phyllostomatidae long have fascinated biolo¬ gists. Known only from the New World, most genera of phyllostomatids are limited distributionally to tropical environments, but some representatives occur as far north as the southwestern United States and others southward to the northern parts of Argentina and Chile; some species also are distributed on the Bahamas and islands of the Greater and Lesser Antilles. With the advent in recent years of improved methods of collecting bats, a tremendous wealth of information on phyllostomatids has been gathered, and it is the purpose of this publication, which ultimately will contain more than 20 individual chapters, to bring these data together in order to assess what now is known about the family and to provide a departure point for further studies. Owing to the large number of contributions, all of which were solicited by us from persons we felt to be knowledgeable of the subject matter, and the fact that several contributions are necessarily lengthy, the decision was made to group chapters into three parts, each separately numbered as a Special Publica¬ tion of The Museum at Texas Tech University. In order to establish a workable approach by which reference could be made consistently to taxa throughout the series, an annotated checklist by Jones and Carter (published in the first part of the series) was circulated to all authors. Each was asked to follow the nomenclature and systematic arrangement in the checklist or, alternatively, to document departures therefrom. This system, it is hoped, will allow readers to relate information! from one chapter to another and one part to the next without the handicap of conflicting names for the same organism. Manuscripts first were solicited from contributors in 1973. Most had been received by the end of 1974, and Part I of the series was published in 1976. As editorial work progressed, some authors provided up-dated information and all authors of chapters in Part II had the opportunity to insert limited materials at the time they received galley proofs early in 1977. Therefore, content is as current as reasonably could be anticipated for a project of this kind. Organiza¬ tion and editorial style follow that established for the Special Publications of The Museum at Texas Tech University. Otherwise, authors were allowed broad latitude concerning material to be included in their chapters. Accordingly, and for obvious other reasons, some chapters overlap others in content. Even though some redundacy has resulted, we thought it best to have a section on the cited literature with each contribution. Citations to manuscripts in Part II and those scheduled for Part III of this collected series are carried in text as “this volume.” For the convenience of readers who may not have seen Part I of the series (Spec. Publ. Mus., Texas Tech Univ., 10:1-218, 1976), the titles, authors, and pagination of its contents are as follows: Introduction (Baker, Jones, and Carter), p, 5; Annotated checklist, with keys to subfamilies and genera (Jones and Carter), 5 f) SJ^ECIAL F'UiilJCATJONS MUSEUM TEXAS TECH UNIVERSITY pp. 7-38; Zoogeography (Koupman), pp. 39-47; Chiropteran evolution (Smith), pp, 49-69; Collecting techniques (Tuttle), pp. 71-H8; Care in captivity (Green- hall), pp. 89-131; Economics and conservation (C. Jones), pp. 133-145; Brain anatomy (VIcDaniel), pp. 147-200; and Lactation and milk (Jenness and Stiidier), pp, 201-218. May 1977 Robert J, Baker J. Knox Jones, Jr. Dilford C. Carter ENDOPARASITES John E. Uhelaker, Robert D. Specian, and Donald W. Duszvnskj The leaf-nosed bats of the New World family Phyllostoniatidae occur from the southwestern United States through tropical Central and South America, Mem¬ bers of this family are also found throughout the Antilles. The ecological associa¬ tions of the species in this family seem to be rather broad; species are found in humid tropical to semiarid and arid subtropical environments. Diversity in feed¬ ing is readily apparent ranging from nectivores {Gioss(^phaga). frugivores {Antbeus), sanguivores {Desmodus), to omnivores (PhyKosionius) (see review by Glass, 1970; Gardner, this volume). To understand better the biology of phyllostomaiid bats, it is worthw'hile to examine their parasites. The distribution of parasites, especially endohelminths, is governed largely by climate, distribution of intermediate hosts, feeding habits of the hosts, evolutionary age, physiology, and availability of the host species. Be¬ cause parasites often evolve with their host, the systematic and phylogenetic ages of particular groups of hosts can be determined, in some cases, directly from the systemalics and assemblages of their parasites if appropriate precautions are taken. The aims of this study w'ere to collect and correlate as much information as possible concerning the endoparasites of the Phyllostomaiidae and present prob¬ lems for future work. Specifically, this report includes a systematic review of ail parasitic species of Protozoa, Acanthocephala, Pentastomida, Platyhelminthes, and Nematoda wcurring in the Phyllostoniatidae; an addition of unpublished parasite collection records; and a preliminary appraisal of various factors that have influenced the dispersal and special ion in the endoparasites of leaf-nosed bats. Historical Review Published works dealing w ith parasites of leaf-nosed bats are few. The earliest studies were probably those of Kolenali (1856) w'ho examined bats in Brazil and described several nematodes of the genus Capillaria Zeder, 1800, Molin (1861) described and reported on the anatomy of Histiosirofigylus coronams from Phyl- iiKstoftta sp. (not necessarily a species of Phyfhsiomus) collected in Brazil. Fol¬ lowing these early reports of nematodes, Braun (1900) described several trema- todes from Brazilian bats. Looss (1907) indicated, however, that Braun’s descrip¬ tions were so inadequate that the species could not be identified. The tremalodes of Brazilian bats were studied later in good detail by Travassos (1921, 1928, 1955). Beginning in the I930’s, Perez-Vigueras initiated research on helminths of phyllostomatids collected in Cuba (1934, 1935, 1936, 1941a, 1941/), 1942). At about the same time, the nematodes of tropical American bats were studied ex- 7 8 SPECJAl. PUHLICATU)NS MUSEUM f EXAS TECH UN I VERS I (Y tensively by Lent and Tcixcra de Freitas (1936, 1940) and Lent e/ al. (1945, 1946). The first reports of helminths from North American phyllosiomatids were by Caballero y Caballero (1942). His contributions to the helminth fauna of Mexi¬ can bats continued until recently. In 1960, he and Grocott reported on helminths in bats from Central American countries. There are many reports of parasitic worms from tropical bats, I he majority of these reports deal with descriptions of individual species and are presented in the systematic part of this report. In addition to the above mentioned reports, several brief surveys arc available, namely, Chitwood (1938) and Slunkard (1938) in Yucatan, Mexico, and Silva Taboada (1965) and Barns and del Valle (1967) in Cuba. With the exception of the haemotlagellates, the protozoan parasites of bats have not been studied well. Most published parasite surveys of phylloslomatid bats are concerned only with their parasitic helminths, as noted above, or with zoonotic bacterial, viral, and fungal organisms (for example, Grose and Marinkelle, 1966, 1968; Grose c/ a(., 1968; Marinkellc and Grose, 1966). In only a few' instances have general survey reports included information of the protozoan parasites of phyllosiomatids and these are usually of a public health nature in '^.vhich attention is given to zoonotic forms. Several reviews of parasites from bats in general are available. Stiles and Nolan (1931) listed all known parasites of bats, including eclo and endoparasitic forms. A general account of parasites of bats w as presented by Allen {1939). Caballero y Caballero and Grocott (1960) published a significant w'ork review ing the ircma- todes from bats, Ubelaker (1970) published a general account of parasites from bats and in the following year, Barus and Rysavy (1971) analyzed the biogeng- raphy of nematodes of the family Trichostrongylidae occurring in mienKhirop- lera. Webster (1973) reviewed the helminths of bats north of the United Stales- Mcxico border. Methods The majority of the specimens obtained for study w'ere acquired by three col¬ lecting trips to Southern Mexico and Central America. Collectors on these trips included Cesar Estrada R. (CER), l.awrcncc M. Hardy (LMH), J. Knox Jones, Jr. (JKJ), rimothy E. Lawlor (TEL), James D. Smith (JDS), Delbert L. Kilgore, Jr. (DLK) and John E. Ubelaker (JEU). Specimens indicated by DWD were col¬ lected by Donald W'. Duszynski in Costa Rica. Specimens collected in Mexico or Nicaragua w^erc fixed in formalin or acetic acid-formalin-alcohol (AFA) and stored in 70 per cent ethanol; those collected in Costa Rica w'ere fixed in warm 70 per cent ethanol and stored in 70 per cent ethanol and 5 per cent glycerine until studied. Wherever possible, museum accession numbers are given for host specimens. The designation (KU) refers to the mammalogy collection, the Museum of Natu¬ ral History, The University of Kansas, l^awTcnce. Due to the misidcniification or name changes of hosts, the practice of depositing hosts in reputable museum col¬ lections is strongly encouraged. BlOl.OGY OF THE PHYLLOS]OMATIDAE 9 [Editors' note; Because the use of host names in the older parasitological liter¬ ature often obscures host-parasite relations for those ill acquainted with the nomenclatural history of host taxa, we routinely replaced a junior synonym w'ith a senior one. When some notation of such changes seemed necessary, we enclosed a brief explanation in brackets; otherv^ise, none was made. Also, mis¬ spelled names were corrected. We made no attempt to verify the identification of any species, although a notation w'as inserted when the identity of a host w'as im¬ probable. A host name was enclosed in quotation marks to indicate that its origi¬ nal use in the parasitological literature could not be applied with certainty to any known taxon. ] All specimens to be studied by light microscopy were stored in 70 per cent ethanol and subsequently mounted on glass microscope slides. Soft-bodied speci¬ mens were stained in acetocarmine, cleared in xylene, and mounted in Canadian balsam prior to study. Nematode specimens were cleared either in warmed lacto- phenol or glycerine prior to study. Specimens studied by .scanning electron microscopy w-ere prepared in the fol¬ lowing manner. Fixed specimens were dehydrated in an ascending series of ethanol solutions to 70 per cent, transferred to 5 per cent glycerine-95 per cent ethanol solution from which the alcohol w'as allow'ed to evaporate, and cleared in 96.6 per cent glycerol-0.05 per cent potassium chtoride-3.35 per cent distilled water, 24 to 48 hours prior to examination. Whole specimens or dissected por¬ tions of the helminths were mounted on metal specimen stubs with Duco cement, out-gassed in a vacuum evaporator for one hour or more, rotary coated w ith gold palladium (200 A or less), and examined w-ith an AMR 1000 scanning electron microscope. Phylum Protozoa The best present classification of the Protozoa is that proposed by Honigberg et ai (1964), as presented by Levine (1973), though we prefer not to use the lat¬ ter’s *’unttbrm endings of higher taxa” (Levine, 1958). Of the five subphyla uti¬ lized in this classification, two of these, Ciliophora and Sarcomastigophora, con¬ tain both free-living and parasitic forms, whereas in the remaining three, Apicom- plexa, Microspora, and Myxospora, all species are parasitic. Only two of these subphyla (Apicomplexa, the coccidia, malaria, and toxoplasma-type organisms; Sarcomastigophora, the flagellates and amoebae) contain parasites frequently found in mammals. Unfortunately, there is a considerable paucity of information on the protozoan parasites of all bats, w'orldwide, and such studies would provide much new' information to future workers. Subphylum Apicompi-Exa Levine, 1970 Class Sporozoa Leukart, 1879 Family Etmeriidac Eimcria sp. Type host .—Any phyllostomatid bat. 10 SPECIAI, PUHIJC Al J<)NS MUSEUM TEXAS TECH UNIVERSITY Siie of infeciiati. —Endogenous stages itsuaJly jn the intestinal epithelial cells; oocysts are found in the feces. Retnarks. —Although there are no records of Coccidia from phyllostomatid bats, we include this section to point out the immediate need for work in this area. Inasmuch as the Coccidia tend to be particularly host specific, the information from such studies could provide data to indicate and help us understand certain phylogenetic relationships. There are 13 named species of but eimerians, but it is questionable whether all should be considered valid species iPellerdy, 1974; Wheat, 1975), Of these 13 species, only Eimeriu enniops from Ettoiops fruoihiilii {Colombia), E. macyi from Pipistrelliis suhflavus (Alabama), and £. fhy/uinmycii'rklis from Rhyn- chonyaeris fiaso (British Honduras) have been reported in the Western Hemi¬ sphere (Lainson, 1968; Marinkelle, I968fv; Wheal, 1975). Presumably, eimerians and related taxa (for example, Kiossui variahiiis, see Levine ei ai, 1955) have not been found in phyllostomaiids because no one has bothered to look for them. The 13 reported species of bat eimerians are only a fraction of the number which must actually parasitize these mammals; Eimcria spp. have been described from only 12 of the 168 Recent genera (7 per cent) and 14 of the 853 living species (1.6 per cent) of bats recognized by Vaughan (1972). Although some species of Eimeria occur in more than one host, we also know that many hosts harbor two or more species that may be unique to them. If we conservatively assume that there is a least one Eimeria species per bat species, as was done for rodents (Levine and Ivens, 1965), w'C can estimate that there may be about 900 species of Eimeria alone in bats. The number described already is only 1.5 per cent of this number. Family Plasmodiidae PotyehromophiJus deariei Garnham tT a}., I 97 I Type hos!.—Myotis tiigricans. Site of inf eel iotL —Red blood cells. Type locality. —Para, BraziI. Other record —This species was seen in the blood of Glossophapa soricina from Para, Brazil, by Deane and Deane (1961), but their identification w'as both incorrect and incomplete (Garnham et aL, 1971; Garnham, 1973). Remarks .—^Haemosporidian parasites of any sort are rare in New World mam¬ mals. According to Garnham (1973), the haemosporidian parasites of bats fall into at least four genera, Plasmodium, Hepatocysiis, Nycteria, and Polychromo- p}}i!us^ with the first three being found only in bats of the Old World. The first re¬ port of a bat ‘‘malaria'’ on the American continent w as by Wood (1952) in w'hich he found W'hat he called Plasmodium sp. in five /I(Vesper til ion- idae) in California and in one A. pallidus and one Pipistrelius hesperus (Ves- pertilionidae) from the Chisos Mountains in Texas. He did not specify whether the California and Texas parasites were the same or different species. Only one report exists of a haemosporidian in phyllostomatid bats, and that W'as by Deane and Deane (1961), who found w'hat they also described as P!as- BIOLOGY OF THE PHYLLOSTOMATIDAE 1 I niodium sp. After describing and picturing the parasite in considerable detail, they concluded their paper by stating they weren't sure whether the forms they saw belonged to the genus Plasmodium or to some other genus within the '‘Haemoproteidae.” Garnham et ai. (1971) described A deanei from M. ui^^ricans (Vespertilionidae) caught in the same general area of Para as the bats examined by Deane and Deane (1961) and speculated that the general morphological fea¬ tures of P. deimei and the Plasmodium sp. seen by the Deanes were quite similar. In a later report, Garnham (1973) synonymized P. deanei and the form seen a decade earlier by Deane and Deane (1961) and, after reviewing the original slides made by Wood (1952), also placed that “malarial parasite” into the genus Poly- chromophiius. Thus, Polychromophilus has been reported three times in the New World, twice from the Amazon region and once from California and Texas, The latter parasite is longer and more oval than P, deanei and the pigment in the fe¬ male is more abundant. Family Toxoplasmatidae Toxoplasma gondii Nicolle and Manceaux, 1908 Type hosi.—Cienodactylus gondi. Site of infection, —Trophozoites and cysts throughout the host’s tissues. Type locality, —Foothills and mountains, Southern Tunisia, North Africa, Other tecordsr —Roever-Bonnet et uL (1969), using the Sabin-Fcldman dye test for toxoplasmosis, found the sera of two Artibeus litenuus from Tibu, Santander, Colombia to be positive for this parasite. Remarks. —Literally thousands of records of T. gondii from over 50 vertebrate species have appeared in the literature since this parasite first was described (for review, see Frenkel, 1973). How-ever, information on the incidence of 7', gondii in bats is meager as few- such surveys have been conducted worldwide {for example, Rifaat et ai, 1967; Kaliakin, 1970) and we find only one report documenting, serologically, the incidence of T. gondii in phyllostomatid hosts (Roever-Bonnet et ai, 1969). Toxoplasma gondii is almost ubiquitous in nature and the role of bats in the ecology and distribution of this most important para¬ site certainly should merit immediate future investigation. Subphylum Sarcomastigophora Honigberg and Balamuih, 1963 Class Zoom ASTI GOP MORE A Family Trypanosomatidae Before beginning a discussion on the haemoflagellales, wc must point out that the classification of the various species and the terminology associated with their developmental stages has changed considerably in the last several years. Thus, to be consistent with current trends of thought, w'e will follow the classification of the Trypanosomatidae as outlined by Levine (1973) and the uniform terminology of body forms introduced by Hoare and Wallace (1966). The study of trypanosomes of bats is important because bats often live in proximity to humans and can migrate great distances; thus, they can act as links J2 Si'ECIAL PUHLJCATKINS MUSEUM TEXAS [ECH UNIVERSlfY between sylvatic, rural, and urban populations. According to DiasU936t;), try¬ panosomes of bats have been known since 1898 when Dionisi in Italy first iso¬ lated and described, but did not name, haemoflagelkites that he found in the blood of three species of vespertilionid bats (Miniopiertts schreihersii^ Vespeniiio murinits, ye.spenty(> lUKfida). Dias (1936(0 also stated that in 1900 Durham examined the stomach contents of a mosquito that had just fed on the bkwd of Pliyllo.sfomns sp. from the state of Para, Brazil, and found numerous trypr indUnti’it hy iifi iisfcrL'ik, BlK hosts l ocality Rffttrcncfs Ariihi’tr'i chieri’im, Ari ihei/s Jn/fittii e/ix/.s Ccn o/Zui pi’r.\pin7/^t/u, Ckt/erant'sct/x ffitftor forn/)i/ftX D( .\/rrodn ,v nr/ir/idus Artiheas lUtifnUts, Phytiostonsas hustnsnx CandUa pi‘ rxp kit lam, Choero >i isci rv /irhntr. GiifSSt rph (lift I x< trie it ui Ca n dfki p cr.sp ic illar a, Gfirssopha^i*t! soricitia CaroflUt pcrsph illaiti, " Lftuchoplosxa etwitiiitir Dcsaiodax roititsdtis Di'xaiodiix romtuitis A ri ih vtix jtttna iev f \ s i\, Urodvnita hilohafuni Art the us I it taw us, Candikr perspiciUafa, Dt’S!funitis roiundus, Gliixsopha}>a sttrk rtia, PItyKoslofiius disri}l<)i\ Phylf(isrpjinc>.soiiii:i ptssoai Jiirco, San Jose, Cosia Rica Para, Brazil Guararema, Sao Paulo, Brazil Cali, Colombia Trjpiinosoina pifanoi Tibu and Tolima, CoJombia rrj piiiio.soiiia spp. Para, Brazil Para, Brazil Rio tie Janeiro, Brazil San Jose, Costa Rica Para, Brazil VFSPERTIl lONlS Trypunosiania erti/i Canal Zone, Panama Western and central Colombia Chilibrillo Caves, Panama Bella Visla, Panama Brazil Trypanosoma cruzi-Hke Western and central Colombia Esquival ct ah, 1967 Deane, 1964/) Deane and Sugay, 1963 Ayala and Wells, 1974 Marinkelle and Duarte, 1968 Deane, 19646 Dias tv td„ 1942; Deane and Sugay, 1963 Dias. 1940 (not T. heyhi'if'i-lik^, see Deane, 19646) Zeledon and Vieio, 1957 Romana, 1940 (ia Dias tV ul., 1942) Clark and Dunn, 1932 Marinkclle, 19666; Marinkelle and Grose, 1966 Clark and Dunn, !932 Clark and Dunn. 1932 Dias, 19360 Marinkelle, 19666, 19686; Deane. 1967 14 SPECIAL PUHl.JCAllONS MUSEUM TEXAS lECH UNIVERSITY Cf^ n^ilia p (■ np k il!ai a , [}v:!iftuHius nHiifiiitis. Glits^ophtipii .\ork imu Miinofi hcftnctlii, Fhy(lt>.\toiiJu.'i discolor, PhyUostttnitis hti.sfdi us, Unulenmi hift/huiuin, yuinpyimn spa irnni A ri ihcusjttiiutici’ijsis, Pinilostontifs httsfafiis Ciinfllki per spies! I tiiu CiirtfUid perspk iHiSfa Carol!its pcrspictlhtfa, Cliiii’rtsfu'-'ii'its ttliuor, Gitfsss/ph(spa Stfrsc!ssa, /.ofsc!)t>p!jy!!(S /aordisx, 1/ icnfssych'ris sssepaItP Is, F/syUos/iossfss cltf/spa/sis Ctif o!/i(f persp!(iHasss, C/soiTttfsiscsss tssifsos', /’!iy//(3sff)ssis{s iussfarsss Dess/sodtss s(>risfi(/sss G1 1 ' v.vi )p h apa st o- k i/tu, Lossc/iop/iy!!(s sitofssas! P/syl!i3S/(}stjsn dofspfs/sss PlsyUosu/sasis isii.'s/dfS/s Vtsss/pyri/ps /i/s asftss CaroHics persp i( if!(tia Cort//!!o perspic!/!(s//s Cssr/fUia persp!ci!!is//t Table 1 .— Ct/stisssssc/!. French Guiana Cojonibia Guarativana, Yaracuy, VeneTiuela Marajo Island. Iklem and Para. Brazil Para, Brazil Panama Para. Brazil Venezuela Brazil. Colombia, Venezuela Try panosoma lineatiis Caracas, Venezuela Trypaiiosoina pliiyliosfoinae Brazil Cuba Guaraiivana, Yaracuy. Venezuela Fioch cs ts!., 1942 Renjifo-Salcedo ei a/„ 19521 Marinkelle, 1966/? Dias and Pifano. 1941 Dias e; ts!,, 1942; Deane. 1961, I964u Deane. 1964//, I964^: Dias, 1940 Wood and Wood. 1941 Gam ham e/ a/., 1971 Dias and Pifano, 1942 Carini, 1932; Deane, 1961. 1964r/, 1967; Dias. 1933, 1936u. 19.36/); Dias and Romaha. 1939; Marinkelle, 1966/?; Pifano, 1964; Renjifo- Salcedo, 1948; Renjifo- Salcedoe/u/., 19.50 Itiirbe and Gonzalez, 1916; W.Y., 1917 Dias. 1940 ( = 7. ers/zi- like?, see Deane. 1964/)) Carl ay a, 1910 (= 7. ers/z!- like?, see Marinkelle, 1968/)) Dias and Pifano, 1941 (= 7. cr/szi-like?, see Marinkelle, 1968/?) tilOUOGY OF THE PHVLLOS'TOMATIDAE 15 Citml!id pk'ilidfiU “ Loiu'ho^IdJSSii evdiicldhr Cardllui persp k ilUiia, Chi?i'n>ttixi ds ni inory Gl{tssitpiiagd xorit ina, “Loiichoglttsxd cadiikitd" Cdfoii id persp it il fd tn Gtoxxophdgd soriciiidy Fhylf<>x(t)m ux futxtatux Mdcroldx H'dti't /iousii Phyll<}xt(}njus fmsldfns Cdroilid pL'i spk il(aui Loncl it }phy f I a fn o nl ax " Tm chops ctongdtdx" Arfihi’ds jdf}jdiccits{s*y Carol!id persp icifhdd *, GlossophdMd soricina *. Phyllosiodttis /wxfdj us* Des rn o ih tx rot ti nil us* DesmoJux ro/tiittiiis Di'suioiliis rotund!!S Dcsuuniux rotu/uiux* Artihcux (itiirdfusy O!osxtiphugd soric inti Tabl.e T — Ctiutiitited. T ry |iaiM).s«iiia vespert it ion is Rio lie Janeiro. Brazil Brazil San Jose, Cosia Rica Coquimatlan, Colima, Mexico Colombia Try panosoma spp. Limon, Costa Rica Brazil Brazil OTHER SPECIES Trypanosoma evaiisi Panama Panama Arauca and Cali, Colombia Valle de Cauca, Colombia Trypanosoma equinum Argentina Try panosoma raiigeli-Hke Central and western Colombia Dias. 1940 Dias (T di, 1942 (see Deane, I964i^) Zeiedon and Victo, 1957, 195H Mazzotti, 1946 See Marinkelle, 19666 Zeiedon and Rosabal, 19696 Romaha, 1940 (in Dias and Pifano, 1941) Dias and Pifano, 1942 Clark and Dunn, 1933 Dunn, 1932; Clark and Dunn, 1933; Johnson, 1936^;, 19366 Ayala and Wells. 1974 Ayala. 1972 {in Ayala and Wells, 1974) Acosta and Romaha, 1938; Hoarc, 1965 Marinkelle, 19666; Tamsitt and Valdivieso, 1970 "not Trypanosoma cruzMike" Choeronixciix minor Para, Brazil Garnham e/fd., 197! 16 SJ'KCIAI (C ATIONS MUSEUM IFXAS TECH UNlVERSJfY Trypanosoma ( = Sciiizotrypniium) cru/i Chagas, 1909 Type }u>sL — Pims!rimi>yhis /,v/ a. Site of infevtion.^in the inlesltne of tlie triatoniid bug (originally)* hut also intercellularly in the blood (trypomasiigoie form) and intracellularly in the reticuloendothelial and other tissue cells (amastigote form) of vertebrate hosts. Type locality. —B raz i 1. Other records, —Sec Table 1. Reomrks, —Trypanosomes morphologically similar to T. cruzl have been re^ corded from more than 100 species of mammals (Deane* 1964«)- Technically, forms identified as this species should be restricted to those which produce amastigote bodies in the organs of inoculated laboratory animals or in tissue cul¬ tures. In addition, the length of the trypomastigote blood form (approximately 20 micron.s), its nuclear index (approximately 1,4 to 1.6), its ability to develop in triatomid bugs, and w'hether or not the bat host(s) came from endemic areas of Chagas' disease should all be utilized as supportive evidence in such identifica¬ tions (Deane, 1961), Only three reports ( l able 1) use much of the above criteria to demonstrate conclusively the presence of 7’ cruzl, either naturally or experi¬ mentally, in American leaf-nosed hats. Trypanosoma cni/i-like Retmirks, —Many of the bats in most of the countries of the Americas are hosts to trypanosomes structurally identical to T, cruzl (Sec Marinkelle, 1965). It is now generally accepted that these forms should be referred to as T, cruzi-Vike when only blood forms are studied or if they fail to produce amastigote bodies in living cells (Marinkelle, 1966 /j). However, Marinkelle (!968/>) stated also that the majority of T, cn/c/-Hke forms {vcspcrilllotiis group) are capable of forming amastigotes in cells of mammals. Deane (1964u), on the other hand, disagreed with this view and summarized well the difficulties encountered in working with bat trypanosomes: “The bat strains, how-ever, remain a problem. At least some bats of the endemic area of Chagas’ disease do harbour flagellates undistinguish- able from 7’. cruzl, on the basis of morphology, biology and virulence and even immunologically. But most bats harbour strains which cannot, at present, be identified to the agent of Chagas' disease: they are of little or no virulence for laboratory animals and, besides, some strains do not seem to develop well in triatomid bugs and others show morphological differences that are said to be con¬ stant.” Dias (1936fd offered somewhat of a compromise position by suggesting the trypanosomes of bats can, after repeated passage, change their virulence and lose the ability to infect other hosts. A translation of his original statements (p. 75, in Portuguese) follows: “One extremely interesting question that should be belter investigated is that of the behavior of virulent /'. cruzl in bats that are natural hosts to trypanosomes. Experiments done to date show' that these mam¬ mals (at least some species) arc %'ery resistant, if not refractory, to infection by strains that are very pathogenic to other animals. One of our experiments demon¬ strated that before the trypanosomes are destroyed they experience an abrupt BlOl.OGV OK THE PHYLLOSTOMATJDAE 17 and remarkable attenuation of virulence in bats. If, by means of repeated pas- sages, one succeeds in obtaining infections that are more and more prolonged, finally adapting the trypanosome to the bat, it is possible that this adaptation will be made at the cost of the loss of infectiveness to other animals, because of a real effect which the organic environment of the mammal exercises on the nagcllaie. If this could be verified, 7". cmzi will have been transformed into T. vcsperfliiotiis, just as T. ve.sperftiionis can he identified as T. cnizi in those rare circumstances in which its inoculations into animals arc positive." Additional confusion in naming such forms stems from: the highly variable na¬ ture of structural dimensions during different phases of infection by a single strain of cruzi‘, the w'ide variation in nuclear indices reported for T. critzi (from 0.95-1.63 by Bareito, 1965); and the possible influence of temperature on the morphology and pathology of various trypanosomes (Marinkellc, 1966^/, 1968/>). Such information points out the need for much additional work before the T. rHi^z/'-like forms in bats can begin to be accurately separated. Some of the first reports of T. rncZ-like parasites from bats in Latin America were by Dias and Pifano (1941, 1942) in Venezuela. However, Zeledon and Vieto (1958), based on their biometrical study of tw'o lab strains of T, cmzi (from mice and trialomids) and of 7’. vespenilkmis isolated from a Glassitphaga Sifficina caught near San Jose, Costa Rica, considered the forms seen by Dias and Pifano {1942) to be different from T. cmzi and 7'. vespenilionis. Zeledon and Vieio (1958) and later Marinkelle (1968/)), in a retrospective look at the literature, considered as 7’. c‘ni;/-like the following phyllostomatid bat trypano¬ somes: those from Curollkt persptcilkifu and described in Cuba by Cartaya (1910) as 7'. phyllostomac., the “phyllostomae" strain from Camilia pcrspicillam in Venezuela by Dias and Pifano (1941); the Brazilian strains from C. perspicklata and Phyllo.sionuis hashtnfs studied by Deane (I964«); and the strains isolated from 11 species of phyllostomatids in Colombia (see Table 1) by Marinkelle (1966/), 1968/)). Additional records to 7. cT//z/-like forms found in American phyllostomatids are listed in Table 1. Trypanosoma equinutii Voges, 1901 Type fum. — Horse.^. Site of infectiotL —Extracellular blood parasite. lypc iocality. "It originates in South America and occurs as far south as the Argentina provinces of St. Fe and Corrientes" (sec Voges, 1901). Other m<)/v/.v.—See Table 1, Remarks .—This species differs structurally from 7. evnnsi, from which it probably arose, only in lacking a kinctoplast (Levine, 1973). Tryparnmwia etfifinimi infects cattle in an asymptomatic form, but produces a severe disease in horses called Mai de Caderas throughout much of South America, especially Brazil. It is unique (as is 7\ evansi) in that it has evolved to utilize the vampire bat, Des/nffiitis rotumius, as a parallel host and as a vector of the disease (Hoarc, 1965). In Argentina, it w'as demonstrated experimentally that vampire bats be- ]H Sf'ECIAL I'UBLJCATJON.S MUSEUM l EXAS I Et H UNJVFRSJ T Y come infected with iyuifUffn from horses and can iraiisinit it by feeding on healthy horses (Acosta and Komaha, I93X). Trypaiiusonia evaiisi (Steel, 1885) 7>7j c ya J -v/.—“ H o rso s," Sife of (tifeaioti .—Extracellular blood parasite. /y/;c localify,- —Punjab, India. Other rei itnis .—See Table 1. Rcfitarks .— 7ryptnuKSofua evansi ( = T. hippiciufi) has a wide distribution in Latin America being prevalent in Mexico, all of Central America, Venezuela, and Colombia, where it causes a disease called Murrina in horses (Hoarc, 1965), Hoarc (195?) stated that mechanical transmission of T. evansi (and of 7'. eipiinnin) probably evolved as a secondary adaptation when it separated from its African ancestor T. hrncei and lost its original intermediate host, the tsetse fly. After these two species became established in the New' World, they acquired, in addition to blood sucking flies (Tabaniidae), a new type of vector, the vampire bat. Vampires are ideal vectors because their infection from cattle harboring small numbers of parasites is ensured by the large amount of blood taken during a meal (16 to 50 milliliters) (Hoarc, 1965). The high rate of reproduction of the parasite within the vampire's body increases the chances of successful trans¬ mission to new hosts, I'hcrefore, vampires play an important role in the spread of bovine Murrina among horses in laitin America. Dunn (1932) first documented that the vampire bat Desnioilus rotundus w'as a natural vector of T. evansi on the Isthmus of Panama, and Clark and Dunn (1933) were able to transmit this trypanosome to other phyllostomatids (Table 1), but all specimens so infected, including the vampires, were highly susceptible to disease and died within a few weeks. Clark and Dunn apparently never found any phyllostomatids with ’'spontaneous" ( = natural?) /’. evansi infections, but felt that the vampire bat, inasmuch as it could be infected experimentally and fed with equal freedom on equine and bovine animals, might be an important vector in transmitting this parasite from reservoir cattle hosts to highly suscep¬ tible horses and mules. Johnson (I936if, 1936b) and Hoare (1957) also demon¬ strated that vampire bats acquire and transmit 7’. evansi under experimental con¬ ditions, but we found records of only 20 individual vampire bats with natural infections (Ayala and Wells, 1974; Clark, 1948; Johnson, I936«, l936/>). Trypanoscniui tineatus Iturbe and Gonzalez, 1916 I'ype ho sL — Vatnpyrops iineatns. Site of infection .-—Extracellular blood parasite. Typ e IfH'a I ity. —Venezuela, Other records ,-—None to date. Remarks ,—Since this species was originally described, it has been mentioned on only three occasions in the literature. The first W'us a rather scathing review- by one of the editors of Tropical Disease Bulletin (W. Y., 1917) and the other two limes (Zeledon and Vieto, 1958; Marinkelle, 19686) the authors c<)nsidered BIOLOGY OF THE PHYLLOStOMATIDAE 19 this form too 7, to merit its own specific status. The validity of this species is, therefore, questionable. ( Vanipyraps litieaiiLs is not known to occur in Venezuela, and the identification of the host is probably erroneous. Eds.] Trypanosmiii) pessoai Deane and Sugay, 1963 In Venezueia, Dias and Pifano (1941) isolated a njcfiadt’ntiac-lypG trypano- some (from Myofis tuf^rkwis) for the first lime in the New- World as these forms were previously knowm only from bats in Africa. Since then, several large un¬ named li 7 panosomes of the /negadcniuie group have been reported from phyl- lostomalids in the Western Hemisphere (Table I), but only Deane and Sugay (1963), Esquival ei al. (1967), and Marinkelle and Duarte (1968) described and pictured these parasites. Since its original description, this species has been reported in several species of phylloslomatids (Deane, 1964u; Esquival ei u/., 1967), Trypanoscfnia pessoal differs from the vespc^nilicinis group (particularly 7. cmzi) not only in size, but also because xenodiagnosis, hemacultures, labora¬ tory animals, and tissue sections and smears arc always negative for other de¬ velopmental stages (for example, aniastigote forms) of the parasite. Trypanosoma phyUostomae Cartaya, 1910 Type hosi.—Cawllia perspicHlafa, Site of infecfkm .—Extracellular blood parasi tes. 7'ype local tty r —Cuba Other records ,—^See Table I. Ret narks .—Most of those who work w-ilh bat trypanosomes believe this species to be too 7. c/v^cZ-like to distinguish it as a separate species (see Table 1). Trypanosoma pifanoi Markinelle and Duarte, 1968 Type hosts.—Artibeifs litiinmts and Phyflostotnas hastatns. She of infection .—Extracellular blood parasite. Type ItH alities .—Tibii and Tolima, Colombia. Other records .—None to date. Rctnarks .—This is only the second species of the tnegadertmie group to he found in the Americas. Like frypanosotna pessoai, developmental stages of this species could not be isolated in tissue sections of inoculated laboratory mice nor was multiplication observed in tissue cultures of mouse fibroblast cells or in the triatomid Rhodniiis proUxas by xenodiagnosis (Marinkelle and Duarte, 1968). Attempted transmission of this species to Carollia perspicillata w'as unsuccessful, but blood forms isolated front a specimen of Artiheas fitiiratus w'ere grown in NNN culture media and these culture forms closely resembled the blond and cul¬ ture forms of Trypanosoma ernzi. Also, when 5000 NNN culture forms were in¬ oculated intracoelomically into three species of triatomids, the parasite (when compared with control 7. cruz/-inoculated bugs) proved highly fatal for the insects. Only three of 264 triatomids so inoculated lived for four weeks postinoc- iilation (PI) and at 30 days PI their hemolymph had numerous, long, slender. 20 SOFX IAI PlJti] it A1 IONS MIJSFUM i HXAS TFCH LJNIVFRSJiY Trypufinsomu cpinrastigote tbrjiis (Mariiikellc and Duarle, 1968). This species difl’ers from T. pessimi in size and by the absence of a twist of the posterior of the body. Trypaiiosoinii rangdi-like Remarks .-—Only Marinkclle (1966/)) has reported what he called like trypomasiigotc fortiis from American phyllostomalids. lie found three hats (Table 1) harboring such parasites, and xenodiagnosis with the triatomids R. pt{>li\i(s and Citvemicoki piUisa showed abundant development of epimastigote stages of this parasite in the rectal ampulla of the bugs. Neither anterior station development nor signs of homolymph infection took place and attempts to infect laboratory mice with these forms were unsuccessful. TrypanosuJiia vesperfilicmis Battaglia, 1904 Type host.—FipispvUtts sp. Site of iafeahih —Extracellular blood parasite. lype iovaiity,- —Italy. Other records. —Sec I'able I. Remarks, —Since the original description of this parasite from vespertiHoiiid bats in Europe, it has been observed on several i)ccasions in bats of the Americas (for example, Deane, 1961), but few reports exist of its occurrence in phyl- lostomatids (1 able 1). This species can easily be distinguished from others with¬ in the vespertiUonis group by its small size (14 to 16 microns), its large nuclear index (2.6 to 2.7), and its apparent inability to infect laboratory animals or tri¬ atom id bugs. Trypanosoma spp. Remarks, —Unidentified forms of trypanosomes have been found in phyb lostomatids on many occasions. In the majority of these records, the organisms seen were reported to belong to the me^adermae group, but no illustrations of the parasite or structural data were provided (Table 1). Phylum Af AN JH(K IiPHAl A F a m i 1 y 01 i gac a n i h o rh y n e h i da e INeunek'oJa novcJJat'(Parona, 1890) Type iiosf.—A rtiheus jamaiceasis. Site Small intestine. Type locaTtiy. —Puerto Rico. Other revords. —^Nonc to date. Remarks .—-The acanthoccphalan fauna of tropical American bats is restricted to a single species described from A. Jafttaiceasis coWcclidd in Puerto Rico. It has apparently not been recorded since its original description. Schmidt (I972f/) in¬ cluded seven species in the genus, all w'ith 30 proboscis hooks. These parasites B[OLOGV OK THE i>H Y[.l.OSTOMA‘rtDAE have been reported in CamK'Ora, Chiroptera, and ducks (?) in South America, Malaysia, USSR, Puerto Rico, and Africa, The life cycle of N. mn’ellae is unknown. In a related genus, PnKst/iefioniiis, species such as P. elegaiis and P. sptmla are reported to use cockroaches {Bkitef- la gcmuwica^ RhyparohLs /tiadarae, and Blabt^ra fusca) as well as beetles {Liisiodemta serriafnie and Sfegohiitni pti/iiccttm) as intermediate hosts. Pre¬ sumably similar insects serve as intermediate hosts for N. mn’cllae. If this is true, the host bat becomes infected by eating a cockroach or beetle containing an in¬ fective larva, the cystacanth. It should be emphasized that intermediate hosts listed above represent experiments based on captive animals; the intermediate hosts in nature are not known. Pathology due to acanthocephalans, in general, is influenced by nunterous factors including the size, shape, and armature of the proboscis, number of para¬ sites present, general health of the host prior to infection, and ability of the host to overcome secondary infection by pathogenic organisms (see Schmidt, \912h). Inasmuch as the effect of N. iioveliae is unknown in Artiheux, a general discus¬ sion of pathology, diagnosis, treatment, and control of related species is not in¬ cluded here (see Schmidt, 1972A). Phylum Pentastomida Family Poroccphalidae Porocephalus crotali (Humboldt, 1808) Type hoxi.—Crotulns diirissus. Site (^f i/ifec!toti .—Body cavity. Type —Unable to locate. Of her rcro/z/.v.—See below. Remarks .—Members of the phylum Penlastoniida, often referred to as tongue worms, are of uncertain systematic position, although evidence is accumulating that they are related to the brachiuran crustaceans. The genus PonHephalus is among the most highly evolved of the peniastonies. All species parasitize snakes as adults, and most may utilize a mammal in their development as does A crofuli, the only species recorded from bats (Self, 1969). Porocephalus crotali occurs as an adult in various species of snakes, and has been reported as nymphs encysted in the liver of Phyilosnmms discolor from Cumana, Venezuela, and BrEizil {sec Penn, 1942; Sambon, 1922; Shipley, 1898). The life cycle of P. croudi has been studied intensively by Esslinger (1962u, 1962/), 1962c). Adult bats probably can be infected by ingesting eggs that con¬ taminate food. From experiments with albino rats, it is known that the larvae hatch in the intestine and migrate through the w-all into the viscera and mesen- taries, leaving a trail of host neutrophils. After reaching the liver or other organs, they molt and eventually form sixth stage nymphs that show marked sexual differentiation. Development of the sixth stage nymph is completed in three months and it is then infective to the snake definitive host. Infection occurs by ingestion of the infected bat host, w'hich may be a more common occurrence than previously suspected (Gillette and Kimbrough, 1970). SE^KCIAI J^UHLK'ATIONS MUSEUM TEXAS I ECH UNIVERSITY Pathology of penlaslon'ics tt> iheir bat hosts probably is related directly to the development of the two pairs of hooks on the head. During metamorphosis to the sixth stage nymph, the adult hooks develop from papillae representing the atrophied appendages of the primary larvae, I’he median and lateral hooks project and have blades that extend above the surface of the head and serve lo anchor the nymph to the tissue. As seen in Fig. I, the lateral hooks project con- spicut)iisly from the surface and undoubtedly cause the primary destruction of host tissue. As the parasite develops in the liver, and in probable response to the hooks, a granulomatous lesion forms. At least four distinctive progressions of the disease can be determined: an initial macrophage proliferation with eosino¬ phils, epithelioid, and giant cells accumulating in the area of the lesion lasting about three weeks; clonic development with involvement of fibrobhustic tissue, plasma cells, and lymphocytes during the second and third months; reduction in inflammation during the fourth month; and production of a dense hyaline fibrous capsule by the sixth month. Again, it must be emphasized that the life cycle and pathology as determined by Esslingcr (1962^/, l962/i, I962r) did not em¬ ploy bats. PorcfcephalKs vrofali is also recorded in man (Stiles and Nolan, 1931). Phylum Plai viii^i MIN 1 nts Class Tkema ioi>A Family Anenterotrematidae Aiicjiterotreiiia aurifiim Stunkard, 1938 7'ypi’ has{. — Micr()nyv!Cf isftu\i;alo!is. Siic of infeefion .—Small intestine. I'ype locaihy .-—Cueva de .Xmahit Teka.x, Xconsacah, lizimin, Yucatan, Mexico. Anetilerotreiiia eduardoeahalkroi (Freitas. 1960) 7'ypc hosL — Eiinu^ps inns. Site of itt/eefion .—Small intestine. Type hHdlity .—Sao Paulo, Brazil. Other reconls .— I'ravassos ei aL, (1969) gave the following host records from Brazil: Moln.ssn.\ rnfiis, M. mujar erassicaiukans, and Phyliosumuis elongotns. Aiienten^trenia freitusi Caballero y Caballero, I 964 I'ype host.—Micronyeleris h irsttfa. Site of lufecilon .—^Small intestine. Type lovuiity .—^Cosia Rica. Aiunterotrema linputianuni (Travassos, 1928) / ype h ()si.—Perop te /'>m canina. Site of infecdofh —Small intestine. Type locality ,—Angra dos Reis, Brazil. BIOLOGY OF THE PHYLLOSTOMATtDAE 2:! Fi(i, L—Scanning eleciron photomicrograph of lateral hook on the head of Pot oce phut as vroiaU from the body cavhy of Epresicus fusi us collected at the Black Gap Wildlife Manage¬ ment Area, Brewster County, Texas. The above report is the first listing of P. notciti in North American hats. I X2l5) —Travossos p/ ij/. (1969) gave the following host records from [Brazil: ‘‘Molossidae sp,,” M. /nct/or fm,v.v/tY;trr/f//rrv, and P/jy/hs/amus e/o/iptuus, Teixera de Freitas and Dobbin (1963) also re¬ ported finding A. /t7ipu//afnp?i in Mo/rmtf.v |The name Ppropuryx can/mi could refer either to Peropferyx kappferi or P. manotis, Eds.J Anentcrotretua stunkardi Caballero y Caballero and GrocoiL I960 Type hasf.—Phyllostomus hasiams. Site of infea km. —Small intestine. / yp e he all fy. —Panama. Ranarks.—AW known species of Anenteroirema have been found in the small intestine of their hosts. Members of this genus are unique because, unlike most digenetic trematodes, they lack a digestive tract. This evolutionary structural modification most certainly restricts their habitat selection in modern day hosts. Yamaguli (1969) examined histologically the parenchymal cells of A. awitnm and later (1971) stated that the nuclei of these cells were involved in nutritional activity. No glandular-secretory cell types have ever been reported (Yamaguli, 1969). Although five of the six species of Ancnicrotretmi occur in phyUostornatid bats, they are not specific. Anenterotn'ma freitaxi and A. sfnnkardi arc both re¬ corded from a single host species and are known only from the original descrip- 24 SPECIAL PUHl JCA t IONS MUSEUM 7‘EXAS TECH UNIVERSITY lions. It is probable lhal additional collections will indicate a general lack of host specificity. The biology of this genus is completely unknown. Inasmuch as these ire- matodes are so unusual morphologically, additional studies are needed. Family Dicrocoeliidae .4lhesmia parktTi Perez-Vigueras, 1942 Type hissl.—A niheifs Jamai( i'nsis, Siie (if infection .—Small intestine. Type iocaiiiy. —Province Pinar del Rfo, Cuba. Renuirks .—The species is recorded only from the type host in the original de¬ scription, Teixera de Freitas (1962) considered this species conspecific with A. heterolecithodes (Braun, 1899) Looss, 1899, common in the bile duct of a variety of birds. The only other species in mammals, A. fhxi Goldbergcr and Crane, 191 I, occurs in primates. The ecology, pathology, and life cycle of A, parkeri are unknown. Paraiiietadelphis eonipacttis Travassos, 1955 Type iiost.—Miaonycteris hehni. Site of infection .-—Bile duct and bladder, I'ype focadty, —Cachimbo, Para, Brazil. Remarks. —This trematode has been reported only in the original description. Nothing is known of its biology. Family Lecilhodcndrium Lecithodendriiim prk'ei Pcrez-Vigiicras, 1940 Type iiosr.—A rtihetis Jof?iaii ensis . Site of infection .—Small intestine. Type localiiy. —Santa Marfa del Rosario, Habana Province, Cuba. Although the pathology and ecology are not known, Koga (1954) reported briefly on the life cycle of LiTithodendriinn (ayenifonne (Ogata, 1947). Virgulate cercariae develop in an aquatic snail, Semisnlcospira iibertina, and encyst in Stenop.syche grissipennis. Bats are infected by ingesting the meta- cercariae transmitted by the trichopteran second intermediate host. The genus Lecithoiiendrinm contains numerous species occurring in bats and chameleons. At least 19 species occur in bats but all species except L. pricei are found in bats from Eurasia. Liitiafiiluni aberrans Caballero y Caballero and Bravo Hollis, I 950 /ype ii(KSt.—Macrotns h ateHu>nsii. Site of infection. —1 ntestinc. Type locality .—^Cuicatlan, Oaxaca, Mexico. Other records. — Njcaragua: Phyllostonnts discoior (KU 97445) collected at Hacienda San Isidro, 10 km. S Chtnandega, 10 m. (TEL 480). H[OLOGY OF THE PHYLl,OSTOMAT[DAE 25 Liinatuluiii isthmicus Caballero y Caballero, 1964 Type host .— Mkronyctet is / 1 irsnfa. Site of inf —Small intestine. Type locality. —Costa Rica. Limatutum oklahoniense Macy, 1931 Type host .— Taiia/ Ula hrasilicnsis. She i)f infectiotL —Small intestine. Type locality. —Aetna, Kansas, and Freedom, Oklahoma. Other ret on/.s.— Mexico; Mat rotas waterhoasih Cuicatlan, Oaxaca; Natal ns tnexicamis. Acolman (Caballero y Caballero and Hravo Hollis, 1950); Para¬ guay: Myoiis tiigricans, Chaco, (Lent et ah, 1945); United Siatesof America: Myo t is y r isescet is, Ka tisas (U beI ak er, 1966). Retnarks.—Liniatnlnm aherra/is and C. isthmiens apparently are restricted to phyllostoniatid bats. Additional records are needed, however, before spec¬ ificity can be established. Seven species occur in the genus and ail except L. okahei (Koga, 1954) Yamaguti, 1958, occur in New World bats. The ecology of this genus is unknow n. Family Urolreniatidae Uroircma scabriduiti Braun, 1900 Type ht)St. —Mithtssns major crassicainlatns. Site of itifet'tion. —Small i ntesl i ne. Type locality.—Braz'] 1. Other hosts,—-Noctilio leporinns, N. labialis, Molossus ater, Promops centralis, Phyilostomus hastatus, Lasinrns intermedins, Myoiis nigricans, Phyl- lostornns sp.; also in numerous bats in North America as reviewfed by Webster (1973) and Caballero y Caballero (1960). Webster (1971) reported that Pieronotns macleayii and Tadarida hrasiHensis from Jamaica w'erc also hosts to this parasite. Remarks. —Caballero y Caballero (1942) review-ed the systematics of this genus and concluded that the following species are synonyms of U. scabridntn: U. lasinrense Alicata, 1932 (see also Chandler, 1938), U. minntnm Macy, 1933, and U. shillingeri Price, 1931. Keys to this complex of species w ere presented by Macy (1933). Caballero y Caballero (1942) further considered Urotrematnltmi Macy, 1933 synonymous with U. scuhridntn and Caballero y Caballero and Gro- cott, (I960) considered U. aeileni Baer, 1957, parasitic in Pipistrelins nanus Cote D’Ivoire as synonymous w'ith U. siabridnm. Inasmuch as body shape, a more posterior position of the ovary from the acetabulum, lobed testes, and vitellaria that begin posterior to the acetabulum are all specific characters, it is doubtful that Urotrematnlnm is distinct. It is more reasonable to consider this species as Urotrenm attennainm (Macy, 1933) Caballero y Caballero, 1942, and distinct from U. scabridntn. SrFClAl I’UHl.lC A t IONS MUSEUM l EXAS I EOH UNJVEKSITY Ifi Class Cf.STO[>A F a m i I y A n o p I ac e p !i a I i d a e OochoristiL-a immature Arandas Rego, 1963 7'yfic htfSl — (rkfSM >p{uif^a so/ k i> m. Site of infect —Small inlestinc. Typ e l< H-a Iity, —Brazil. Hetmirks. — (kH’horistica imnutftint was assigned originally to the genus Miithevotaeniii hy Arandas Rego (1963). However, Della Santa's (1956) synonomy of this genus with Oochorisiica appears valid (sec Flores-Barreota et u/., 1958, and Prudhoe and Manger, 1969). Other species of Oocitori.stica in bats include (). antrozoi Voge, 1954, from Amrozoit.s palikins in the United States, O, nyctophili Hickman, 1954, from Nyct<}phi!ioi gcoffroyi in Tasmania, and O. ke/ivoulae Prudhoe and Manger, 1969, from Kerivonlu sp, and Tylonycteris from Malaya, F am i 1 y 11 yme nol cp id idae Vaiiipirolepls doiigatus Arandas Rego, 1962 Type hosts and localities .— Glossophaga soricina, Rio de Janeiro, state of Guanabara; Fhylk)st<}tnits hast at as, Conceifao da Barra, state of Expi'rito Santo; Molk>ssas ater, Tingua e Sao Goncalo, state of Rio de Janeiro, Br^izil, Site of infection. —Small intestine. Other recofds.—Glossophaga soricina: Mexico: Chiapas, Ruinas de Palenque, 300 m. (KU 102308); Nicaragiia: 3 km, N Sabana Grande, 50 m. (KU 97589); Daraili, 5 km. N, 14 km, E Condega, 940 m. (KU 97533). Specimens of Artiheas Utnratus from Cali, Colombia (collected by M. E, Thomas) contained several cestodes of this species. The specimens differ slightly in some measurements and at the present time they are provisionally considered as Vampirolepis eknigatas. Specimens from this latter collection have been depos¬ ited in the United States National Museum Helminthological Collections, Belts- ville, Maryland. Remarks .— Vampirolepis elongaias belongs to the subfamily Hymenolepidinae Perrier, 1897, and represents one of 27 species in the subfamily recorded from bats. Five species of Vampitidepis are knowai from the Western Hemisphere: K chiropierophila Pcrez-Vigueras, 1941, in Moios.sas tropidorhynchas from Cuba; V. decipiens {Diesing, 1850) in Fteronotns rahigimmh and Eamops perotis from Brazil; V, chistensoni (Macy, 1931) in Myotis lacifagus and other bats in North America; V. gertschi (Macy, 1947) in Myanis califnnicas from North America and V. roadahashi {Mncy and Rausch, 1946) in various bats in North America. An excellent review of host records of Vampirolepis spp. from North American bats is available (Webster, 1973). Vampirolepis elongatas is most closely related to V. chiropterophila but differs principally in measurements of the rostelluni and eggs, Vampirolepis eiongatns is potentially dangerous to its hosts for the rostellum interrupts the integrity of the intestinal epithelium and may produce ulcerous conditions in infected animals. B[OLOGY OF THE PHYLLOSTOMATIDAE 27 Fici. 2.—Scanning electron photomicrograph of posterior proglottids of the sirobila of I'timpiroU’pix cloui^'aiifx from the small intestine of Oloxxopimuii soricifiu collected in Nicaragua. Note the decreasing dimensions of the proglottids and the opening of the ex¬ cretory system (at arrow). ( X 7001 Ihe surface of Vampirolepis efongiKtiS is clearly similar to other hymenolepidiid cestodes in that it is cellular and covered with a dense microvillar surface that presumably aids in the absorption of available nutrients from the host (see Ubelaker et uL, 1973). Examination of the strobila by scanning electron microscopy reveals that even the terminal proglottids (filled with eggs) are covered by a dense absorptive surface (Fig. 2). As groups of proglottid.s become gravid, they detach and are passed out with the feces. Although intermediate hosts 28 Sj>PCJAi- PUH[-[CAT[{)NS MUSPUM TEXAS TECH UNtYERStlY of other hymenolepidiid cestodes involve various insects, the life cycles of all Vampirok'pis are unknown. Kochseder (1969) suggested that Hymenoleph grisea (van Beneden, 1873) had a higher incidence in younger animals {Myotis myotis, M, Riiinolopfiiis fernouequinunu and Barbasrtkla harhasrelliis) than older ones. Phylum Nematoda F amily Dipctaloneniatidae Litoinosoides artibei Esslinger, 1973 7'v/;c host-—Artiheus cif wrens. Site of infection^ l horiicic or abdominal cavity, lype locafity,^ —Vicinity of Buena Ventura, Valle, Colombia. Litomosoides brasiJiensis Linsde Almeida, 1936 Type h(fst.—Myotis sp. Site of infection. — fhoracic or abdominal cavity. Type !(>caliiy .— Brazil, Syno/iy/ny. —Esslinger (1973) synonymized the following species with L. hrasiiiensis: L. carolliae Caballero y Caballero, 1944, and L, cahafleroi Gsirchi- Rodrigo, 1954, Other records. —The following arc those listed by Esslinger, 1973: CaroUia perspicilliita in Brtizil (Sandground, 1934), Mexico and Panama (Caballero y Caballero, 1944), Costa Rica (Jimenez-Ouiros and Arroyo, 1960), Venezuela (Garcia-Rodrigo, 1959; Diaz-Ungria, 1963), and Colombia (Esslinger, 1973); '’"Caroilia snhfhiyns" in Colombia (Martin, 1969, personal communication); Glossopha^ai sp. in Briizil (Arandas Rego, 1961 id: Giossophaga .soricina in Brazil (Arandas Rego, 196B;); Phyiiostottms s>p. in Venezuela (Diaz-Ungria, 1963); and an unidentified phyllostomatid bat in Brazil (Arandas Rego, 196U/). Renuirks .— Filaria spicalanon was poorly described from specimens oCPhyl- bhstoma" sp,, CaroUia per.sptciliata, and Stnr/ura HI tarn in Brazil (Mol in, 1858). Although positive identification cannot be determined until the original specimens are reexamined, they are probably Idtotnosoides hrasiiiensis. Litoinusoides ealien.sis Esslinger, 1973 Type htrsf,—Stnrnira lilin/n. 57/c o/iu/ccr/cju.—Unknown, microfilariae in blood. Type locality. —Vicinity of Cali, Valle, Colombia, Other records. —None to date. Litomosoicles chandleri Esslinger, 1973 Type host.—A rtibens jamaicensis. Site of infect ion.^ThomCw or abdominal cavity. Type locality. —Vicinity of Buena Ventura, Valle, Colombia. Other records. —Vicinity of Cali, Valle, Colombia (Esslinger, 1973). RIOLOGY OF THF FHYLLOSTOMATIDAE 29 Litomosoides Esslinger, 1973 Type hosL — Vampymps dorsalis. Site of iufectioti. —Unknown. Type ioeality. —Vicinity of Buena Ventura, Valle, Colombia. Other records,—Artibeus jamaicensls in the vicinity of the type locality also were found to be infected (Esslinger, 1973). Litomosoides fosleri Caballero y Caballero, 1947 Typ e /1 ost.—Glossop /uigct sorkli ut. Site of infection. —Thoracic or abdominal cavity. Type locality .— Pan am a. Other records, —None to date. Litomosoides guiterasi {Perez-Vigueras, 1934) Type host.—A rtibetisjamaicemls. Site of infection. —Body cavity. Type ioeality. —Santa Clara and La Havana, Cuba. Synonymy. —Esslinger (1973) listed the following synonymies: fitdaynema giiiterasi Perez-Vtgucras, 1934; L. hamletti Sandground, 1934; and L. penai Jimenez-Ouiros and Arroyo, 1960. Other records.—Glossophaga soricina in Brazil (Sandground, 1934), in Mexico (Chitwood, 1938) and in Colombia (Esslinger, 1973); Glossophaga 'sp. in Brazil (Arandas Rego, 19616); Tadarida latlcandata and T. brasiiiensis muscala in Cuba (Barus and del Valle, 1967); and Pteronotns parneUi in Jamaica (Web¬ ster, 1971). Wc recovered a single specimen of this species in Glossophaga soricina (KU 102354) from Las Margaritas, 1500 m., Chiapas, Mexico (DLK 358, 23 July 1965). Litonio.soidcs leoitilavazquezae Caballero y Caballero, 1939 Type host.—Macroins waterhonsit. Site of infection. — Body cavity. Typ e /uru//Yy.—Mex ico. Litomosoides teshi Esslinger, 1973 Type host.—CaroUia perspicillata. Site of infectkm.—Thox^ciQ or abdominal cavity. Type locality. —Vicinity of Buga, Valle, Colombia. Remarks. —Filariid nemattxies of the genus Utomosoides are common in leaf- nosed bats; of the 12 recognized species within this genus (Esslinger, 1973), nine are reported from phylloslomatids and the existing records seem to indicate that these parasites are relatively stenoxenous. Of the 10 species found in bats, seven are recorded from a single host species and another, Litomosoides colombiensis, is recorded from only two host genera, Artibeus and Vampyrops. Only two species, Litomosoides brasiiiensis and L. guiterasi^ have been recorded from 30 Sj^KClAt- PUHi [tVVnONS MUSHUM TEXAS f hX’H UNIVHRSITY more than one family of bats. Although these adult filariids tend to be relatively host specific, a given bat may serve as the definitive host for several members of this genus, for example, Ariiheus JwuaicL'nsis has been found to host Lifomosoidt's chatidieri, L. Kdofnhiensix^ L. i^uiiemsL and Litomosokk’s sp. of Chitwood {1938). rfie adult parasites occur in the body cavity of bats. Mature females give birth ovoviviparously to microfilariae, which migrate to the circulatory system and are picked up from the peripheral blocxi by miles that serv'C as the vector. Unidentified microfilariae have been reported from numerous bats including CarotUa custimeiu C, pcrspicUkihi, Ghssophaga sork'huu PhylkmorntT^ sp., and P. hastaius. Such microfilariae probably represent members of the genus Litomosokivs, but no author prior to Esslinger (1973) has attempted to identify these nematodes on the basis of larval structures alone. Fam ily Tr ic h ost rongy 1 idae Biucantha desjtioda Wolfgang, 1954 Type host.—lyesmodus rotmulus. She t)f infevtkoL —-Small intestine. Type locuiity. —Trinidad, West Indies. Other records .—We found this species in several Desniodus rotundus at La Pacifica, Costa Rica (OW'D I66-LP-8, I68-LP-I2, I69-LP-t3, 170-LP-14, 176- LP-6, 12 July 1967), and it has also been reported from D. rotiouins from Jalpa, Zacatecas, and San Bals, Mexico (Wolfgang, 1956). Remarks ,—Specimens of this species are identified easily by the tw'o asym¬ metrically placed cephalic hooks {Fig, 3) and a scries of longitudinal ridges that extend the entire length of the body (Fig. 4). Bidigiticauda vivipara Chitwood, 1938 Type host,—A rtibeiis januttcensis. Type tociilify .—Puz Cave, Oxkutzeab, Yucatan, Mexico. Site o//u/et7/ou.—Small intestine. Other records,—Artibeus jamaivensisr. Costa Rica: l.a Pacifica (DWD 162- LP-48, 12 July 1967) and the Osa Peninsula (DWD 253-OP-42, 28 July 1967); Mexico: (KU 102469) Chiapas, Finca San Salvador, 17 km, SE San Clemente, 1000 m. (JDS no. 927, 4 August 1965); Nicaragua: (KU 9779) 2 km, N Sabana Grande, 50 m. (JKJ no. 4359, 15 July 1964); (KU 97726) San Antonio, 15 m, (JKJ, 6 July 1964); (KU 97773) 14 km. S Boaco, 200 m. (JKJ no. 4569); (KU 97804) 11 km. S, 3 km. E Rivas, 50 m. (TEL, 24 July I 964); (KU 97785) Moyo- galpa, NW end Isla de Ometepe, 40 m. (JDS, 31 July 1964); (KU 977130) Finca Tepcyac, 10,5 km. N, 9 km, E Matagalpa, 960 m. (TEL no. 591,7 August 1964). Artibeus lit unit iis\ Mexico: {KU 1025329) Chiapas, Rutnas de Palenque, 300 m. (JDS no. 721, 17 June 1965); (KU 192469) Chiapas, 4 km, NE Pichucal- co, 100 m. (DLK no. 254, 30 June (965). Remarks ,—The characteristic posterior extremity of this species is presented in Fig. 5, but the functional significance of the divided appendages is unknown. HIOLOGY OF THF PHVFl.OSTOMATEDAE .11 Fio. y ,-—Scanning electron photomicrograph of anterior end of Bincitnffut des/tnuia from small intestine of Dts/ntulitx roiundux collected at La Pacifica, Costa Rica- The irregular surface is an artifact resulting from alcohol fixation. Note platelike teeth in vestibule (arrow). ( X 765) Fk;. 4.—Scanning electron photomicrograph of body surface of Bimantha desnuxhi. Ridge.s (arrow ) are raised above the general body surface, (X 1175) Sl^HC JAl J'UHI.K A r [ONS MUSEUM [ EXAS I'Et H UNlVEKSf lA^ 32 Fifi. 5,—Scanning elec iron photomicrograph of posterior end of body of /^kiinifkatahi rivipunt. (X 675} Eio. 6.~Scanning electron photomicrograph of head of IHifif^iiictnuht yivipura. Fapiiiac (arrov\) and teeth in vestibule NS MUSEUM TEXAS l EC’H UNIVERSITY Siic of Infect ittH.—SnuiW intestine. Type Fazenda Bento, stale of Rio de Janeiro and Canipo Grande, Mato Grosso, Brazil. Other records.^Phyliostiwins hastatiis: Nicaragua: (KU 97418) Daraili, 5 km. N, 14 km, E Condega, 940 m. (TEL no. 396, 25 June 1964); (KU 97416) same location (JKJ no. 4463, 23 June 1964). Ariiheusjwttatvefisis: Nit ARAtitJA: (KU 97800) 11 km, S, 3 km. E Rivas, 50 m, (CER no. 41, 24 July 1964). Remarks .-—Based primarily on the shape of the spicules, Perez-Vigueras {I94hd renamed H. ocfaca/itha as the type of a genus, Parahistiostro/i^iiylus. Yamaguti (1961) and Barns and Rysavy (1971) did not accept the new genus based on spicule characteristics, and vve consider it as a member of the genus Histiosiro/Jifylits. Records obtained from the Index Catalogue of Medical and Veterinary Zoology at Bellsville, Maryland included a report of H. parcuhhxtts Travassos, 1918, from P, spienlatitfJi as referenced by Travassos, 1920. We have not verified this report, Torrestrongj liis toirei Perez-Vigucras Type host.—Macrotns waterhoasIi. Site (}f htfeethm .—Small intestine. Ifpe /ora/zVy.—Cueva del Rincon dc Guanabo, Habana Province, Cuba. Other records. —-This species has also been reported by Barns and del Valle (1967) in PteronoiHS macleayii from Caba San Jose del Lago, Mayaijign, Las Villas Provence, Cuba. Trichtdeiperia leiperi Travassos, 1937 Type fu)Sl .— Trach(}ps cirrh(?sus. Site (ff infeefioti ,—Small intestine. Type i<}ea!ity. — Brazi 1. Other reptmfs .—Caballero y Caballero (1951) also reported this species in T. eirrhosus from Mexico (in Index Catalogue, USDA, Beltsville, Maryland, not verified). Unidentified strongyJict nematodes Host. — Gl(K\si>phaya sortcina. Site of i/tfection.— In embryo. Type locaiity. —Arapua in eastern Mato Grosso, Brazil. Remarks .—-Uamlcil (1934) identified these nematodes only as being hook¬ worms, These specimens undoubtedly belong to the family Trichostrongylidae, but without a reexamination of HamletLs specimens, no further conclusions can be made. B[C)LOGY OF THE FHYLLOSTOMATtDAE 37 Family Trichitridae Capiltaria sp. Type hosr.—Mk ronycieris Site . tin/niami even though both rats occupied the same geographical area and presumably have similar feeding habits. Inasmuch as col¬ lections of coccidians are so easy to obtain under field conditions (oocy.sts are found in fecal material), it is surprising that only three species are recorded from bats in the Western Hemisphere. I'he haemoflageltales of bats have been rather extensively inve.stigaled, though, perhaps, they are still not well understood. The study of blood sporozoans (for H[Ol.OCn' OF [ HE PHYLLOSTOMATIDAE .^9 example, malarial parasites) at' phyllostoniatids, however, is an area about which virtually nothing is known. Again, such organisms are also easy to obtain under field conditions. Only a single adult acanthocephalan is recorded from phylloslomatid bats. In¬ asmuch as that parasite is known only from the original report, it is difficult to determine any degree of host specificity. The genus Neomict^la possesses species widely distributed in carnivores. Bams (1973) reported an acanthella of Pachi- semis sp. in the body cavity of Taphozaus nwiivefuris and an acanthella of Monillformix sp. in the body cavity of Otonycieris hemprichi from Egypt. Barns suggested that bats exhibit reservoir parasitism of an active accumulating type. Whether or not this is true for acanthocephala cannot be ascertained until addi¬ tional reports are available. The remarks concerning the acanthocephala are also generally tme for the pentasiomids. If the few available reports arc indicative, reservoir parasitism is involved here also. The potential value of treinalodes as indicators of host phylogeny or zoogeog¬ raphy has been suggested by Szidat (1955, 1956^/, 1956 /j) and effectively demon¬ strated in some hosts by Kabata (1963), Margoli.s (1965), and a number of other w'Orkers (sec reviews by Kabata, 1963, and Cameron, 1964). Although trematodcs arc reasonably common in phyllostomatid bats, they generally lack specificity in these hosts. The genus Anenteroiremu seems to be mainly associated with the leaf-no.scd bats with some members occurring in the Molossidae. There are no life cycles available for any ircmatode species in phyllostomatid bats. Until studies involving allometric growth (Marlin, 1969) and individual variation arc made and addi¬ tional distribution records are available, this parasitic fauna w ill be of little use in examining host phylogeny. Because the various members of AncnftriHteniu lack a digestive tract, the establishment of this parasite as a laboratory model would allow important advances in the biology of trematodes, especially in nutrition. Only a single species of cestode, Vampimlcpis t'lonffiaus is of interest in light of this discu-ssion, for it appears to be restricted to the Phyllosiomatidae except for a single report in Mitiossns ater. The numerous records from Brazil, Nicaragua, Colombia, and southern Mexico, suggest this organism is the major tapeworm of leaf-nosed bats. Inasmuch as this organism is not rare in occurrence, studies on life cycle, pathology, and so forth may be feasible. In the only life cycle known of tapewx>rms in bats, Kochseder (1969) recovered cysticercoids within the intestinal mucosa suggesting auto-infection of fiynietuflcpis grisea, perhaps similar to that of H. fiatui. Inglis (1965) examined patterns of evolution in nematodes. According to this author, generally, parasitic nematodes arc not host specific but they tend to occur in animals with similar feeding and ecological habitats. Barus and Rysavy (1971) evaluated morphological relationships, specificity, and geographical dis¬ tribution of trichostrongylid nematodes in their respective bat hosts. Their re¬ sults suggested to them that phylogeneiical devcloprnem of these parasites and hosts proceeded along parallel lines. Because more information is available con¬ cerning these ncmatcKles they are reanalyzed here. 40 S0EC IA[ J>UB1.1C ATIONS MUSEUM TEXAS TEC H UN[VtKSlTY The first morphological group of ncmatodc.s listed by Bams and Rysavy (1971) included the genera Sirotit^niavaiuha and /Hacamhn; the former species occurring in Rhinolophidae, the latter species occurring in DestiKnius and Nakiius. We prefer to consider Btcicaiuha as belonging to the sect>nd group for reasons presented below. Because of morphological features exhibited by Slr(}Ui>ykicafj!iuL this genus is the most primitive. Dougherty (1951) and ("haubaud (i9654i, 1965/>, !965r, 1965c/) present arguments that the trichostrongylids evolved from primitive strongylids and the placing of S(r(}/ii>y(aciini/ut in the Ancylosiomatidae retlects this relationship. Both seem to have existed before the Paleocenc (Patterson, 1957) and perhaps split as early as the Eocene. It is tempting to suggest that the origin of trichostrongylids of bats occurred in Ec^cene limes in the Megachiroptera. Subsequently, and prob¬ ably closely correlated with the origin and radiation of the Microehiroptera. these nematodes gave rise to the second group of nematodes described below. 'I he second group of nematodes according to Barns and Rysavy (1971), in¬ cluded the genera SpinosfroH^n-ius, Hisno.sirotigylus, Neohisiiostnmgylus, and provisionally C7?tT/‘op/eroucu/u. This complex (excluding Chciropienme/mi) is characterized principally in having a reduced cephalic vesicle, sclerotized spine¬ like hooks on the head, and a general conical tail, usually with spines. I he genus Bkaantfia is known only from the Neotropical region with B. desf}ihisriostr(}/igyliis and Spitfoxtnfiif^ylus, occur in Old World bat hosts. The remaining genera of bat iiemati^des show little specificity. Although several species are recorded from only a single host species, additional records are badly needed before confidence can be placed on the degree of host specitT city. At present, it is impossible to make definitive conclusions on the evolution of any bat species by examining endoparasites. Such work shows promise, however, particularly in the nematodes where certain genera show- relationships with the hosts: Cheirupferotuatia and Bklit^uficaiukt with Artiheiis; ilisiiustronf^ylus and Tarrestmngylus miU vespertilionids. Ba.sed on such relationships (however ten¬ uous) Barus and Rysavy (1971) speculated that the phyliostomatids served as a sieni host group for development of the trichostrongylids of New^ World bats. Phyllostomatid bats are similar to other groups of Chiroptera in ser\'ing as intermediate, reservoir, or definitive hosts. I heir role as intermediate hosts is minimal. Although Parocephalux vrutali functions as a larval parasite in Phyilos- totuus disc(?k)r and is later transmitted to the snake definitive host, it has not been reported in bats in the last 50 years. .According to the classification suggested by Odening (1968), bats in general are “cureservoir, stationary hosts." This classification would hold true for phyl- lostomatid bats also. Most endoparasites use phyllostomatid bats a.s definitive lilOl.OGV OF THK FHYLFOSTOMATIDAE 41 hosts and not as transitory bioreceptor hosts as suggested for other groups of hats by Rysavy and Barus (1965), Barus and Tenora (1967), or Shulls and Davtyan (1955). Additional collections should clarify these relationships. Summary The cndoparasites of phyllostoniatid bats are reviewed for the first time, A historical review' emphasises the lack of systematic collections of parasites from this group of hats. The major parasitic groups reviewed include the Protozoa, Acanthocephala, Pentastomida, Trcmatoda, Cestoda, and Ncmatoda. New host and distributional records are as follows (a single asterisk indicates that a parasite was known previously from a given host; double asterisks, known previously from a given locality): T’rematoda: Liniamlufn abenans in Pbyllosio- mns disi{flor from Nicaragua. Cestoda: Vampirolcpis ekfnyaius in Glosst^pfiaga sc^ricifia from Nicaragua and Mexico, and in Aritbetts iitumms from Colombia. Nemaioda: Bkicamha titsfmxla in Desniodux rotutulus* from Costa Rica; Hidfgitiau(da vivipara in Artiheiis janiaicensis* from Costa Rica, Nicaragua, and Mexico, and in Arttheus liiuratHS from Mexico; Cheiropteronenui globocephaia in Ariibens liturams from Mexico*’*' and Nicaragua, Caroiiki perspicHkita from Costa Rica and Nicaragua, Arfiheus Jantaiceftsis* from Nicaragua and Costa Rica, Ariihciis phacotis from Nicaragua, Costa Rica, and Mexico, and in A, rol- {ecif.s from Mexicr>; HistioMrongylus con?nan(S in Phyflosfomas discolor from Mexico and Nicaragua, and in Phy(ios!omns hasiaiK.s from Nicaragua; /T ocUi~ canifm in PkyliosfonitLS bustutus and in Arnbeus Januiiccnsis from Nicaragua; Litomofnosidcs hrasdiensis in CaroiHa subfktvus from Colombia; Liioffufsoidc.s gtdfcrkis in Giossophaga sorictna from Mexico. Scanning electron photomicrographs are presented for Vumpyrolcpis idonga- liis, Cheiropterottema glofyocepbctla, Biacufufui dcsoioda, Bidigidcauda vivipara, and Porocephalas CTOtali. Porocephaias croiali is reported from Epicsicus fusca.s for the first time. Addendum After the present review was submitted, several articles have been published, and others brought to our attention, which should be mentioned here. Marinkelle (1976) reviewed the biology of all bat try'panosomes and listed four subgenera, in¬ cluding 20 species, as occurring in these mammals. His first subgenus, Mcgatry- panatu, included the large, broad forms listed in the megadermae group of this review' ( /', pessoai, T, pifanoi). He divided the smaller forms of the classical vesperfilkmis group into two subgenera, Schizoirypanimi (T. cruzi, T, trucZ-likc, ■/'. phyilosfotnac, T. vespenUionh) and Herpaosoma {T. line atm). His fourth subgenus, Trypanozoon, included T. evami. He emphasized, as did we, that the forms in the subgenus Svhizottypanum are difficult to .separate into defined species. An excellent review' of cestodes in the genus Hymetwiepis from bats in North America and Hawaii w'as w'ritten by Rausch (1975). This paper critically evaluated the taxonomic status of the hymenoleptid cestodes and added a new 42 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY spectes, H. lasiouyaeridis, from eight species of bats in North America and Hawaii. Rausch (1975) also discussed briefly the zoogeography of cestcxles of bats. Chabaud and Bain (1974) described a new genus and species of muspiceid nemati>de, Lnkonema iukoscimsi, from Noctiih labialis, Tonatia varnkerk CaroUia persplciikita, Desmodua ronmdus^ Sacvopieryx ieptum, and Epiesicus mekmopterus collected in Surinam and French Guyana. The biology, host-para- site relationship, and life-history of L. lukoschusi also are discussed in this paper. Other papers that merit attention include those by Caballero-Deloya (1971), Durette-Desset and Chabaud (1975), and Chabaud and Durette-Dessel (1975). The first paper redescrtbed Bidigiikuuda vivipara collected from Anibeus lituratiis palmarum in Guerrero, Mexico. The latter two papers reviewed nema¬ todes from European bats, analyzed the trichostrongylid nematode fauna of bats, proposed a hypothesis for the origin of these nematodes, and indicated a possible phyletic relationship between the Tupaiidae and the Chiroptera. Teixera de Freitas and Machado de Mendonca (1960, 1963) assigned several species of nematodes from bats to the genus ParaHintoschius, but Durette-Desset and Chabaud (1975) considered ParaUintoschius to be synonymous with AiPm- toschius. We have not been able to kxate the original papers of Teixera de Freitas and Machado de Mendonca (1960, 1963) for confirmation. Acknowledgments Wc wish to thank Dr. J. Knox Jones, Jr,, for encouraging us to participate in this volume. Many of the parasite records w'ere obtained by J. E. Ubelaker who par¬ ticipated with Dr. Jones in collections in Nicaragua in the summer of 1964 and southern Mexico in 1965 under United States Army Research and Development Command, The University of Kansas, Contract DA 49 193 MD 2215, Specimens collected in Costa Rica were obtained when D. W. Duszynski w'as supported in part by an NSF-Ford Foundation summer fellowship in conjunction with the Or¬ ganization for Tropical Studies and in part by Training Grant 5T1 AI 94-08 from the NIAID, NIH, United States Public Health Service. Review of host-parasite records from the Index Catalogue of Medical and Vet¬ erinary Zoology, United States Department of Agriculture, Beltsville, Maryland, was made possible by funds from the Office of Research Services, Southern Meth¬ odist University. Dr. Ralph Lichtenfels is gratefully acknowledged for hts assistance. Thanks are also due to Mr, Maurice E. Thomas, Tulane University, New Orleans, for allowing us to examine specimens of cestodes from Anibeus (hura- tus from Colombia. To the many additional collectors, contributors of specimens, and individuals who assisted in the identification, we are deeply grateful for their generous cooperation. Special thanks are due Dr. J, Teague Self, Department of Zoology, University of Oklahoma, Norman, and Ms, Lindy Andersen and Mr. John D. Kimbrough, Department of Biology, Southern Methodist University, Dallas. Drs. Edelberto J. Cabrera and Marke W. Talley, Department of Biology, BIOLOGY OF THt: PHVLLOSTOiVtATIDAE 4.1 The University of New Mexico, were of invaluable aid in assisting with the trans¬ lation of the Portuguese and Spanish literature. Literature Cited A(t>siA, J. L,, C, Rom;\n,v. 19.18. Infcccion del murcielago DestnoJn.'; nifundu^ rotn/ulus E. Gcoffroy St.-Hilaire) por Tnpu/u/so/Ho eq/iiNt/tti (Elmassian) y trans¬ mission del "Mai del eaderas” por an mordednra. Mem. Inst, Oswatdo Cruz., 11:291-295. Ai.i.KN, G. M. 1919. Bats. Harvard Univ. Press. Cambridge, x + 168 pp. AraMus Rego, a. 196hv. Sobre aignmas espccies do genero LVo/nfWj/f/e.v Chandler. 1931 (Nemaloda. Filarioidea). Mem. Inst. Oswaldo Cruz.. 59:1-9. --l961/>. Noia sohre a especie Lifof/io.whhx Vigueras, 1914 (Nematoda. Filarioidea). Alas. Soc. Biol., Rio de Janeiro, 5:11-! 4. -, 1963. Nova especie do genero Akhumian, 1946 Parasita de Quiropteros (Cestoda. Anoplocephalidae). Rev. Brasil Biol, 21:11.14. AVai iV. S. C., AND E. A. Wells. 1974. Disappearance of T/ypa/muf/tn/ ryu/isi from a vampire bat colony in western Colombia. Trans. Roy. Soc. Trop. Med. Hyg., 68:76. Baer, J. G. 1951. Ecology of animal parasites. Univ. Illinois Press, Urbana, x + 224 pp. Bakretto, M. P. 1965. Tripanosomas .semelhanies ao Trypartt/sofiw truzi en animais silvestres e sua identificacao como agente etiologico da doen^a de Chagas. Rev. Inst. Med. Trop.. Sao Paulo, 7:105-11.^. Marls, V. 1971. 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L„ V. J. Tipion, and a. Kiewjic/.. 1966. The Streblid batflies of Panama (Diptera Calypterae: Streblidae), Pp. 405-675. in Ectoparasites of Panama, (R, L, Wenzel and V. J. Tipton, eds.j. Field Mus. Nat. Hist., Chicago, xii + 861 pp. Wueai, B. E, 1975. Einii^iiii nnnyi sp. n. tProtozoa: Eimeriidae) from the eastern pipistrelle, Pipistrellns Mihjlavn.s, from Alabama. J. Parasitol., 61:920-922. WoiJ'GANc., R. W. 1956. An additional note on the distribution of Hiacanfha desmodu, Canadian J. Zoo!., 34:209. Wood. F. D., and S, F, Wood. 1941. Present knowledge of the distribution of Trypnno- sonni trnzi in reservoir animals and vectors. Amer. J, Trop. Med., 21:3,35-345. Wood, S. F, 1952, Mamma! blood parasite records from Southwestern United States and .Mexico. J. Para.sitol.. 38:85-86, VV, Y, 1917. ta review of the paper by Iturbc and Gonzalez, 1916, describing Trypn/io- .Siwiii lifn’iitns from yampirops iineafafi], Trop. Dis. Bull., 9:342. YaMaguti, S. 1961. Systema Helminthum. Voh 3. The nematodes of vertebrates, pari 1. (nterscience Publ., New' York, 679 pp. - — , 1969. Special modes of nutrition in some digenetic tremaiodes. J. Fish. Res. Bd. Canada, 26:845-848. —-. 1971. Synopsis of Digenetic trematodes of vertebrates. Keigaku Publ., Tokyo, vol, 1, 1074 pp. BIOLOGY OF THE FHYLLOSTOMAKDAE 51 Zeii-don, R., and R. RosaHal. 1969. Trypa/ioxomu leonichtsdeaiiei sp. nov. in in-sectivoroys hats of Costa Rica. Ann. Trop. Med. Parasitol., 63:221-22fi. Zei EDON, R., AND P. L. VtETO. 1957. Hallazgo de Schizotrypa/tutn vesperiilio/iix (Bat¬ taglia, 1904) en la sangre de murcielagos de Costa Rica. Rev. Biol. Trop., 5:123- 12B, --, 1958. Comparative study of Scfitz.\\opfui}’ti Mork itui loAoplasmatidac 'I'o.xtiphi.'irmi yioulii Artihens tiiiir(iin\ li ypanosomaiidat; Trypeino-yonui oriizi Aniht'us janiait Arsihi’tis finirmn.'i. Ctindfio pvrspii-illiihi Dosnukho ronmdn^ Gh >.\-Wjp liapn .\orh'inn rhyllo'<{onii(.'idisroff)r Phylio.'^ltonif.x hii.xfIIIu,x Ufodenini hiiohnlton Trypiiiii>M>nM ike Anihen.', cina-rn.'^ Aril hens jdnuiivioi.xis Arlihi'ti.x liltfrnni v Cnroflid ftcrspicilhuii Chn( liop/iylln niordnx lAiin'hophyfli! lininnixi \Iixrtutyvf('fix niepnlofis Xfifiion hctnu'llii I’hyfliisiofnnsdi.scolor P/iyihi.sfoniifs clonpitfnx /‘hylloxlofuns Itti.stiiins Urodxrnm hifolktann Vanipyrinn spxi innn Trypiniosonni ('(.iHfniff}} Di smodns rottindnx* Trypanosikna evnnxi ArlihiPix jn/nnii-enxis* Ciindtia pvrspk iUaui* Dcstnodux I'oltnultn Di'snitHtnx rotnndn.s* Ol(>xsiiphiipii stnicinn* fdtylltfsUnnns hnxtiitns* Trypnno.sonni lineatns Vn/npyrops iint'nms I'rypni josof r i a pe ss < hi i Ariihens i itifrens Artthi'ns jnin a ivens is C(irtdlia pi’rsp ic ilhitn Choi'ronisciis ininor Desniodns roHindns 'Irypanosotint phyllosfonnu’ ('nro/lia ptrspicdiaia Trypni11 >stnini ptfanoi Arlihi’us Htnrnins Idiyfiosionuis host (it ns 'S'rypnntisonHf Art the ns liininins Glossophnpn sork'inn TiypniUisonm vexpertiHonis Aiionnt vandifer Carfiilin per spiv a In l a Ckoeronisvns minor Giossiiphnpn sorh inn Mnerotus wiuertniusii Hi ytlostoinns hast a 11 ts Tryptinosomn sp. (>ney;ndei/nne-iypc) Anonrn enndifer Cnrollin pvispivillnnt Ciioeroniscns minor DesnuHins rotnndns Gh issop}!(ipa S{irieiiin Trypnnosoma sp. ( verpertiiionis-iypQ) C(!roUin perspivillain Lt I n<'hopityl{a mordax Tiypanosonm sp. (not 7. (‘jv(.'f-like) Choeronisensntinor At AS T Hot J.PIIAl A <>1 igacan t hot hy nch idiie i\'eonrivnln novidlae A rtihens jarnaicensis Pf.n 1 AST ()M in A [’orocephal idae Pi trove phi ilns i' ro ta I i Phyliosi{>nin.\ disvoU/r TkKM A KJLiA Anenieroiremat idae Anenierofrenns iV/i(v o;i V( / eris mepnlotis Ancnteroirenni ednl■ndoeaha!|eroi Phyilostomns elonpafns A iwnierotrcma fre ilnsi Mieronyel eris h irsnta BIOLOGY OF IME 1>HVl.LOSTOMATJDAE >3 Af'^’E^Dlx I.— Cotiti/mtii Ani'fUt‘ii>}rcma filipiifiafiu/>i PhyUoxiotn us fhffipunix Aiienu'rotri'iiia MUtikardi f’hyllifsifJinitx hustoiiix Dicrocoeliiduf Afht'xinid parkcri Ariiheii-sju/imiveft s is f\innnL’fiuii'(ph!X cty/upurtux Mii rfuiyi U’ris hchtii Leciihodendciidiie Lci'iihoth'/idi intu prictd' Ardbi'us jufiniict’/i .v is iJ/iunuhtni aherruns .MiH’iit/us miierlufdsii Phyllt.)st<}/fins discoit}!' Liiiiatidufu isihniit'us .\7 icftitiyt UTis hiistita Liftiiinduift /iit‘nsis Mm'nuns H‘a!t‘rhousii Urotremiitidae Urtitrcnui sctihruln/i) PhyllitsiDHUis sp. Fhy!tt>s/f>iiius Cj;xt(>da Anoplocephalidae OiH-horisi k u itnn himni Glossophny’a st>riri/}(t Hymenolepididae Vil tup iralcp is cio/ipiifus Acnhi'i/s /iinniiiis G!assi)phtt^>u soric ina Fhyltu.MDtUKS /utslitfiis NtNtATODA [Jipetalonematidae Litomosoidi’s sp. Anihetis jitmtiicensis 07f^ v.w }phii}i(i sork ifui Liu>tiios<>hii’s ariihei Ardht'iis cincrciix Liroiiiosfddcs hrasiHcnsis Condlia pi'rspi( iliiim Condiiti suhjliivus Clossopluipti sp. Gtossoplhipii St>rii'ini{ Phylli>s/i>i)ui.\ sp. L i(oi!u>st>idi's cdtiensis Sfnrnint lilintii Lifi^nufsoidi’s chandieri A rt ibcns Jatnoii'ettsis Liiof/iosoidcs cotomhiensis Arrihens jaiiiait I'fisis Vciinpyn>ps dorsaiis LiliitmKKoidt’sjdsli'ii GiiKssopfuipti soririmi LiltonosoidI’s pa il onisi Ai I ihet IS ja nmii' o n sis Giossophai'ii sp. G7^m(^p/^^^^'^ll sto'icithi Liloiih)soidi's h'ouiliivozyius ociiuunffm A riibeus jumuicousis Pbyliosromus basiadis Ttnresinm pyl u s lon o i Macroius o'ofi'rhousii TrichoUdpcriii Icipori Tnichops firrliosiis Unidentified stmngylid nematodes Glossophupci sork'inu T richuridae Cupiilnrki sp. Mii'lonycn’ris IIIopu!itk‘s ariihei Protozoa Trypanosoma (vvf'zFIike Trypaiio.soma p fs,u?soide .v gt / iteras / Protozoa Trypanosoma era zi Trypanosoma ike Trypanosoma pessoui Trypanosonuf evnnsi Trematoda Athesmia parkt ri Lecithodendriam privei Artiheus litnrasas Ccstoda Vampindepis eionpaius Nemaioda Bidipifieanda vivipara Protozoa Toxopfasma }>ondii Trypanosoma ernzi Trypanosoma c/v/^lFlike TIypimostrma pi/anoi Trypanosftma ranpelTWke Aniheas phaeoiis Nematoda Cheiropteronema phhoeepbala Ardheiis ltdfecus Nemaioda Cheirop(ert>nema plohocephata Carollia perspivi/tafa Nemaiotla Clieirt/ptenmema plohtKephaia Fiiaria serpicidnm LiloniifSifidi's hrasiliensis Lilomosoides leshi Protozoa Trypanttsifma trnzi Trypanosoma trnzi* Try pa nosoma craz i-l ike Trypantruona eeansi Trypantrsif/na pesMn/i Trypantrstnna phylltnatnane Trypa/nrstona vesperfi/hmis Trypamfsoma sp. (mepadermae-type) Trypanosimia sp. (i esperfifhmisAypc) "Carid/in snh/Ias as"' Nemaioda LdtnnifSifides hrasdiensls Clroeronisias min/n Protozoa TrypantJstnna rrnzi-like Tryp a ntfstmia pesstfai Trypi^mt.awni v'£',vpr) 7 ( 7 /o/n'.\ Trypantsstnna sp. (mepadernu/eAype) Jrypanosoma sp. (not T. trnzidikc} Desmodf/s nnimd/ts Nematoda Binamrlnt desnaida Protozoa Trypiintrstnna cm zi Trypanifsoma erazTIike Trypaniiumta c i'ansi J'rypani?S(}ma e l ansi* Trypanefsomn epn/nam* J'rypanirsoma pesscfn i Trypanosoma sp. (mcfiadermaeAype) Glossciphapa sp. Nemaioda /V(j/1f£J.vo id <'.v hrasiltensis Liiomosoid cs p niter a s i G!ossi>phapa soricina Cestoda Ooehoristicn immatnra Vampirotepis cl<>n.i'tifits Nematoda Lifomosoides sp. Litomosoides brnsiliettsis Lit on losoidcs ft >sf er i Lifoinosirides pniterasi Protozoa Potyctiromttphiins deanei Trypanosoma ernzi Trypanosemta crazTlike Trypa nosoma evansi Jrypanosoma ranpelTWk^ Trypanosoma sp. UnepaderntaeAype) SPEClAl, PUBl.JCATtONS MUSEUM TEXAS I’ECH UNIVERSITY ApPtN'D1X 2.— Cl>tt(ililted. TryfhiiiitMiitiit ve.\peti i(k m is Liitichophy/lu null iliix Protozoa Ti ypunosotno ike Ti'ypttihi:s<}f}i(t sp. ! vc-speniHiJitis-typc) Lo/it fuiphyliu (iuiiiHisi Protozoa 7 rypt 11 ui.sotnti rr //-1 ike .V/( leniru.s etj/i/t)f/i(c/i.v Nematoda G/yp t < ist n iit^y I US rofUii' i.s Miu rofux wiilerhtiu.s it Nematoda Cupillufu I yipueru.si L itii/tutst) iift’\ le fI ///hi i 'tizpiie~(iV Torrestmispylii.s ton ei Protozoa rrv/>^j):o.vo//!f f ve.speti ilk in i s T rematoda fjitituuluni aheiran.s LiiiiiUuliiin okltihiiiueit.sis Micnmyt (ei i.s hehni Tremaloda hini/iietiHleiphis: eoiiipai tii.s M irn>ny 1 7 eri.s n j eptiitifi.s Protozoa T. f7f^:(-likc Nematoda Capilktyhi sp. Tremaloda A iie/itcrtiftenia (iiiri} run :Vfit ronyc/eri.s hirsutii Tremaloda A tienteio/n'/na //r itasi Li mat III it III ist hi 11 k 7 / ,v .Mitnon hcuiH’tdi Protozoa Ti'ypoiui.soniti (ike Fh yfIo/iy rte ri.s p< >c'>' i Nematoda CitpiUuriii phyliottyeteris His / /oA7 ro /! ,ey/ f/.V f' f ovjf; / ^ (A FhyUosioiiiiis sp. Nematoda Faiiria .serp iciilnin Lift I/! lost I id t’.s h ras itie / is i.s T rematoda Urinrcifitt M tihridiiH! Fhyi1 1 fsttype) UrtHierniu hilohutuin Protozoa TrypafifiSiintcs entzi Tiypano.sontti crtizi-h\:^c ytintpyrups dtnsali.s Nematoda Liititiio.stiides [7 iftnnhiensi.s Vtinipyraps Iineaf us Protozoa Tryptinosonui /ineuins Vtinipyrtitii spectnttn Protozoa 7’r.\' p f; /j oAf j// a; ( 77 / r /-1 i k e [Possibly a htfi'^us cuhniit for CtirftliUi ui/inipt. Eiis.] ECTOPARASITES James P. Weimi, Jr., and Richard H. Loomis Phyllostoniiitid bats harbor an assemblage of ectoparasites numbering more than 230 species that represent 1 5 families of acarines and two families of dipteran insects. Among all chiropieran families, only the vespertilionids, with 18 acarine and six insect families, have more parasites. Streblids account for the greatest num¬ ber of species of any of the phyilostomatid-infesting groups, having 83, comprising 20 genera. The nycteribiids are represented by 13 species in the genus BasiluL The remainder consists of 150 species of miles and ticks presently recognized in 49 genera. We found no substantiated literature records of siphonapterans, anoplurans, cimicids, or polyctenids regularly associated with phyllostomatids, although Oeas were listed as questionable parasites of Venezuelan phyllostomatids (Tipton and Machado-Allison, 1972), tw'o species of Hesperoaem’s (polyctenids) were men¬ tioned (Hoffmann, 1972) from Gfossophaga species, and H. funiarius (W'cstw'ood) was reported (Maa, 1961) from Venezuelan PhylhKStoniiiS hasunus. In addition, Gerberg and Goble (194!) recorded two species of mallophagan lice from leaf-nosed bats, including Physconclloides (near gaUipi\i,H’tisis) recovered from a Panamanian Canillia pet-xpicillaui and thought by the authors to represent a possible bird-bat association, GeoniyiI(fecns geomydis, which normally infest pocket gophers, was also mentioned from a Mexican Lepitmycieris nivaHs. Other insects, for example cu lie ids, psychodids, and ceratopogonids, which may be associated with bats were not included in this review. A comprehensive worldwide list of bats and their acarine parasites, including phyllostoniatid-associated mites and ticks, w-as compiled by Anciaux de Faveaux (1971). Additional recent citations are listed under each family. Macronyssidae Oudeinans, 1936 Sixteen species of macronyssid miles comprising six genera have been described from phyllostomatid bats. Radfordielia Fonseca, Parichoronyssns Radovsky, Macronyssoides Radovsky, and ChitoecePes Herrin and Radovsky are in the Mucronyssus- group and are considered by Radovsky (1967) and Herrin and Radovsky (1974) to be the more primitive and closely aligned with the laclapine stock, whereas the Sfredionyssits- group has more highly specialized representa¬ tives. Nycteronyssus desnu?diis Saunders and Yiinker (1973), based on a single female from a Diaemits youngH, was considered by its authors as possibly of another demianyssoid family. Radovsky (1967:59) suggested that the macronyssids evolved from progenitors that closely resembled extant laelapids because certain features are common to Neokielaps Hirst and Nohdaelpas W'omersley in the Laelapinae and to Bewskdla Domrovv, khoronyssns Kolenati, and Patichoronyssiis in the Macronyssidae. This relationship between the laelapines and macronyssids seems to suggest a relatively recent association wuth bats. 57 .^8 SPECIAL PUHI.ICATIONS MUSEUM TEXAS ] Et H UNIVERSITY Parusilk development by an invading organism on a new host Uonc of diversi¬ fication. Parichorottysstis and the others of this group apparently are now in the early phases of this process, A number of the RailfonHeila species have adapted, as depicted by the protonymphal stage, to a higher level of specialization. Radford- iidki ntOfutphyUi, R. orkola, and R. a/uairae, all described by Radovsky el al. (1971), are known only from protonymphs found in tlie soft palate tissue of several phyllostomaiid species. Each of these species may cause denial and peridontal destruction in bats (Phillips. 1971; Phillips et ul., 1969; Radovsky et ai, 1971) and at least one bat, Leptonycterh scuibornk may have evolved certain tongue mod¬ ifications that prohibit the establishment of mites in their mouths (Greenbaum and Phillips, 1974). In most other species, the protonymphs and adults feed on skin tissues or blood while situated in the fur, rarely on the naked wing or paiagial mem¬ branes. When compared w'ith species of either Mucrottyssimtes or Park honmyssus^ the occupation of the intraoral niche by three species of Radfordielkt suggests a longer affiliation wuth phyilostomatids, BkK)d feeding may occur in Mac- ro/mwk/e'A (Radovsky, 1967: 12), possibly accounting for its large number of host species as seen in M. kocht. Once adapted to hernatophagy, the procurement of new' hosts seems less difficult than in the case of adaptation to dermal hisiophagy, w'hich involves exposure to a greater variation in nutritional and other components. Ch 'troecetes is represented by C kmihophyila and is know n from a single fe¬ male specimen. Herrin and Radovsky (1974) included this species with subgroup C of the Muenujyvv/M'-group based on chaeiological criteria. Other stages of the life cycle are unknown. Although Stea!ouysstisjoaqiuiui{¥omQC‘eL) has been recorded from Giossophaga sorteina, it probably is more commonly found on vespertilionids as demonstrated by its occurrence on Myotis aihescetts from Paraguay (Radovsky, 1967) and nu¬ merous other records of Sicatonyssus species from vespertilionids (Anciaux de Faveaux, 1971). Bat macronyssids are probably in or near the bat roosts when not on the host. Mating, however, most likely occurs on the host. Only unembryonaied eggs have been found (Radovsky, 1967:13) in most species of Maenmyssus, Macronys- sokies, and Radfordie{ki\ however, Radovsky (1967) postulated that ovoviviparity may occur in some species of Macronyssus. Aside from morphological correlations and overlap of features of egg development, relationships may be draw n between hosts and geographic ranges of Macronyssus species and the phyllostoniatid- parasitizing macronyssid genera. Macrouyssus has a cosmopolitan distribution as do its primary vespertilionid hosts. One line probably provided the common origin of Radfordielki^ Chlroevetes, Parkhoronyssus, Mucranyssoides, and Mae- ronyssus. Evidence for the rise of these genera from a Macronysstis-hkc progenitor may be seen in the relationship found today between Old World species of Mac- roiiyssiLs and Old World vespertilionid, rhinolophid, and hipposiderid bats. Macrotiyssits granuiosns (Kolenati), M. kfriginiatiiis (Kolenati), M. ridnolophi (Oudemans), and M, ctfreamts (Ah) all are found on vespertilionids and on rhi- nolophids or hipposiderids, or both. In the New' World, remnants of Macronys- At/.v-vcspertilionid associations may still be found in the numerous reports of M. B[OLt)GY OF THE PHYLLOSTOMATIDAE 39 croshyi (Ewing and Stover) on vesperttlionids, especially species of Myotis. MacroHyssiis jonesl (White) is known from North American vespertilionids as is M. imkiens Radovsky, w'hich also has been recorded (Radovsky, 1967) from the phyllostomatid Leptonycteris nivalh. Host specificity coupled w'ith the adaptive strategies of certain macronyssids, for example, species of Radfordlella., on phyllostomatids suggests a long host association and a New World origin, possibly on vespertilionids. SpiNTURNiciDAE Oudemans, 1902 Sixteen species of wing mites in one genus, Penplischrus Benoit, presently are recognized as parasites of phyllostomatids and are numerous and widespread from Mexico and the Antilles to Paraguay. Cameroniem was established as morphologically distinct from Petiplischrus by Machado-Allison (1965) and its apparent host specificity on species of Ptero- notiis and Mormoops prompted Machado-Allison (1967) to suggest separate familial status for the mormoopids. Later, Smith (1972) separated mormooplds from phyllostomatids on the basis of various morphological criteria and referred to Machado-Allison's (1967) statements about parasite-host relationships for additional support. An apparent parallel to the Periglhchrus-^Cameronieta di¬ vergence in the New' World may be seen in two Old W'orJd genera. Eyndhovenia Rudnick (I960) monotypic with E. enrya/A (Canestrini), is found principally on rhinolophids, whereas Paraperi^lischrus Rudnick (1960) is known both from rhinolophids and hipposiderids. Spinturnicids on both phyllostomatids and Old World rhinolophids and hip¬ posiderids probably arose from a line common w ith that of SpintHrnix Von Heyden. Spinmniix is nearly cosmopolitan, primarily on vespertilionids, especially species of My Otis and Eptesicus, genera common to both the Old and New' World, Al¬ though not recorded from Neotropical leaf-nosed bats, at least two species of Spimurnix have been recorded from rhinolophids (Anciaux de Faveaux, 1971), An Old World origin for the spinturnicids seems to be suggested by the diversi¬ fication of taxa and by the geographic and taxonomic ranges of their hosts. Dis¬ persal of wing mites to the New W'orld probably occurred on vespertilionids w'ith subsequent infestation of phyllostomatids. Additional information regarding the new taxa and parasite records from phyl¬ lostomatids may be found in Dusbabek, 1970; Dusbabek and Lukoschus, 1971/7; Hoffmann et ai, 1972; Kingston et ai,, 1971; Machado-AMison and Antequera, 1971; Tamsitt and Fox, 1970h; W'hiiaker and Easlerla, 1975; and Whitaker and Wilson, 1974. Spelaeorhynckidae Oudemans, 1902 Originally described as a tick, Spekteorhynchits praecitrsor Neumann served shortly thereafter as the type genus for the family Spelaeorhynchidae, The origin¬ al specimens have been searched for and are presumed lost (Fain et ui, 1967), Considered for some time to be closely related to ixodids, later workers (Baker SMKt’IAl- !>UHLJCAI IONS MUSKUM 1 EX AS J Et H UNIV ERSH Y ftO and Wharton, 1952; Fain ei ai,. 1967) have placed this group in the Mesostig- mata. Two of the three kmnvn species are found only on phyllostomatids. Spe(ari>thy}uhiis pniecursof has been reported (Anciaux de Kaveaux^ 1971; Dusbabek, 1970; Hoffmann and de Barrera. 1970; Tamsitt and Fox, 1970/0 from leaf-nosed species taken in Puerto Rico, Cuba, Mexico, Dominican Re¬ public, Colombia, and Venezuela, Mofiophyllns n'dtnanl from Puerto Rico (Tamsitt and Fox, l970/>; lamsitt and Valdivieso, 1970) is the only known host for 5. /nifnopJiylii Fain tv al. The species S, chilonycief i.'i Fain tv ai, is based on a single female from Ptcronom.s nthiyituKStts, a niormoopid, taken in Guate¬ mala (Fain tv vA, 1967), Twt> female specimens of Spclaei>rliyn(iuts species i/iccrfae .vet/Av also were mentioned by Fain tv t;A, (1967) from Brazilian CaraiUn hfvvlcaifikt. Both S. pnierursot and S. tnonopiiylii have been removed from the lower portion of the ear, often from the tragus, and usually embedded deep in the skin. Only female spelacorhynchids have been collected although larvae have been dissected from gravid females. Fain tv al. (1967) postulated that the males of these mites arc free-living nidicoles, although parthenogenesis also may he a possibility. I'he origin of Spelaeorhynchidae may be from the laclapoids and the highly specialized features that make it distinguishable from extant relatives are des¬ cribed as fixation to its hosts (Fain tv vA, 1967). This restriction to the hosts may retlect an inability to adapt to new hosts and may account for the small numbers of spelacorhynchids encountered tt>day, lxo[>inAE Murray, 1877 I vvo genera and two species of Ixodid ticks have been reported from leaf- nosed bats in the New World. Atnhiyomma huigirosin' (Koch) is known from a single nymphal record re¬ covered from a Venezuelan Arfibcun Itfiintitis and also was listed (Jones tv af., 1972) from a number of prehensile-tailed porcupines and a squirrel, Svtitnis j^ranaie/isis from Venezuela. In addition, Cooley and Kohls (1944) reported A. longirosire (as A. aveculcns Cooley and Kohls) as occurring on birds from Texas, Belize (formerly British Honduras), and Panama. A single larva of /.vo(7c.v (Kohls, 1957) was listed from Anonni geof- froyi and a male, female, and three nymphs were found crawling on a cave wall in Trinidad, As mentioned further by Kohls, this larval tick represented the first bat record for the genus Lxades from the New W'orld, but it is unknown whether /. dowmi regularly parasitizes bats. Jones tv al. (1972) subsequently have recorded a number of cases of AvfWc.v species from Anihats jamaicensis and two species of Stunitra from Venezuela. Species of Ambiyonima and Ixodes arc generally not hat affiliates and the records from phyllo.stomatids probably represent accidental associations, al¬ though three species of Ixodes apparently are usual parasites of Old World leaf-nosed and vcspcrlilionid species ( Anciaux de Faveaux, 1971). HIC)l,OGV OF THE PJ!YD.OSTOMATlt>AE 61 Argasidae Canestrini, 1890 Nine species of Onuiiunioros Koch and two species of Aufricala Cooley and Kohls are the soft ticks presently known from phyllostomatid hosts, (huhho- doros vi^uerasi Qoohy and Kohls has been placed in the subgeniis Subparmtuus Clifford, Kohls, and Sonenshine; six species belong to the subgenus AUTtorohius Pocock; and two species, O. luhuon Kohls, Clifford, and Jones (1969) and O, peruvkuuts Kohls, Clifford, and Jones are unassigned to subgenus. Only a single larva of fA mimon has been recovered from a phyllostomatid, Minwn crenufatwu from Bolivia, whereas 38 specimens were removed from a number of Uruguayan Eptesicus hmsiUcnnis. The larvae of fA penivkums have been re¬ ported by Kohls a cd. (1969) to resemble superficially (A {Alectorohhis) kettyi Cooley and Kohls. The species (X (/4.) pitcnoricensis Fox has been recovered from lizards, various mammals, including burrow dwellers, and a numberofem- balloniirid, molossid, mornioopid, and noclilionid bats, and there is a questionable record (Jones ct u/., 1972) from a Venezuelan Aniheus Ihunaus. Looking at hosl-parasite relationships, indications are that the members of the subgenus Alevtorobius may have arisen from a line closely related to the Old World Puvtovskyeiki, as each subgenus parasitizes a wide variety of hosts such as reptiles, birds, and burrowing and nonburrowing mammals. The mode of distribution from the Old to the New World, w here Aiectorohius may have origin¬ ated and radiated, is unclear, but avian or rodent hosts, or both, are suspect transporters. However, because members of both subgenera are found on birds, O. (/I.) capeusis Neumann, taken from marine birds that live in tropical and temperate regions world-wide, and O. {A) dentuarki Kohls, Sonenshine, and Clifford, also from marine birds living on islands in Pacific and Caribbean waters, are of particular note. The possibility of the original migration of Aivcfor- ohius progenitors via a bat carrier common to both the Old and New Worlds seems remote, at least relative to present day distributional patterns of Ornhho- doros species. Even though specimens of Alcaorohins have been collected from vespertilionid, emballonurid, and molossid bats of the New World, only one record of this subgenus has been recorded (Anciaux de Faveaux, 1971) from any Old World representatives of these bat families. Already adapted to feeding on a w'ide variety of mammalian hosts, invading species of the Alevionybim line adjusted to using bats, especially phyllostoniatids, as hosts. A number of mainland licks developed as species limited to the phyl- lo.stoniatid host type, then apparently spread to adjacent Caribbean islands, and then from island to island in the Antillean chain. An example of this is provided by (). {A.) azteci Matheson, which has been reported from Mexico, Panama, Venezuela, and from the islands of Trinidad. Jamaica, and Cuba (Kohls a ai, 1965; Anciaux dc Faveaux, 1971; Jones a ak, 1972). (huidiodoro.s {Subpar/uatus} vigifcrasi differs greatly from the other species of Oruhhodoros (Clifford el ai., 1964). However, its affinity for certain bats suggests an origin from the Antillean Aiectorohius line, as (). viguerasi para¬ sitizes Cuban Phylhmycteris poeyi and BrachyphyUa miua^ and Puerto Rican ErophyUa Nuuhifrons (Tarnsitl and Fox, 1970/f), all phyllostoniatids. A strong 62 SPFX’tAl. PUBLlCAf iONS MUSEUM TEXAS TECH UN[VEKSITY association with mornioopids is indicated by the records of O. vi^iientsi on Ptenntoiu.s from Cuba and Jamaica, and on Ptenmoius and M[ormo(?ps from Trinidad. Furthermore, Pi trout >1 us parueUii has yielded a number of these ticks in Panama (Fairchild a a/., 1966), as have Mornioops uie^^alophyiiu and species of Pieronoius from Venezuela (Jones ei ciL, 1972). The species of Afi(ria}la seem to exhibit the same host and general distri¬ butional patterns of O. viguorasi on the mainland and Caribbean islands. Two of these, A. margiuaius (Banks) and A. silvai Cerny, have been taken from Phyllofiycieris poeyi, endemic to Cuba (Hall and Kelson, 1959), but are also known from Cuban rnormoopid species (Anciaux de Faveaux, 1971), and A. silvai was recorded by Jones ei ai (1972) from Venezuelan Mormoops megalo- phyiki and Picronotus, Mainland phyllostomatids have not yielded Antricoiu, which may suggest that Aniricola arose from island-inhabiting Oruithodoros types on phyllostomatids or on mornioopids. Is land-hopping mornioopids may have brought the ticks to the mainland where A. ttiexUamis seems specific to rnormoopid species such as Mexican and Panamanian Pteronotus in addition to being found in bat caves in Guatemala and Mexico (Fairchild et r//., 1966). Amricofa coprophilus (McIntosh), although never recorded from a chiropteran host, is knowm from caves and mines in Arizona and Texas inhabited by ves- pertilionid bats (Cooley and Kohls, 1944), and numerous A, gramisi de la Cruz (1973) were recorded from a cave in Cuba. The overall trend of development of species and distributional patterns seems to indicate an origin of a line of phyllostomatid-infesting Ornithodoros {Alec- {orohius) species in southern Central America or northeastern South America. Radiation has occurred northward into Mexico and the southwestern United States, eastward onto the Caribbean islands, and southw-ard into other parts of South America, The northern and southern distributional patterns appear to be retlections of one another with adaptations by these ticks to more temperate- ranging bat species such as certain vespertilionids and molossids. In the north, O. {AA yumatensis and O, (.4.) rtKSsi, both parasitic on tropical and subtropical phyllostomatids, parasitize vespertilionid bat species as far north as Arizona (Cooley and Kohls, 1944; Kohls ct ai., 1965; Jones et aL, 1972). South¬ ward, the species O. (A.) boll vie us is, for instance, has been recovered from Bo¬ livian Mytuis nigricans and Moi(}SSus major (Kohls and Clifford, 1964), and O. eptesicus is known (Kohls ei al., 1969) from Venezuelan Epiesicusfurinal is. Erevnetidae Oudemans, 1931 Five species of speleognathine ereynetids all of the genus Speieochir Fain are known to infest the nasal passages of phyllostomat id bats: SpeletKhir aitkeni Fain (1966) from Anoura geoffroyi taken in Trinidad; S. brasiliensis Fain and Aitken (1969) from Vampyrodes caraccioloi and Ariibeus jamaicensis’, S. bar- bulaia Fain and Aitken (1970) from Mimon crenidaiuni of Brazil; S. phyi- lostomi (Clark, 1967) from Phyltosiomus fiastatiis of Colombia; and S. carol- iiae Fain and Lukoschus (1971) taken from Caroltkt per.spicillaia in Surinam. BK».C)GV OF THE FHVLLOSTOMAT[DAE 63 Some ereynetids are free-living predators, but most are adapted to a mucosal environment and the origins of the Ereynetidae seem to He with a ground or plant-living ancestor that adapted to mucosal secretions or mucosal secreting environs. Rodents are the most numerous and widely represented mammalian hosts although many bird species also harbor speleognathines. A common, rel¬ atively recent, ancestral history seems to be suggested for both mammalian and avian ereynetids as both types are w^etl represented and included together in the Speleognathinae. Comparisons of rodent and bat liost records have revealed certain related patterns. For instance, species of Panispeieo^mtthoijsis Fain arc known from Africa, Europe, Korea, and Australia from different species of murid rodents; species of Speicorodem Fain have been reported from Africa, Europe, the United Stales, and Panama from sciurid rodents, and from Australia and Panama from murid rodents; and cricetid rodent records are cited from Panama, Trinidad, Brazil, and Holland (Fain, 1970 /j). A similar Old and New' World distributional pattern for species of the same genus is exhibited by Neospeieopmahopsis on vespertilionid bats from Europe (Belgium) and from the United States. All but one of the species of Speteovhir arc know'n from Neotropical phyllostomatids; S. duhoisi (Fain) is from an African Nycieris. Interpretation of these patterns leads to the assumption that two possible routes of dispersal may have been involved. The Old World murid rodents ap¬ parently provided a possible dispersion mechanism as they spread worldwide, with speleorognathines secondarily infesting sciurids, cricetids, and other mammals. The other possible route centers with the vespertilionid bats, which may have transported speleognathines to the Neotropics where the leaf-nosed bats became hosts and sites of development for these mites. The genus Speieochir appears to have originated in the New World on phyl- lostomatid bats, from Old W'orld progenitors carried to the Neotropics by one form or another. The occurrence of congeneric species in two widely separated regions— S, duhoisi in Africa and the five Neotropical Speleoihir species—may suggest a greater distribution for nycterid bats in the geological past or that the generic position of S. duhoisi is in need of reevaluaiion, Mvobiidae Megnin, 1877 More than 50 species of myobiid fur mites are recorded from chiropteran hosts. Eudusbabekici Jameson contains nine of these species that arc known exclusively from leaf-nosed species. Five species of Eudusbabekia have been found on Cuba or Isla de Pinos (Dusbabek, 1967), a small island near Cuba: E, cemyi (Dusbabek) from Brachyphyila tuina, E, danieli from Phylionycteris poeyi, E, safnsinaki from Macro!us waterhousii, and £. viguerasi from Arli- heusJauiaiceusis, Jameson (1971) subsequently named two species, E. lepidoseta from Srurnira (ilium and £. phyllostomi from Pbyilosioniux discolor taken in Nicaragua. Later, Vomcro (1972) described E. argatioi from Desmodus roluudus taken is San Luis Potos'i, Mexico, and Fain (1972) described E. urodermae from a single Brazilian Uroderma maguirostnim. Two other, £. Jimenezi (Dusbabek) SPECIAI. HUti] 1C A'JIONS MUSEUM 1 EX AS TECH UNIVERSITY (i4 and E. sa^uei (Dusbabek) arc found on species of Ctihan Pferonomx, a mormo- opid genus. The genus kmnncla (Dusbabek and Lukoschus, 1973) contains L (known from seven females and three tritonymphs) from a Surinam Mbnou crenukidtm. Each of these niyobiids has been taken in association with an individual phyllostoniatid bat species. The reccm recovery of a female E. viguermi (ident¬ ified by Dr. E, VV. Jameson, Jr.) from Aniheus Jantaicensts from Veracruz, Mexico, suggests a recent connection between tlie mainland A. jatnaicctisis and insular A. Jamaicensis on Cuba. A chcyletoid ancestry for the Myobiidae was proposed by Dusbabek (1969) who also suggested a close phylogenetic affinity among EudushahrkUi, Ewingana, and UgLifuiohia, all found on bats. Species of Ewingana are parasites principally of molossids found in the Old and New Worlds, Eyvingana ttiolossi Dusbabek from Mifiossits /noic^ssus and E. yuguajayetisis Dusbabek from Tadarkla kukan- (iaicu both from Cuba, may indicate a host link between the Old and New W'orlds. Evidence for a vespertilionid transport system may be seen in the relation¬ ship between Old World Neomyobki Radford and species of vespertilionid and leaf-nosed bats. Ncon}yi>hia durnpieralijy (Michael) has been found on Pipis- ftellns pipistndkis, Epicsictts tiilssonii, and Rhinok^pkns hipposideros in Europe and several other Neoniyohia species are reported from European, African, and Asian RkifUfloplius species. I’hese associations may parallel those between vespertilionids and phyllostomatids in the Neotropics. Additionally, several species of Pieracarus are known from the Old and New W'orlds from vespertil¬ ionids and one, P. chalkudohus (Womersley), is known from Australia, North America, and Czechoslovakia (Anciaux de Faveaux, 1971). A New World origin is suggested for species of EtfdiLshahekia specifically, but the bat-infesting myobiids in general probably arose in the Old World, as there is seemingly less taxonomic differentiation and host specificity exhibited by the myobiids on New World chiroptcrans when compared to their Old World counterparts. Demodici[)ae Nicolet, 1855 Three species of demodicid mites {Dv/)u>dex}, all from Surinam, are known from Phyllosiotnii^s husfaius {D. phylloslonuuis L.eydig, 1859) and Carodki pcrspicilliiut {D. carolliae Desch c/ «/., 1971 and D, lofigis?;i/}ius Desch e( ak, 1972). It seems likely that many other chiroptcrans harbor Demodex mites, as only these three phyllostomatids, six vespertilionids, and a molossid arc recorded hosts (Anciaux de Faveaux, 1971; Desch e/ a/., 1972; Fain, I960; Lukoschus 1972). A commcnsalistic cheyletoid ancestor was hypothesized (Nutting, 1964, 1965) for demodicid mites, the intermediate, less specialized forms of w'hich are exemplified today by species of Sf{>nuiU}dix Fain and Rftinodex haeti Fain, Both SfomcKodex gakigoefists Fain and /?. haeti occupy intraoral and nasal cavi¬ ties of Galago sc tie gale ns is, a lorisid primate, whereas the other three Stomaktdex species inhabit the same microhabitat of a pteropodid, a nyctcrid, and several BiOLOGV OF THE PHYTLOSTOMA'rfDAE vespertilionid bai species. Because they are only slightly modified morphological¬ ly, it seems that these mites invaded the relatively stable nasal and oral cavities early in demodicid phylogeny and have changed little since. The association of more primitive Stoniniodex species with five bat species that range from Central Africa to Europe suggests a comparatively long affiliation by demodicids with chiropterans. The origins then of Dc/nodex may have been with bat hosts in the Old World. Vespertilionids are the most numcroirs and widespread ho.st bats of Deuiodi-’x species and are recorded from the Old and New Worlds (Anciaux de Faveaux, 1971). The tumors or small papules on the skin from which the phyllostomatid- infesting Demodex were recovered may be tissue reactions, suggesting a recent incorporation of leaf-nosed bats as hosts, perhaps by demodicids previously associated with vespertilionid bats. Nutting (1964:443) has expressed doubts about recent interspecific transfer. He further staled that phylogenetic palierns and species specificity are indeterminable, PsoRERGATiDAF. Dubinin, 1955 Three of the I 1 species of Psorerf^atoides Fain are presently known (Liiko- schus et a!., 1973} to infest phyllostomaiids— -P. lanchorhinae from Lffnehorhina aitrira from Venezuela, and P. glossop/uii^ae from Glossopfuiga soricinu and r. artihei from AriiheKs Uiu/aius, both from Surinam. In the Old World, the species of this genus occur on rhinolophid and hipposiderid bats throughout Europe and into Africa, one species is found on vespertilionids in Africa and another species was taken from molossids in Surinam, which may indicate sonic evidence for a vespertilionid or rnolossid host link between the Old and New Worlds. The Psorergatoides species are intradermal inhabitants of the cars and wings of their hosts, apparently feeding on dermal tissues. Host tissue reaction has been observed and discussed (Lukoschus e! «/., 1973), and is especially pro¬ nounced in the phyllostomatid hosts of P. glossophagui' and P. anihei, and in P. mofossi (found on Molossus molassus), perhaps suggestive of a relatively recent invasion of these hosts. This possibility and the fact that P. rhinolophi Fain has adapted to nasal mem bran e.s, wings, and ears, and is widespread on eight species of rhinolophid bats in Europe and Africa may suggest an Old W'orld origin for the group. Nutting (1965) noted the possibility that psorergalids arose from a cheyletoid ancestor common with the basal stock that produced the demodicids, myobiids, and several other families parasitic on vertebrates. The Psorergamide.s group shows close affinities with the species of Psorergaics Tyrrell, which are restrict¬ ed to nonehiroplcran mammals. Movement of P.vo/w^e^^ire.v-like species onto bats from rodents or other small mammals may account for the seemingly strong similarities between species of the two mite genera. It is also feasible that the Psitrergaioidcs laxa evolved along with the chiropterans from insectivore-para- sitizing ancestors as there are many species of Psorergaies known from extant species of insectivores. However, this seems unlikely as one w^ould expect to find ■SPECIAI, PUIJ1.1CAT[()NS MUSRUM t EXAS TECH UNIVERSITY 6fi the psorergatids more widely distributed on bat hosts and greater differences between species of Psorcrefutes iind Psotrrgaii^ides. Trombscui lOAE Ew'ing, 1944 Tronibiculid mites (including Leeuw'enhoekiinae) are parasitic on many kinds of vertebrates, including bats. More than 1500 species are known from temperate and tropical regions of the world. The larval stage, commonly called a chigger, is a frequent ectoparasite and occa-sionally an endoparasite of phyb lostomatids. Most of the tronibiculid literature concerning bats can be found in Anciaux de Faveaux (1971). Several regional papers include those on Mexico (Loomis, 1969), Panama (Brcnnen and Yunker, 1966), Surinam (Brennen and Lukoschus, 1971; Brennan and van Bronswijk, 1975), Trinidad (Brennan and Jones, 1960), and Venezuela (Brennan and Reed, 1973, 1974, 1975; Reed and Brennen, 1975). Major taxonomic papers deal with the genera Bca/ficrflla, Hooperclki, and I eatnuifiafki (Vercamnien-Cirandjean, 1967), Chiropiclla and Lt^pk/irombidinni (Vercammcn-Graiidjcan and Langston, 1971), LtH^ntisia (Brennan and Reed, 1972), MicRXromhlcula (Webb and Loomis, 1971), Nyaennastes (Brennan and Reed, 1973), Parusecia (Brennan, 1969ii), and Pertssopalla (Brennan, 19696, 1970). t he generic status of certain taxa has been questioned by Ver- cammen-GrandJean et ciL (1973); however, they are recognized as subgenera. riiere are 51 species, belonging to 22 genera of two subfamilies (Trombi- culinae and Leeinvenhoekiinae) known from phyllostomatids. Records of three species seem to be based on accidentals or errors in handling— lilankaania .si/imimaryi normally parasitizes birds, P.seiHioscfuH'fii^asfia hklhifera is abundant on small mammals, especially rodents, and Trtfnthicula dutitii is known from a variety of terrestrial mammals (Brennan and Yunker, 1966). Occasional and possibly accidental records include seven species of the abundant and wide¬ spread genus Eufmnthiada (Brennan and Reed, 1974), and LepunromhuHitm hafnaviaiiinj known to regularly infest rodents and rabbits. Xenodoniacanis serratus Loomis and Goff (1973) was described from a single larva taken from a Mexican Aniheiis lifnraitts, the only record of a Xctiikioniacarus from a bat. The remaining 40 trombiculids regularly infest bats of the Americas and are documented from phyllostomatids. These consist of species in the genera Aiexfai/iia (one of two knowm species), fii’antendla (two of three), Chiropiella (one of three American species), Hoopvndia (three of four), Loomisia (five of six ), Mivroirnnihkiila (three of about 20 American species), Naxicola (one species), Nycte/inasies (two species), ParascoscfuH^/jgasiia (two of five), Para- secia (three of 12), Perntes (one of two), Perisstypalla (six of 12), Speleocoia (two of five), Tevofuadcimi (two), Wayenmiria (one), Wharlonia (four of eight American species), and Xenodonfacarifs iyf four). The usual life cycle consists of the egg, deiitovum, a parasitic larva (which attaches to the host, feeds on lymph and histolyzed tissue, and then leaves the host after engorgement), inactive proto nymph, active and predaceous BIOLOGY Oh THE PHYLLOSTOMATIDAE 67 deulonymph, inactive tritonymph, and the figure-cight’Shaped predaceous adult, either male or female. The parasitic larv al stage is relatively brief, whereas the free-living nymphs and adults must have the proper substrate and prey. A good host will pick up the larv'a, provide a favorable parasitopc and nourish¬ ment, and deliver it after engorgement to a suitable drop-off site, frequently the original or a similar niicrohabitat in w hich the larva w'as picked up. rhe larvae usually attach singly or in clusters on or in the cars (Micro- irombicttUif Spelcocohiy Iccoouukuia, and Xenodonfacurus), wings and tail membranes (Bcamerclla, Chiropieflay Hooperelki, Loooiisia, PerissopaUa^ HcraicSy Farasccki, W'harfonkh and some Tcco/natUifui and Farascttscfiocn- ^iptsiki), toes (Fenites), and nasal passages (Alcxfuiniu and Nasicola). Chiggers are occasionally found on the head, lips, body area, and the genitalia. The en¬ larged larva is oval in shape, rarely larger than one millimeter in length, and may be red {lica/nerella^ Hooperelki, Tcconiatlami^ Wkanofiiak orange [Peris- sopalla, LtHmiisia), yellow- ( Chiroptelki^ Nasicola, Farasccia, Feraics), to w-hitish ( Spe! coco I a. Paras ec ia ). Emergence of the unfed larvae of most temperate and many tropical chiggers is seasonal, cither correlated w ith temperature in alternately hot and cool regions, or synchronized with wet or dry' periods. Modifications in larval morphology that seem correlated w-ith chiropteran hosts include greater sclerotization of legs and palpi, projections on certain leg and palpal segments (Vercammen-Grandjean, 1967), and enlarged and serrated cheliceral blades on those that attach externally (but usually there is a moderate blade on those that attach w-ithin ears and small blades on intranasal species). The oval-shaped larvae may be red or orange in larger ectoparasites and yellow- or nearly white in those that normally attach deep in the ears or are free in the nasal passage. Virtually all of the seven most frequently parasitized phyllostomatids (Arii- heas, Carol I ia, Dcsmodiis, Clossophaga, Macrofas, Micronycteris, and Fhyl- losiooiKs), as well as other heavily infested tropical American bats, such as Balantiopfcryx plicata, Moroioops, and Pteronotas parmdiii, are regular or wholly inhabitants of caves and rock crevices. 'I"he remaining host phyl¬ lostomatids normally roost in hollow- trees. In addition, nearly all of these hosts usually roost in clusters or large colonies. Although most free-living stages of these and similar genera are known only from laboratory-reared material, most if not all of those listed probably inhabit cracks in rocks or decaying wood (frass). Closely related species in the genera Parasccia and Microtrofahiaila are known to inhabit decaying logs, stumps, and dead .standing trees. Other Microrromhicida and Wharionia are associated closely with rock crevices in cliffs and caves. Bats taken in caves and mines w'cre infested heavily with larvae of the genera Bcamerclki, Hoopcrcka, Lo- omisia, Microtromhicula, Perissopaila, Speieocokiy Tccomatlana, and Wharumia, Bats recovered from bridges, houses, and other artificial structures rarely possess trombicuHds, nor are they found on bats that usually or alw^ays roost among leaves on living branches of trees. NPECIA[- J>UIU.JC A i JONS MUSEUM 1 EXAS TEC’J-J UNlVbKSi l V 6K None of the well known genera and few of the species have been recorded only from phyllosiomatids, although most of them are known only from bats, including emballonurids. molossids, monnoopids, and vespertilionids. Genera associated with bats and of probable Neotropical origin consist of Nasicola and Spcleovoia (members of a world-wide group including afki)y L(H)t}iisia (a distinct group), Pcrissopalla (possibly related to Old World genera including Ricilliiua and Irisciira, according to Vereammen-Grandjean e! t;/., I 973), and the distinct American irombiculine laxa consisting of Bcamctrl- ia, Hooperelku TcconiatUina, Alej/ainku Nyaerifuistes, and Perates. Leeiiwenhoekiine genera consist of ft'c/ee/mr/r/V/ (closely resembling the genus Sasavani.s, which is abundant on desert and tropical American terrestrial mam¬ mals); Xt^tunkf/ifavarus sernims, one of four known species in a group regularly on small terrestrial mammals; and Wluuftfnici, which is world-wide in distribution and found on many different kinds of bats. Wharti^nki tmdoscKhsa and W\ pachywkanoni represent typical species and are restricted to the New' World tropics, whereas Wharfoniu and fU. ytfan’/etisis arc mostly northern Neotropical. These Wharumia also have been found on a w ide variety of Ameri¬ can bats including emballonurids, molossids, and mormoopids, and all American genera of Iccuwenhockiincs seem to be northern in origin and mostly northern Neotropical in distribution, liats, including a number of phyllostomalids, are common hosts of Para- m}sviuH’H}*as}(a and Parasaia, which seem to be acquired by liosts in holknv trees and other rot)siing sites associated with decaying wotxi. Paranamiiocn- aatuiiafa and P. ntc%>asTyraA are recorded only from bats, as is Parasevia sanaatyatifi; however, Parasecia lon,ific(iIcar and P. niannep also are known from a number of other small jiiamnials and bird.s ( Brennan, 1 969^v). Rosi-NS i iziNiiDAi: Cooreman, 1954 Ihe subfamily Nyctcriglyphinae Fain consists of two genera and 13 species (Anciaux dc Faveaux, 1971) associated with bats or bat roosts. Only one of these, Nyc{eri}>lypftiis stiintirai" Fain (1963), based on a single tritonymph, has been found in association with a phyllostomatid (a Brazilian Sutrttira liliitm). Dusbabek (1967h) subsequently reported a male and female N, sfarnkae from Mol(K\siis aiolossiis Uiken on Isla dc Pinos, Cuba. LahioocARP tDAE Guntlicr, 1942 The family Labidocarpidac consists of eight genera previously placed in the families Listrophoridae or Chirodiscidae. Species of four genera have been reported from phyllostomatids. Akihidocarpus furntani was listed from Trinidad (Pinichpongse, i963a) from Anottra peoffroyi and also from A. geojfroyi and Cla.ssopha^ut soricina taken in Nicaragua (McDaniel, 1970). Other Nicaraguan species include A. nkaraf^uac from Urodenna hiktbatuni and A. jonesi from ya/npyn^p.s /udleri Three species of ParakihkiocarpK.s' Pinichpongse have been recorded exclusively from phyllostomatids. Pundahidocarpus aridud Pinich¬ pongse (1963/9 was originally described from mites on Arfthcas {ifttroftts B[OLOGV OF 1 HE PHYjJ.OSTOMAT[DAE 69 from Trinidad and later reported (Tamsitt and Fox, I970r/) on A. and Sfc/K/ih^r/fUi r/tfi/ni taken in Puerto Rico. Subsequently, de la Cruz e/ a/, ( 1974) described f\ from S. nijii/fj and P, fb.x i from Anihens Jwfuticensis, both from Puerto Rico. LawrenceiKarpus nikropUiis Dusbabek and dc la Cruz was originally described from specimens obtained from the mornioopid Picro- uofus fitltuitioxus taken in Havana Province of Cuba (McDaniel, 1970). A later record of L, miempilus was noted by Tamsitt and Fox (t970«) from Brachy- phylla cavenuuiifii collected in Puerto Rico, which were also the host and locality records for the species L. de la Cruz, Tamsitt, and Val- divicso (1974). Fain (1970 /j) has reported some recent records from Surinam phyllostomatids and McDaniel (1972) reported numerous labtdocarpids from Venezuelan leaf-nosed species, Labidocarpids have anterior appendages specialized for clasping individual hairs in the fur of their mammalian hosts. Food habits have not been noted, but, as with other listrophoroids, they probably feed on dermal tissues and se¬ cretions at the base of the hair. The life cycle includes a hexapod larval stage, which after parturition may molt either into a nyniphal male or female. The nymphal male molts again to become an adult, whereas the nyniphal female undergoes another molt in which the eedysiurn remains around her, thus forming a puparium or chrysalis. During mating, the male clasps a hair while a female is attached to his posterior end III copuh) with the anterior end of each facing in opposite directions. How the male and copulatory female initially unite is unclear as the legs of the pupar¬ ium have no apparent morphological adaptive quality for holding fast to hair or skin and, before pupal formation, were of little locomotive value to the nymph because of their diminutive size. Furthermore, the fate of the copulatory female after disengagement from the male is questionable for the same reasons. The “three-legged,” fully chiiinized stage w'ith the next stage developing inside as seen by Law'rence (1952:137) was possibly a representative of a separate species in wliich the short or nearly absent legs of the nymphal female are character¬ istic of that taxon. Ovovivipurity is seen in these mites tor fully formed larvae have been observed inside normal females (McDaniel, 1970) and mature fe¬ males give birth shortly after shedding the pupal skin. The genera LahkiocarpHS, Aiahidocarpus, and Okihidocarpus, w-hich para¬ sitize phyllostomatids, have both New and Old World species. All three also have representatives on vespertilionid bats either in the New' World {Labkio- tarpus) or in both the New and Old Worlds (the other two genera). The relatively greater differences among the seven genera in Mexico, Central America, or South America suggest the site of origin was in the New' World. Only a single monotypic genus is know-n from the Old World. Furthermore, phyllostomatids harbor the greatest number of labidocarpld species, suggesting a relatively long association. Additional evidence for the initial establishment of labidocarpids on phyl- iostomalids may be implied in the Old and New' World distribution of the mite species. If they had invaded the Old World rhinolophids or hipposiderids or 70 Sl’EClAl. PUfiJ [CATIONS MUSEUM TEXAS TECH UNlVERSf l’Y vespcriiJionids of cither realm in the beginning, we would expect to see a much broader range of these host species, something that is not in evidence. If original¬ ly on New World molossids, then why are they not found on Old World free- tailed bats? T he accumulated evidence seems to indicate a New W'orld origin for labidocarpid mites on phyllostomatid bats with radiation to the molossids, noctilionids, and vespertilionids followed by dissemination to the Old World rhinolophids and hipposiderids via the cosmopolitan vcspcrlilionid species. CniRORiiYNcnoiuiDAE Kain, 1967 This sarcoptiform family consists of two species known only from phylloslo- maiids. Chhorhyitchohia {(nHierniac Fain (1967) was based on a single female from a Panamanian UroJtrfna hilohmuni, and the seven females of C mat son i Yunker (1970) were attached by their mouthparls to the trailing edge of the wing membrane of a single Anottra i^eoffroyt netted in Zulia, V^cnezuela, Sarcopfidae Troiiessart, 1892 One genus of sarcoplid mite is known from phyllostomatid bats. Chirnys- soides was proposed by Pain (1959) to include C. capani recovered from Arti- bens Jamaicet}st\\ C. nmazonae from Ca/ollia perspU illnfa, C. hrasilie}isis from Stnniira lilinni (all from Brazil), and C vetieznelae from a Venezuelan 7b- natia vetiezneiae. Fain (1962) later described C candiiae from Panamanian Carollia perspicillata and C. vastatiea. Only phyllostoniatids were known to be the hosts for Cbif/ryssoides until Fain and Lukoschus (l971/>) transferred Notoedres ni>ctiliofiis to Cbietiyssoides and erected a new' subgenus (M>c7/Y;‘- oc(}pfes} for it. Chirnyssoides /loaiiiom’s is known from Cuba and Surinam from noctilionki bats. Two other species, C. snrituimensis and C. zafideryensis both taken from Surinam Caroilia pe/spk idaia were placed in a separate sub¬ genus, Ca r(} II icopt es. Known parasitopes for Chirnyssoides species arc the skin of the leading or trailing edges of w ings or ears. Ihc Chirnysseddes species group may have arisen from stock common to that including the species of Noitn’dres. Certain evidence suggests an Old W'orld origin for the No toed res group, possibly from an ancestral line that includes the genera Chirnyssns and Nyeteridtteoptes. Support for this idea seems to be implied by several things. There arc many more species of Noii>edres in the Old World and they are found on a greater variety of bat hosts. A number of species of Nyeteridoeoptes and C/iirnyssns are inhabitants of the biical cavity in some bats, a condition that may be seen with other groups of aearines that have long associations w itli their hosts. Nyeterhhteoptes and Noteodres have representatives on pieropodids, rhinolophids, hipposiderids, molossids, and vespertilionids in the Old World and numerous species are known from vespertilionids in the New W orld. The developmental sequence in the phylogeny of these mites seems to indicate an early establishment on pieropodids followed by a movement onto rhinolophids and then to vespertilionids, which carried them to the New World to phyllostoniatids. HlOL,OGY OF THE PHVLLDSTOMATJDAE 71 GaS[ronyssidae Fain, 1956 FhyiiasfottHmyssus cofuadyunki'n Fain ()970f) of the subfamily Rod- hainyssinae Fain, is the only gasironyssjd reported from phyllostomatids. It has been found in the nasal passages of Venezuelan Aniheus liiumat.s and Uro- (Inma bilohaiuf}}, Vampyrops helleri, and A. iintrcuiis from Surinam (Fain and Lukoschus, 1972). LeaFnosed bats may have acquired this mite from associ¬ ations with vespertilionids. I’hc few additional reports of rodhainyssines from the New World include the vespertilionid Hisiiotus vekims from Brazil and Chile, the molossid Eufjiops uhrasits from Surinam, and the emballonurid 8^ll- (lunopii^ryA plicala from Mexico (Anciaux de Faveaux, 1971), All were infested with species of Rsis also tended to have more streblids. It was further suggested that within the host’s preferred habitat, however, host specificity was great even though the same streblid species may be found on a number of other species of bats. Earl¬ ier, Ross (1961) Slated the same situation for several Nearactic phylloslomatid and two vesperiiliunid species. Hts observation of the apparent non infestation of molossids cohabiting with Infested vespertilionid and leaf-nosed bats may be due to recent streblid dispersal to these temperate regitms as a number of molossids serve as hosts for batflies in the tropics (Jobling, 1949; Wenzel, 1970; Wenzel c/ 1966). Although Jobling (1949) stated that streblids probably arose from an ances¬ tor that was not blood sucking, it seems more likely that they originated from hematophagous, calypteraie, muscoid tlies as Theodor (19,'i7) has postulated, and that the adaptation was originally to bats. I heodor (1957) further staled that because of a complete lack of streblid fossils no conclusions about the evolution and phylogeny could be drawn regard¬ ing battlics and their hosts. Certain observations may be made, however. Kor instance, the greatest differentiation of species (94) and genera (23) has occurred in the New' World as compared to 62 species and 4 genera of the Old W'orld (Wenzel, 1970). Indicative of this great taxonomic range is the spectrum of morphological features of New World streblids from the generalized ealypterate taxa to the small-winged and nigiillcss Hies (certain species of Sfn'hia, for ex¬ ample) and to the w ingless species of ParaJyachirki. Further rcllections of adap¬ tations by Neotropical battlics may be seen in the polyctenoid appearance of Si rtf h la and other species and in the siphonapteroid NycurophUia. These diversifications seem to indicate a New' World origin for streblid battlics. The endoparasitic mode of AsvtHlipn’ton on Old World bats may merely rcllect an adaptation to competitive factors w ith other celoparasiles, such as nycteribiids. Additional evidence for New World origins may be seen in the hyper- parasitic relationship between certain streblids and the trombidiid mite, M(fnunguis sirt'hUtia, first noted by Wharton (1 93H) as an ectoparasite on streb¬ lids— Mi'gi.stopcxhi arantai (Coquilleit) and I'rkhtfhitfS tlugesii Townsend— from caves in Yucatan, Mexico. From other material obtained from California State University, Long Beach, more specimens of M. sirehliiia have been recov¬ ered from (identified by Dr. B. V'. Peterson) taken from Macroius waierhousii from Sinaloa, Mexico. We assume the strebiId-bat association occurred before the mite-tly relation¬ ship, and M, sirehlkla probably encountered the streblids on the cave Boor where the parasitic tlies emerged from the puparia. It would seem probable that the mile is a relatively recent parasite on streblids as it does not exhibit host specificity or selectivity. It is probably also more closely associated with a suit¬ able cave environment, large colonies of bats, and adequate populations of suitable streblids. BIOIJXIV OK THE HHYl.LOSTOMAI IDAE 73 Other published papers regarding streblids include those of Guimaraes (1944), Peterson and Hurka (1974), Peterson and Ross (1972), Reddell (1970), Star¬ red and de la Torre (1964), and Whitaker and Eslerla (1975). Discussion Ectoparasites are recorded from 39 of 49 genera (89 of 136 species) of phyl- lostoniatids. There are two general host-parasite categories: either the parasite remains on the host throughout the entire life cycle (for example, demodicids, my obi ids, and sp into rnic ids), or it spends part of the cycle off the host in the bat roost (for example, argasids. ixodids, macronyssids, and irombiculids) and may parasitize a variety of bats and even other vertebrates that trespass into its ter¬ ritory. Nyaeris^iypfitts snitnime, a rosensteiniid mite, is a commensal with two species of leaf-nosed bats. For part-time ectoparasites, the host is visited at least once, and sometimes two to three lintes. This type moves onto the available host, attaches to some part of the body, and commences feeding. Tlie feeding site is termed the para- sitope (Fain and Vcrcammen-Grandjean, 1953; Wrenn and Loomis, 1967) and seems to be selected by the parasite although it may be influenced by grooming or scratching behavior of the host. The microbiotope refers to the area of normal activity of each parasite, which may be on the entire surface of the host for battljes, or in a single dermal pore by dcmodicid mites. Others, such as soft ticks and macronyssids, also must sur¬ vive in micro habitats off the host. The parasitope and microbiotope may be virtually the same for endophilic demodicid, sarcoptid, and psorergatld mites. Myobiids and labidocarpids have niicrobioiopes where they clasp individual hairs, and their feeding parasitope is visited pcricKiically at the base of the hairs, Spinlurnicid miles and the battlies have extensive microbiotopes. Wing mites usually are found on the wing and leg membranes and rarely invade the fur, whereas batllies move over much of the body surface. Argasids and most tromhiculids find and climb onto the host and move quickly to feeding sites, so the parasitope and biotope are essentially the same. They remain at these feeding sites and on the host for relatively brief periods. Gastronyssids, ereyneiids, and some trombiculids occupy the respiratory passages, and certain macronyssids ( Rmifordiclla) are found in peridontal tis¬ sues. The females of chirorhyiichobiids have been recovered from w ing mem- branes and spelaeorhynchids w^ere embedded in the skin of the ear. Three feeding categories are suggested for these ectoparasites: hematophagy, histophagy, and mucophagy. The hematophagous soft and hard ticks, the two families of batllies, spinturnicids, and some macronyssids derive most if not all of their nourishment from blood meals. The remaining parasites, except for intranasal and iniraoral taxa, feed on dermal tissues, fluids, or skin secretions. Those inhabiting oral and respiratory cavities apparently feed on mucus, although histophagy also has been suggested. Protonymphs of several species of Rmi- fonitella feed on tissues surrounding teeth, resulting in extensive damage in some instances. 74 SE^ECJM. I>U in. I CAT IONS MUSEUM t EXAS' TECH UNIVERSITY The highest degree of niche niid host specialization occurs in ectoparasites of birds and bats and is attributed to their relative ecologic and geographic isola^ tion (Wenzel and Tipton, 1966). Furthermore, the degree of host specificity is correlated with the extent to which a parasite is host limited. Examples of familial specificity of ectoparasites include Slreblidae, Nycteribiidae, Polyctenidac, and Spinturnicidae and certain trombiculid species exhibit host-species restriction. Howe%'er, most literature on phyllostomatid ectoparasites does not contain carefully documented information about host-parasite relationships. An approach to bypass the shortcomings of a hosl-para.site list is to find two or more separate studies listing a particular parasite from a specific bat species or other taxa. On this basis, it appears that few phyllostomatid infestors are monoxenous. Ex¬ amples of moiioxeny, however, include Peri^iischnis herrerai taken only from Dt'S/uodns rotsituiiLS and Speteachir p/iyUosKnjii recovered solely from Phyllosto- tfuis futstaiKs, both of which corroborate Wenzel and Tipton’s (1966) idea of higher host specificity in host-limited taxa. The usual hosts for several strcblids appear to be AniheusJanuikefisis for Mes^i.siopoi/a amitca, Gi(>ss(/phat,Hi soricina for Trkiufhius dnge.v//, Carolfh perspicillafa for 7’. johli/igi, PhyllosKmufs hastaiiis for T. longipes and P. kiscalor for Strchla herti^i. Macrr accitlenuif (.x;eurrence I I 4 3 ■y 1 3 4 I I I 3 la [ 4 3 1 * 4 4 ectoparasites, most of them not host-limited. The extensive infestation of these bats may be explained by their colonial habits and their practice of cohabiting with other species of bats, sometimes, as w'ith C pt'rspiciHma, actually mixed with colonies of other bat species or in the same roost w'ith other cavity-inhabit¬ ing bats (Pine, 1972). Otrollia perspiciilata may be the focus of infestation for these and other bat species through cohabitation as it is probably the most wide¬ spread and abundant species of colonial fruit-eating hat in the Ncotropics. Fifty- eight species of ectoparasites are recorded form C per.spicilUmt, 18 of which are shared entirely or partially w'ith /(. Jamaicensi.% G. soricina, and D. roiundtts. Although colonial, Lt’pitfnycferis and Choeronycferis harbor few' ectoparasites, probably because of their migratory habits as they are flow-er feeders and migrate to stay in the “dry season" (Humphrey and Bonaccorso, this volume), extending into temperate regions where there are fewer parasitic arthropods. lb SPECIAL PUBl tCAf lONS MUSEUM TEXAS TEXH UNIVEBSKA Most of the phyllosio mat id-infesting families of ectoparasites seemingly originated in the Old World; many probably wore transported by vespertiiionid bats to the New W orld where transfer to phyllostomatids and other bats occurred. Evidence for this mode of exchange may be seen in the macronyssids, spintur- nicids, myobiids, sarcoptids, gastronyssids, and nyctcribiids, Molossids may have been important for dissemination of the psorergaiids from the Old World al¬ though vespenilionids once again arc inferred strongly as the transport mechan¬ ism. V'espertiJionids are likewise the major possible means of early dispersal to the New World for demodicids and ereyneiids even though rt)cients, especially murids in the case of the ereynetids, may also have been significant transporters. Either rodent or avian hosts, or both, provided the ways and means for move¬ ment of the argasids to the New World. Emballoniirids seem to have been important in older parasite transfer between the Old and New' Worlds, parti¬ cularly in the temperate climates and especially with the trombiculids and gastro- nyssids. The families Slreblidae, l.abidi)carpidae, Spelaeorhynchidac, and prob¬ ably Chirnrhynchobiidae apparently arose in the New W'orld. Sec Table 2. Present day geographic distribution of certain ectoparasites possibly may be explained by the effects of continental proximity during early geologic limes (Traub, 1972). Generally, however, today’s geographic placement of nearly all families of phyllostomatid ectoparasites may be explained by Palaearctic migra- tional patterns of vespertiiionid bats in late Cenozoic times or by over-water migration on birds of long-distance llight. Continental drift separated the land masses of Laurasia, Africa, and South America between 70 and 10.5 million years ago during the Cretaceous {CTacraft, 1974), forming water barriers to chiropteran and other vertebrate migrations in later periods. The earliest bat fossil from the early Eocene of Wyoming (Jcpsen, 1966) is similar to extant microchiropterans. Even if bats had existed as early as the Palcoccne (Vaughan, 1972), the oceanic gaps between continents still would have been a restrictive boundary to bats as most species generally do not traverse even small expanses of salt water. Movement of tropical lowland mammal hosts, especially of rodents, and their parasites between South and Middle America apparently has been with¬ out great obstacles (Wenzel, 1972). Ectoparasites reflect the extensive range of a number of leaf-nosed species, several recorded from Mexico to Peru and Brazil, and a free exchange between Mexican and South American tropical species may be scon with several examples: the streblid Meii'tsiapoda anmea found on Ani~ hens janmicefists collected from Mexico, Central America, and northern South America; the spinturnieid Pcri^Hsiiints iheriiigi from A. jafmtkensis from Mexico, several Caribbean islands, and Venezuela; and the irombiculid Loomiski desmodus from Giossophapa soricina recovered from Mexico, Nicara¬ gua, Panama, and northeastern South America. Others show an interchange between Central and South America, for example, Tfichobius jobtingi (streblid) from five Central American countries, Trinidad, 1 obago, and northern South America. Because phyllostomatids are principally tropical species, the adjacent temperate climates represent barriers to them and their ectoparasites. The HJOLOGV OF THE PHYl.LOSTOMATIDAE 77 Taki 1 . 2 .—M'li’ World btu fomtlies and thetr ectopantsiffs. A single osierisk huikutes a pndHihle itiridinkii rt'cord; a dtfuhie as!vrisk indkafi's u cofiimcnsal firtoip: a/id a (tiplv ifSK'ri.sk ifidictHes u sit!}ile record only. Parasitic yroup* a "3 3 C 0 "rt £ E LU Cl n 5 o o 'Z rt [3 E o % £ 7i c. 0 c e u O 3 'rt H 7, ij- S G. O >s £ c o V fX > n "3 ‘7i o o 3 Acai'idae X X Anoetidae X Argasidae X X X X X X X Chi rorhy ncho hi idac X Demodicidae X X X2 Ereynesidae X X Castronyssidac X X X X Glycyphagidat' X Ixod idae X* Labidocarpi*^!^^^ X-ii ■ X X X X Laelapidat' X Macronyssidac X X X X X X Myobiidac X' X X X X Psorergatidae X X X'= x« Pyroglyphidae X Ro.sensieiniidac** X X X Sarcoplidae X x^ X X X Spel aeorh y nch j dae X X SpinUirnicidae X X X X X X Trombiculidae X X« X X XH X X Nycteribiidac**- X X = " X X Sireblidae' ^ X X X X X X X X Cimicidae'- X X X Polyclenidae''* X X Siphonaptera'^- X X 1, kewrds iire fmni Ancsjuis dc Faveaux. 1971, unless oi her wise indlealed; 2 , Desch ff It!., 1972: .L Fain. 1970.; 4. .VlcCJanie!. 1970. 1972; 5, Dusbahek and l.n knschns. 197l«; 6, Liikoschus ft til.. 1975; 7, Kiiin and Liikttschns, 1^71 1 'j; K, unpuhlisihett records oC Loomis; Guimaracs, IV46; 10. Ciutmarac's and D'Andrella. lA'Sfi; U. Wen^d t‘! nt., l'#66: 12. Usingcr. I?. Ueshima, l^'72; 14. Tipion and Mcndcjf. 196(1; 15, Tipion and Mach ado-Allison. 1972, lemperate ccloparasitic fauna on bats, such as on species of the Myotis ni^^ricans complex, shows little restriction to the temperate areas in the tropics in contrast to the rodent parasites (Wenzel, 1972), which is to be expected because of the volant nature of their hosts. The distribution and numbers of the ectoparasitic taxa on phyllostomatid species may give some information regarding the relative duration of host-para¬ site associations. For instance, if one or two genera comprising only a few species of parasites are generally widespread on leaf-nosed bats, as in the case of the demodicids and psorergatids, then it may be assumed that the phyllostomatid- parasile relationship is a relatively recent one. In those with many species and genera, for example, macronyssids and labidocarpids, a longer period of as- 78 SPEC]A[- PUin.tCATlONS MUSEUM TEXAS TEOt UNIVERSITY sociation is suggested- If the ectoparasites arc represented by only a few genera containing numerous species, they probably have been affiliated for an inter¬ mediate period of time. Based on this interpretation, it appears that the streblid battlies with 20 genera and 82 species have had the longest association with phyllostomatid bats. Fleas, polyetenids, and cimicids do not appear to be normal parasites of phyl- lostomaiids, nor are they found regularly on Neotropical species of furipterids, cmballonurids, mornioopids, natalids, noctilionids, or thyropterids. However, polyctenids arc found almost exclusively on molossids and there are a few species of Heas and cimicids on the Neotropical members of the Molossidae and Ves- pcrtilionidae. These asstx;iations may suggest a recent arrival of these parasites from the Old W'orld on members of these two advanced and widespread groups. Parasite-Host List Ectoparasites, and hosts for each, are listed alphabetically by genus and spe¬ cies. A single asterisck indicates an unpublished record from the Chigger Lab¬ oratory, California State University, Long Beach; two asterisks indicate an un¬ published record from The Museuni, Texas Fech University, Lubbock. Geo- graphic origin of records is given if known. Since this list was prepared, four publications (Brennan and van Bronswijk, 1975; Brennan and Reed, 1975; Her¬ rin and Yunker, 1975; Reed and Brennan, 1975) have appeared that should be consulted for additional records. Aliibidociirpuh Funiiiiiii Hinichpongsc (Laiudoc Ammrti i tiuiiift'r, V'enezuelH Amjuni Mcojfroyi, Nicaragua and Trinidad Ciif'ifUut hiiH'knuJii, Venezuela per.'tpirUtaiii, Venezuela (J!t/SM/phtt}>a lottfiiroMris, Venezuela CHosMiphtiiiid soritifuh Nicaragua Vonipyraps .hvfU'ri, Venezuela Atabidocarpu.s Kuyaiieiisi!> Fain Ariiheiis cincn u,s, Surinam AiabidocarpusjonesiMcDaniel Vtinipyrops helli'ti, Nicaragua and Venezuela Alabiduearpu.s nicaraguae McDaniel Urodertnn hiiobtituni, Nicaragua Urodcrnui ftiafiftirosirum^ Venezuela Ala bid near pu.s phylloslnmi Fain Phyllitstonmx hastaiiix, Surinam Alexfairiia chilonycteris Yunker and Jones (Tromiucui lOAr.) Ctiroitici perspicilldUu Panama Amhlyiinima sp. (t\(iDiD.\K) Ardheuxjiinniicettsis, Venezuela Arriheus (iturutii.y Venezuela Cnrollia hrevkauda, Venezuela Cttrolfki perspickiddu Venezuela ChiriHienua viUosu/n, Venezuela C!u)eto/iisciiS inortifr, Venezuela G!osM>phiigd lofigirostrfs^ Venezuela BlOl.OGV OF THE PHVLLOSTOMA'KDAE 79 (jlossifphat’d 'iiu'icifith Venezuela Slur/Iins (ifitui‘, Venezuela Unuli’rftui hihhiiluu)^ Venezuela Vtiiiipyntps fu'lleri, Venezuela Ainbljfomma lon^iru^re (Koch) Ariiheus lituruius, Venezuela Aiiastrebla nuittadeni Wenzel (Stheui in.\M) Afioura sp.. Venezuela Aututni vaudifen Colombia AfUfuru a ultraku Panama Atsoursi iriudjroyi, Panama and Venezuela Anaslrebta mode!»lini Wenzel Auouru s^uaJlfoyL Guatemala, Mexico, Panama, and Trinidad Aiiastrebla nyeteridix Wenzel Liuschophylla rtdfusiu, Panama Aiiafriehobius scor/ai Wenzel (Strehlidae) Lttuchopltylfu rohuMts^ Panama Aiilricola sp, (Aroasidae) Lvpiotsycii'ris cuiasoui\ Venezuela Antricola niar^iiiutiiA (Banks) fdiyfloisyaerfx ptwyi, Cuba Ajdricolii silvai Cei ny Phylfouyi icrix /?oeyj, Cuba Aspidoptera buAcki Coquillett (SrREKi.rOAE) Ariihs us sp.. Puerto Rico Ariihi'iisjiuiuiiccsissx, Colombia. Cuba, Guatemala. Mexico, and Panama Ariihestx iiiuruius, Panama Carol!ia pfrspiciUani, Panama CliinHh’f tusi vil fox Si flit Panama Fliylfoxtofiiits ilixcofor, Panama V(i/iipyresxu tiyttiphtseat Panama AxpidopJera ddatorrei Wenzel Carolliti perxpk'iUatu, Panama Siuruira filiumt Guatemala and Panama Aspidoptera pliyllostoinutis (Perly) Afufura Trinidad? A f lihi ux sp, Artfhcssx (isuriilfis Phylloxfoinitx sp., Brazil Sr (O ft fill I Hi ft lilt Paraguay Ela.silia sp. (Nve terhuidae) Cioiturio .vtuicA Basilia arilrozui (Townsend) Lcpfoisycferix sa>sh({>pti'rii\ tiuriiu.'^, Hrazil BasUiu myutis Curran IJriHleruhi hift)htiiii/}! Bus ilia roridanii Guimaracs and D'AnJrella Artityeii.s Janiaiccitsi^ ffyliniyfici'isuniirr'^votHii B n s i 1 i a spt' istT i (M, R i be i ro) A lit Him }:i‘tfj]hiyi Ctiroifia pct\spicil!iihL Hra/il PhyUt>.\ioiut(S sp. Basilia tiploni Guimaraes XfpitiHt Panama and Venezuela Basilia Guimaracs and D'Andrciia Ainhciis Viin'Anrd LiHichorhi/Ht itiiriut, Venezuela Beainerella actilaseuta Hrennan (Tromiiu ui in\M} Ciiralfia sp., Cosia Rica* Caiolliii pi rspii iUitiiK Niearajjua*, Panama, and Trinidad (ilos.sophayti .sortcimu Mexieu’*' Loui'hophylfa ciHu-tiViu Cosia Rica* Mitronyf/t'ri\ hif.MfUi, Panama and Trinidad Mii ronycn'i is Panama PhyUttMofiiii^ tliM olor, Nicaragua''' Be a me re] la .siibaciitaseula V cream men-Grandjean Miaonytwris liicsatii, Trinidad Biankaailia sinnamarji Floch and Fauran ( [ in ie>ae) f^liyfloMiHinis htisttiKiy Panama CJiirnyssoides sp. (Sahi oi> i m.\h) CViiro//n7 pi’i'itpii-iUuiiH Brazil Chiriiy.ssoides ania/oitae Fain Cat oil hi pi'r.spifilhtiii, Brazil Chiniy.SMriiles brasilierisis Fain Sf Iff film III in in, Brazil Cliirnyssiiides capadi Fain A f!(funis cinerciis, Panama Arfilu'its jtiKuiici'/isis, Brazil, Mexico*'*, and Panama Artfluufs (oliccns, Mexico** and Panama Clifi'oik‘1 tiiii Mifvitii, Panama Di'sittoihoi i-oHiniluy Panama Voinpyrewii pnsilht, Panama Viiiiipyrodi's turiia-ioloi, Panama Vaiitpyrops viihiim, Panama Chlniyssoides caredtiue Fain Clin (It in sp., Panama Ctirofliii ptuwpicillttun Panama and Surinam Oiroliia Mihnifti, Panama Gl(*ss((phiip(t soricithn Surinam M it I'lliiyi t r/7A iiu'piitoriu Surinam C'Jiirnyssdides surinuniensis Fain and Eukosebus Cctnifliii pt'rspit tHadi, Sunnum mOLOGV OF THb PHYl.LOSTOMATJDAE HI Chiniyssoides venc/uelue Fain Tofutn'a vi‘ftcznel{u\ Venezuela Cliiniys-soidcii zatiderjen'^ts Fain and Lukoschus Ciiroltia perspu illuiit^ Surinam Chiruecele-s totichopli>IJa Herrin and Radovsky (Mac ronyssidae) Loin hophyflu rohaski, Venezuela CInroptellu m> cips» Viizihum (Tkomiiiclh idah) A rf ih f 1IX jii n j a ii ■ ensis^ M cx i co Cliirorhyiiehnbia iiiatsoni Ylinker (Chiroriivnc [iobudaE) A/i(}uiu izeoffroyi, Venezuela CJiirnyssoidc-S urodennae Fain L/yoili’rtna hildlnittufK Panama Demodex carolliae Desch, Lebel, Nutting, and [.ukoschus (DemcidiC[Dae) Cit> t){lui pcrspicilidta, Surinam Denuidex loiiuissimiis De.sch, Nutting, and Lukoschus Ctii'oUia persph illitki, Surinam Ueincidex phyllo^ifcimalis Leydig Fhyllosiomux hoxtafiiSt Surinam bldunnia breviceps Curran (Streui inAE) Li>m hop/iyIki rdhitstiu Panama Eudusbabekia arganoi (Vomero) (M vorinDAH) Desmoiiiix roiu/uius, Mexico Eudusbabekia ceniyi (Dusbabek) linn hyphyUii na/up Cuba EudiiKbubckia danidi (Dusbabek) Pftyifonyi tvris pot’yh Cuba Eudiiisbabekia lepidoxtta J ameson Suifnira litiiitiK Nicaragua Eiidiishubekhi phyllosluTiii Jameson Phyiloxfoniiix discolor, Nicaragua Eitdusbabckia roskkyi (Dusbabek) Mtfnophyllus ciihuniis, Cuba Eitdusbabckia saiminaki (Dusbabek) Macrofiix wafcrfioiisii, Cuba (Isla de Pinos) Eudu-sbabekia urodfrmae Fain Urodenna muf’niro.strfiftp Brazil Eudu-sbabekia viguerasi (Dusbabek) Artiheuxjdfmtk eftsi.s, Cuba (isla de Pinos) and Mexico*’*^ Eutrumbieula alfreddiigesi (Oudemans) (Tromrk ui ume) Artiheus azteciis, Mexico* Ariibeus ja/nuicensis, Cosla Rica* Eiitrombicula batatas (Linnaeus) Microitycierix fnejiulotis, Venezuela Urodenna hlUfhation, Trinidad Eutrumbieula gueldii (Oudemans) Arfiheiix cinerens, frinidad Carol(kt hrcvicanda, Venezuela Glosxtjpkitga lotifiiroxtrix, Venezuela Phyl/osrofnnx dixcolor, Venezuela Phylloxroftiiis hasiattix, Venezuela Siiiraini liHum, Venezuela Eutrumbieula uadchatrami Brennan and Reed Va/npyropx hcllcri, Venezuela Eiitrombieuta pacae{Floch and Fauran) Ctiroliia hrevicaiida, Venezuela SPEClAl- PUBLICAl lONS MUSEUM TEXAS TECH UNIVERS[TY K2 Eutrtnubicul:! vuriiibilis Bren nun und Reed M(u nIPhyUfitii inm rophyllNtu, Venezuel u Eiitrombieuli! wehhi Brcnnun and Reed Arfihi'ns jiifmncen.sis. Venezuela ExastinUiii dnvisi (Pessda and Criiiniariies) (Si rehlidae) A notira sp„ Venezuehi Afuxtra i nltnirn, Panama Anonm gcoJfrt)yi\ Brazil, Colombia, ]^anama, Irinidad. and Venezuela Snirtlira h'fiiifn Hooperdla siietopteryit Brennan and Jones (Tromhk ulidae) Artihi'iisjiifHaicvn.si.lutstiu\. Trinidad (rh>s.st>pluiy(i MffH'iiiu, Costa Rica* (i!<}s.soplui! 4 fiia pi'i-spk-iihutu Nicaragua*, Surinam, and Trinidad CitroilUi Mihrufti, Mexico* and Nicaragua* ChroUfpti'i it'i Nicaragua* yi>iftndu\, Nicaragua* and Trinidad G(phiiyu itlliiithK Mexico* and Nicaragua* Gltissopfuiya ci}»iniis.uu-i\i, N icaragua* Ghn.sttphtiya sitru'itui, Mexico*, Nicaragua*. Panama, and Surinam :\fk t(ff}yi n‘fi\ hirsiiui. I rinidad Mirndn i leris mc^diiotis^ Trinidad FltyfloMtunns iii\ci>h>r, Nicaragua* Stuniira lilimii, Nicaragua* Ikiiupyrn/}! \pevlrHnt, Panama and Trinidad luaitnela marine Dusbabek and laikoschus (MvohuijaE) Mi/Ut}n cn'iuiliifutn, V'enezuela Ixodes sp. (Ixodidae) ArrHyi'tisjiinHiici'n.sI.s, Venezuela Srnr/tira IHiu/ii, Venezuela Stdtttira liuhn ici, Venezuela [xotles dovviLsi Kohls Afiouf d pi’ojfxtyi, Trinidad Labidocarpus lukoschi Fain (LAHiDot Aki’inxE) ,V/KT(oiy(7er/.v ftiepakni.'i, Surinam luiwreiieeoearpiis lobiis McDaniel (Laiudoc arimdae) CiiroUiu pi’f.spk illdta, Nicaragua Lawreiieeoearpiis micropilus Dusbabek and de la Cruz lii(H hyphylin i tivi'rfhu nfn, Puerto Rico [aiwrencettcurpiis phylIcKStomu.s McDaniel Mfcndiyaerix hii Mita, Venezuela f‘hvlh}s!oinits clo/igitnfs. Venezuela Lawrenceoearpus puertorieensis de la Cruz, Tamsitt, and Valdivieso Hriic/iyphyllti vavcrnani/tK Puerto Rico l,eptolrinnbiid(um haniiixiaium (Brennan and Dalniat) (Tromhiculidae) Aniheifs Co-sla Rica* Arsihciis fi)iu'cu.'>, Panama inOLOGY OF THE PHYLLOSTOMATIDaE 83 Tooiiiisioi akitlume Brennan and Reed (Tkomhiculidae) CaroUia Venezuela Loomisia desniodus (Brennan and Dalniat) Amfura fico/fnityL Venezuela Ariihi'ns lofteens, Costa Rica* and Nicaragua* CtimdUi %p., Venezuela CitmUia castanea, Costa Rica* CtiroliM perspk Ufatts, Colombia, Nicaragua*, Surinarn. Trinidad and Venezuela CtirifUia snhruftt, Nicaragua*, Panama, and V'enezuela Desnu/iiiis rotifudfis, Guatemala, Nicaragua*, and Venezuela trophyIhi sezekorni, Bahamas Glos.ufplui^'u lon^hdstris, Venezuela Glossopluipa soficimt, Mexico, Nicaragua*, Panama, Surinam, and Venezuela Liffii hophyllti rtthnsKt, Costa Rica* and Venezuela Micnffiycteris /neiialotis, Panama and Trinidad Mi/itoti vozioneliw, Mexico Sitd Hini iifiii/n, Venezuela Trtahops eirrhosns, Mexico* Vd/upyrops vit/uttts, Costa Rica* I.oomisia stmressi (Brennan) Oirofiut rasta/tca, Nicaragua* Gfossoplhi)*ii sork ifia. Mexico Lom hophylla etnwava, Costa Rica* Mticronts cudfoi'/ticKS, California Toomisia univuri (Brennan) 07 osstfphiiftu sork’itui, Mex\co* LoomLsia yunkeri Brennan and Reed Cun>///f/sp., Venezuela Macronyssoides sp, (Macronvssidae) Enchis/heiies fuiriik Panama UrotSermo hiiohotfon, Panama Vittnpyresstt ptisidu, Panama Macroiiyssoides coitciliatus Radovsky Viimpyrops vifkitifs, Panama Mliertiiiyssoides kochi (Fonseca) Ariiheas nzteens, Panama Artibettxjiiiitciicettsis, Cuba, i^anama, and Trinidad Ariihens iiturafus, Brazil, Colombia, and Trinidad Artihens toftecus, Panama Urochyphylla mimk Cuba Desinoiifts rotnneins, Brazil and Trinidad Giitssophiipu sork inu, Trinidad Phylloitycteris poeyi, Cuba Macroiiyssus iinidens Radovsky (Macronyssidae) Leptoftyclcris Minhor/Ut Arizona Mastoptera guimaraesi Wenzel (Strehlidae) Ctirollia pcrspicilhita, Panama Phyllosrotnus sp., Panama Phyllostofnus hast at ns, Colombia and Panama Mastoptera iiiiiiuta (Costa Lima) Phyflosttt/nits hastatns, Colombia Tifnatia sp., Bolivia. Colombia, Ecuador, Peru, and Surinam Tiffuitia tticarat^iftii', Panama Toiiiitiii sylvicoUh Brazil and Panama 84 SPECIAL, i’UHJ.lCA'l IONS MUSEUM TEXAS TECH UNlVEKSi rV Mcnistopodn sp (SiRi,»i iiXAt) Anihcus vitH'ri'us, I rinitkid Mc^isiopodii ariiiK'it {Coquilleitl Ar!ihi‘u.\ jtiniuicensi.'!, Brazil, Costa Rica, Colombia, Cuba, El Salvador, Guaicmala, Mexico. Panama, Pueiio Rico. Surinam. Trinidad, and Venezuela Arfihai.s fimruft/x Colombia. Pananra, and I rinidad CtirifUiu pcrspk'ilhtiu. Panama Di'Miiiuliis linn/ulus^ Panama PhydoMittnif.^ sp., Brazil and Cuba Phylh>s/t>nm\ t!isfoit>f\ Panama VlcKi''(o|M)d:ii pilakM Macquart I'unipymp^ Brazil. Cuba, Mexico, and U.S,A, Mc^isiopoda provima (Seguy) Stuniint litiiniK Colombia and Panama Sfuntitii huhn’ici, Costa Rica Megistopodii llicodori Wenzel Sfurtiirii luiioviri, Panama MeteLasiiHis pseitdopItTiix Coquillctt (S i Rtim lUxti Anihi'ii^ j(iniaic(’n.si\, Panama ArtilH'us lifiicttiiis, Panama and f^araguay CdfoUiit pcfspicilliitiu Panama Viiftipyre.\sa nytnphunu Panama MkndnHiibiciilii boiieti (HofTmann) (I'kOMHk ui tDACl Ariih(‘u.y fohcca.y, Mexico* Di'.stiuniifs Mexico* Ei ophyila st'Zt'^^orni, Bahamas G!os.u}pliftf>fi soriviiiti. Mexico* Micrutn’i it‘yis C uragao PhyUiksltnims husta/u.y, Panama Microtronibieulii carrncmic (Brennan and Jones) Artihi'ii.y jaiitaic Panama ^’hylii).yrt)flI|t^ tii.st aUn-. Costa Rica. Nicaragua, and Trinidad PhylfoMotiiiis lui.ytiUitx, Panama Smitiini huhn'ici. Cost a Rica and Panama S/tfi fiim tfnuiht.w Costa Rica Mierotrombicula sturnirae Webb and Loomis Sna nins lifinni, Mexico and Nicaragua Sturniro Iwfovici, Costa Rica and Panama StKi’iiini moniiix, Costa Rica Nask'ola uiiiKTcauxi Brennan and Ylinker ('[ romhic ut idae) iuislittfix Veneztiela [Neolridiobius delicakis (Machado-Alliwn) (.SiftEai ioae) Ariihcifs i ineiX'u\, Panama and l i inidad Ai'dht'us jiumiirvnsiy. Panama Phyllti.sitnnity fuixuim.y, Surinam Unnicrma hihfhtintiiK Panama Viiinpyrexw pu.yUht, Panama and Venezuela NydiTiglyphtiS slurnirae Fain (Rost nstfiniiuae) Smr/iitii liHiinK Brazil Nycreriiiiisfes primus Brennan and Reed (Trox)HK ulidam) Afioura sp., Veneziiela Atuxi/'d szeoj'frttyi, V^'enezuela Carol!ill perxpicilltna, Venezuela Dcstiioilitx rofu/nliis, Venezuela Kini.OGY OF THE PHYLLOSTOMATIDAE Ci!<}ss()phii),^ii .u)riciiHi, Venezuelii Lionyc/i ii.s spurn'!!i, Venezuela I^ttfu lhtrltiua aurittu Venezuela Nyeterinastcs scturuliis Brennan and Reed Afjount pi'affniyi, Costa Riea* and V^enezucla Li)nc/it>pliyH(i lohifMa, Costa Rica* NyeteroJiyssus desinudus Herrin and Radovsky ( Mac konyssioae} Diiteniiis yoitnpii, Venezuela N y ft e r( tp li il i a sp. (S t r e r i i d a e ) Mavrorii'i Mexico* Nycleropliilia eoxahi Ferris A ri ih i'us jauitiictuisis Hnuhyphydti cavi'rmirurfu British West Indies \hu rouis ciilifonticu.s. Arizona and California Mat rot us iuiterltimsH, Mexico Nycterophilia purnelli Wenzel CuntlHa pt rspk Ulciia, Panama Marroiux waterhiutsii, Cuba Or n it!iodirros sp. (ARt, as i da e} Lt'puuiycuri.s cuniMHW, Venezuela Lottciiorhitui oriiufce/ixis^ Venezuela tlfm ruuis aififoniicits, Arizona i rt'fiNfafuui, Venezuela Stuf/um at in UK Venezuela Tmc hop.s cin lu>sus, Venezuela Ornilliodciros azteci Matheson ArifheiisJtifutiiceftsis, Cub'A, Mexico, and Venezuela /hf;( /i.vp/n //(7 Cuba Carol fin sp.. Venezuela Lh'smotiiis rotiiuiius, Mexico, Panama, Trinidad (jfosstipiuipa !onpirt}stris, Venezuela Ciiosxopitiipa sorh'iua, V'enezuela Lt)nchoriiithi aariun Cuba. Jamaica. Trinidad, and Venezuela A /utv o/* /;y //;/ /u / cri y / n /, V e n e z u e I a Macro/us watcrhotixii, Cuba PhyUostfOiuis htisusms, Venezuela Trarhops cirrhosiuK Venezuela Ornitliodoros brodyi Matheson A rf!h c us jan i ai i I'/i sis . M e x ic o Cart)!fHi sp,, Venezuela Carolfia piTsph illaia, Panama and Venezuela Clo'ohfpU'ras aurifus, Mexico Lonclarrhina a aril a. Venezuela Travhops cirrhosus, Panama Onilthodoros dusbabeki Cerny Afiihras januilceaxis, Cuba (Isla de Plnos^ ()riiith(»dor(»s haset {.Schulze) Ariihcas jamaicensis, Venezuela Artiheas liiaraias. Costa Rica Untc/iyphylla cai e/tafrt/wn Guadeloupe* and Martinique Caroflia sp., Venezuela Ca roil hi pL'ispirifiata, Venezuela Cliiroik’nna salvini, Venezuela Gloxxophuj^a iongiroxiris, Venezuela Sj^EClAt. PUHIJCA TJONS MUSEUM '[ EXAS TECH UNiVERS[TY L(>n( hoi hifia iitirftii, Venezuela L(>/tcht))-ltiiui ori/ttn c‘nsi.\, Venezuela Miu rotitx H'iUt'rhiitf.'iii, Jamaica'*' Minitfft cn'mihitifm, Venezuela PhyUifnyvteris aphyUiK Jamatca'*‘* PftyliitMiniiiis hasitims. V'enezuela Smniiru III in III, Venezuela Siuniiiti luiitivici, Venezuela Tti/uiiid sylvicnla. Panama Unulc riiui hiftihaiifiiu Panama Unnit riiKi nuiy/iinisfi'uiii, Venezuela Vifinpyrofis ht'Ueii, Panama Oniitliodoros minion Kohls, Clifford, and Jones crcnfifatiiiit, Bolivia Onillliiidtiro’ii peruvianus Kohls, Clifford, and Jones Dfsiiiodu\ |■(H^^tuhl^, Peru Giossoplutya sp,. Peru Omirhodoros rossj Kohls, Sonenshine, and Clifford Glnsxophdfftt liiitf’inixfrix, V e n ez u e I a Leptoiivi U'l is nivalis, Arizona /a /i (j; ji f u ( u /Vj oc (^j ,v A, V e n cz u el a Macmfns i'tilifornicits, Mexico Oniilliodnros viyuerasi Cooley and Kohls /JjfK/?y/j/n7/u /itiiu!, Cuba Eniphylla luwihi/mns, Puerto Rico P/jy/yfinty/em/jfoeyi, Cuba and Haiti** Oniilhodoros yuniiilt'rjsis Cooley and Kohls Ai'iihetfs a-terns, Mexico Aitiheiis litnratns, Mexico Cnioiiici perspicillntn, Venezuela Desinodns rt^miulns, Mexico Paradyschiria pan utuides Wenzel (Si rilhi idae) Antinra j^edJJrnyi GItisstiphaya St irk inn Paraeuctenode-s lon^ipes Pessoa and Cuimaraes (S irehlidae) Ananru eantlifer, Brazil Amnira }>roffi (>yi PliyUdstiniUfs hnyratns, Brazil Parakosa maxima McDaniel (L.AHiDOt arridae} Euc7;i,v//u'/je.v hnriii, Venezuela Ghisstiphnya Inn^irtisfris. Venezuela Parako.sa ladarida McDaniel and Lawrence Cartiilin hrevientain, Venezuela Ghisstiphnya loni>irt}Str!s, V'enezuel a Stnrtiira I Hi inn, Venezuela Paralabidocarpus artihei Pinichpongse (LAiUDOt arfudae) ArfiheiiS iitnrafns, Trinidad StefHitlerifui rafnni, Puerto Rico Sinriiira I if in in, Nicaragua Paralabidocarpus caruMiae pain Cartiilin perspieillnfa. Surinam Paraiabidocarpus de.Miictdus Fain Desiiunliis rtnnndns, Surinam Paralabidciearpuv foxi de ia Cruz. Tamsiti, and Valdivieso Artihens jnmnireiisis, Puerto Rico Sienodertna rnfnrn, Puerto Rico BIOLOGY OF IHH PHYLLOSTOMATIDAE 87 P:ir:iliilM(toc:irpiis miicropLyBiitii Fain Mucrt.>pliyflufn nua ropltyflitf}!, Surinam HariiIal>jdoeurpii.*i .slcn(>d(?rnii de la Cruz, TamsitL and Valdivicso S/enotiernia nifum, Puerto Rico Parahibidocarpus (nnaliaf Fain Tofuitia vc/iczt/i’ltie, Venezuela Piiralabidocarpus Irachops Fain TttH hop.s cirrhosns, Surinam ParaseoschoetiKastia aemiilata (Brennan and Jones) (Tromhk ui lOAt) Atiouiii ciuulifcr, Venezuela Siurnira liltmtu Mexico’*' Parascosehoenii'asthi Jiiejiiasfyrax (Brennan and Jones) Carollia pcrspicHUncu Panama Parasecia longicalcar ( Brennan and Jones) (Thommicui ume) DesiiKuitiS nitaiidii.s, Trinidad Vtimpyruiti xpi'csntni, Panama Paraseeia inanueli (Brennan and Yunker) Uroihrmii hilDhtitnin, Costa Rica* Parasccia souccniyanH (Brennan and Yunker) Sfurflint (udovici, Panama Parastrebla handleyi Wenzel (Strehi idae) Xlicroiiyciei i.s tticcfari, Panama Pa rat rk‘bob I IIS sp. (Strehi.idae) Aniheiix azti'cas, Panama Anihi'itx fitiaattix, Colombia and Panama Aniheus loltectts, Panama Chinnkr/tm villoMtui, Panama Vitmpyrops fiefk'ri, Panama Vitfitpyropx vittaftix, Panama Paralrichobhis ainericaiuis Peterson and Ross Cinfcmtiyi feris mc.xUntui, Arizona Paratriehobius dimni (Curran) Arliht'ux jiitiiciici'fixix UnKk'rffiii htfohmu/ti, Panama Paratrkbobiiis lonsicrus (M, Ribeiro) Ariihi itxjattuik i'tixis, Brazil, Colombia, and Panama Af/iht‘ii\ iitunitttn El Salvador and Trinidad C(i ro 11 hi p i‘ rsp k it f a f a Urotierma hilohahuft Vit fi ip yropx I ift eti s t ts Pararriehobius lower Wenzel Aitihi-fix watsonh Panama Pa rat ric hob ins sahini Wenzel Chiroderma xak'ifih Panama Paratrichobius sandiezi Wenzel Envhistheties hat tit, Panama and Venezuela Paricharoiiyssiis sp, (Macronvssiijae) Arfihetis aztevtts, Panama Aniheus toiteens, Panama Vitiftpyrodes caraet iuioi, Panama Parklioronyssijs erassipes Radovsky Carol (la perspiciUaia, Panama Paritboronyssus eiithysterriuiii Radovsky Sturnira ludovtci, Panama ■SPECIAl. PUIHJCATKJNS NtU.SEU\f [EXAS lECH UNiVERSETY Paridioronyssus sdcriis Kiulovsky CJlo.wttpiuifio ^oricithi, P^aniinii’i Phyilt>.%iit/nuM ^p.. Costa Rka Peralfs uiioplitluiliua (Hoffmann) ( rKOMHii Li lo.vt) Arnln ti.^ tizfemx, Mexico* CttroUki pcr.'.pk lUiitti, Trinidat-I Dcsnuulii.'^ ri)iit/}tlu\, Mexico and Punuma Erophyihi u’zckoi Hi, Bahamas Micntnycieri.'i iru-fzaiotis, Colombia and Pcrii Periylisdirus sp. {Si’in iurnk iDAid Ctiroiiia ffei‘\pu atiitu, Panama Ltftuhophyilu nthtiMa, Panama Xltirmphylfiiin mttinfphyiluffK Panama PeriKlischrus acutistermis Mach ado-Allison Ariiben.\ Venezuela PhyUitsiottui.’i lUscitliff, Trinidad and V'enezuela Pliyl!i)stonii(~s i'ltfftptifi/s, Trinidad and Venezuela Phyllo\tonu('i htisiam.s, Colombia. Panama. Ti injdad. and Venezuela PericUseliriJS ealkiis Kolenati Atufio’d ccuuSifer, Venezuela AftiHfid duitruSa, V'enezuela Ghf.i\(iphiii>u sp,. Mexico and Panama tonpiriJi\fri\, Venezuela (/htswphapii 'iork'i/uK Brazil, Panama, Surinam, 'Prinidad, and Venezuela PeriKlisdirus cubanus Dusbabek Bi’tH hyphyffti naruu Cuba Erophyiiu sezekorni. Cuba PhyllofiycicrL’i poeyi. Cuba Perijjtisdirus ddfiiiadoue Dusbabek Men io(n\ n tiicrlitfiisfi, Cuba Perijulisdirus diisbabeki Mach ado-Allison and Aniequcra creniihitiifii, Venezuela PtTiffliselirus yiniieroi Niachado-Allison and Anlequera l,of}i hin hifui iinritu, V'enezuela P e r IK I ise h r us h erre ra i M ac h ad o - A 11 i so n roiufidiix Panama, Trinidad, and Venezuela PeriKliscliriis hopkhisi Mach ado-Allison LiddyUfris \purrcili, Venezuela Khitiophylhi psffuiluh Brazil and Venezuela PeriKlischriis ihennyi Oudemans Aniheu.s ^p., Panama Ai{ihcusitz(evti\. Mexico. Panama, and Venezuela Ai ttheifs t imreiis, Panama. Paraguay, and Venezuela Arrihi'tfs loltci tis. Panama and Venezuela Ai'!ihi'u\ conrolin'. Venezuela Artihetisjtitudiccnsis, Cuba, Mexico, Panama. Puerto Rico. Venezuela, and Virgin Islands Arriheifs lilurafus, Brazil, Colombia, Guatemala, Honduras. Panama, Paraguay, Surinam. Trinidad, and Venezuela Chiroth'nna sp., V'enezuela Chiroth’tHui stifviiii, Panama and Venezuela DestimJii^ rt}[u/!dtaloi{s, Peru Perisxopalla exhuniatus (Brennan) Ctindlia perspkilitita, Peru and Trinidad SPECIAL PUBLICATIONS MUSEUM lEXAS TECH UNrvERSlTV y() Di'.sfiiifihis rift Hint Its, Trinklad D lit ( '/! I as >' f) utit^ i /. T r i n i lI ad GhnM>pltiii,’it styficinti, Trinidad MicntnycU'i h Peru PL^rissopuUu ipeani Brennan Citroltia pcfspicHiuui, Brazil and Surinam Perissupalla precaria (Brennan and Dalmat) Dt xn!<}iiif\ ri>tu/nln.\, Trinidad Gli>ss(fphu^u Mtririnti, Mexico Miaonyiiei is mcf>{t!o{i\, Panama Plijllcistoinonyssiis canradymikcri Fain (GaSI ronvssidae) AiiihiULsJauuiicvH^i.s. Venezuela Arfihi'us Surinam and Venezuela UrtHh’niia Surinam Vtinipyrops helivri, Surinam Pseiidoahibiducarpiis secus McDaniel (LAainH:>c \RPin,\r 1 FhyUttMttnuf.'y d/u o/or. V'enezuela P/iyllt>\it)intiM V''enezuela Pscudoschoenyastia hulhifcra Brennan (TkoMiitc ui.idae) Slut nhit tt(dtn>ict\ Panama Pseudovtreblu ('reenweUi Wenzel (Stkehi idaeJ Toitiiiiit Panama Pscudoxtreblu ribeiroi Cosla Lima lotuina sylrirolit, Brazil and Panama PvorerKaloidcs artibei Lukoschus, Rosmalen. and Fain (Psorercmtidae) Artihenx iituntntx, Surinam Psfirerji;atoides j»lossopliagac Lukoschus, Rosmalen, and Fain Glitwtfplutpti soficimi. Surinam pNurcrKatoide.v londiorbinac Fain Lonvhorfiinu auriitK Venezuela Radfordielta atiourae Radovsky, Jones, and Phillips (Macronyssidae) Antuint f’eojfroyi, Mexico Radford id b carol Mae Radovsky CtintHiii caxiiineu, Panama CiiniUht pi’fxpicitifiup Panama (Canal Zone) Radfordidb desniodi Radovsky CiiroUia pi'rspicttfiiui, Trinidad Dextiioilns ri}iuttJifx, Panama and Trinidad Radford id la iitaiiopbytli Radovsky. Jones, and Phillips Monophyllux n'llmanp Cuba Radfordidla oricola Radovsky. Jones, and Phillips Aiuuit’u yaijfn j_v /, Mexico Li'p/o/tycferh nivulix, Mexico Radford idla oudeiuansi Fonseca BrttchyphyUa cavvrfwrum, Puerto Rico Dextimtius rtfitifuii/x, Brazil Duii fitiix y(tnftgii, Trinidad Spelseria ambi^ua Kessd (Streiii (Dae) Arumfo I’coffroyp Trinidad Cariflliii caxuineth Panama Carol!ill perspk iUtiia, Colombia, Panama, and Trinidad Caro! I id xuhrufa, Panama Desmadux mtutidiix, Panama Gft>xsopha^d x<>rk-iiiu, Trinidad Li/ruhophylla rohuxiid Panama BIOLOGY OF THE PHYLLOSTOMATIDAE 91 Loiii horhitui tun tin, Panama Mkronyt ieris hrachyotk^ Trinidad PkyUosjomKx husmtits, Panama TifniHia hUivns Tnu hops ciryhosHs, Panama Vaiiipyrops viltatHS, Panama Spelaeorhynchiis sp. (Spelaeorhync hidae) CitrofUa perxpk UksiiL Brazil Spetaeorh.viK'hus inoitoplivlli Fain, Anasins, Camin, and Johnston MtmophyllHs ndniuui, Pnerio Rico SpeJaeorhynchuK pniecursor Neumann Anihvux sp., Mexico AfilbeHs jamuicensis, Cuba. Dominican Republic, Mexico**, and Puerto Rico Carokki vasiunetu Mexico CttroKki perspk'iUaus, Brazil, Colombia, Mexico, and Venezuela Glossophitpa sork imt, Amazon(?) Spidoocliir aitkeiii Fain (Erevnetidae) Anoum pvoffroyi, Trinidad Spcieochif barliuhita Fain and A it ken Minuiti Brazil Speleochir brasiliensis Fain and Aitken A nihvHs jamuiven^is, Braz i I Va})tpyriHii‘s curuvcioiok Brazil Speleocliir curotliae Fain and Lukoschus Citrotiia pt’yjipk'iUaUu Surinam Spcieochir phyllostomi (Clark) PhyUostomu'i hastuttis^ Colombia Speleocola davisi Webb and Loomis (TrombiculiDaE) Desmodtia rofundns, Mexico CkossophapLi sork imt, Mexico Lt pionyaeris sanhornk Mexico Speleoeola seeunda Brennan and Jones CuroUkt castauvuy Nicaragua* CuroUkt perspk'ilkiSiu Surinam CttroUkt suhiiifa, Nicaragua* Dcsmoditfi roumdits^ Trinidad Gio.sMfphapd viHUrnkstifisk Nicaragua* Giossophiiga Sifrk inu, N icaragua Minanyvteri.'i hirsuia., Trinidad Mkronycteris nreptdoiis, Trinidad Sleatonyssii-sjoaquimi(Fonseca) (MACRONYSStDAE) Giossopha^it sork'inti^ Brazil Stizoslrebla longirosiris Jobling (Streblidae) TiffukUi sp„ Brazil and Colombia Strebla sp. (Sirebiidae) Dku'fuusyoungiu Trinidad Strebla aJlniani Wenzel Cundikt perspk'iUatiU Panama Liinvhorhinu uurku^ Panama and Venezuela Mitcrophyllum mttcrophyllnm^ Panama Tyiu'iuyps viryhosHS, Panama Sirebta alvare/.i Wenzel Micyottyvfcy'ts megtdosis^ Panama M /l i t >/ly (7 er is nit efoyiy Panama Micyo/ivi teris xv Ivemris, Panama SPEC’lAl- PUBLlCA riON-S MUSEUM TEXAS IECH UNJVERSITY Slrt:h[ii carol line Wenzel A riiheit.sjatfiaiccnsis, Panama Caioilut sp„ .Surinam Ctiroflui aiMafica, Panama Oirtilliii pi‘rspici!tittu, Brazil, Colomhia. Panama, Trinidad, and Venezuela Camllia siihrsf/tu Panama Di’s/hihSils ro!tfnthi\. Panama CilossDpliiipit st)rU i/!ii, El Salvador, Panama, and Venezuela L<>ui'lH>phytfa nthtiMu, Panama Lonchtfi hinti aurita, Panama Mtu rophylluiii imicri>phyilntn., Panama I’hythfMoiiius hciMafus, Panama rrai hops cinhosn.'i, Panama Strebla Christ inne Wenzel Pliylitnk rfnu sietutps, Panama Strebla ccjiisoclus Wenzel CtintUui pcr.'ipicikiKih Trinidad Fliyllostointis sp., Peru and Surinam Phylhniti/nns di\rt)f,)r. Colombia PiiyUifMofiUis htistiPus, Surinam, Trinidad, and Venezuela Ti ariiopx sp., Peru StrebJa tJiaemi Wenzel Dine in US ytnntjiii, Colomhia and Panama Strebla diphyllae Wenzel Dt'snunhis nituftilns, Guatemala Diphylln ei duihiia, Guatemala and Mexico Tnit’httpx i'inhtf.sn.s, Guatemala StrobJe suli'^doi Wenzel Tonatitt sp., Trinidad Ttnuasu hUli’n.s^ Panama Strebla liertliti Wenzel Artiheux jaiiiaii'eitxi.s, Panama Dt’xinoilus rtffuiuhix. El Salvador and Panama Phyllostnmiix iIixrott»\ Colombia, Costa Rica. El Salvador, Mexico Nicaragua, Panama, Surinam, Trinidad, and Venezuela Pfiyliosfomux Ita.sidHix, Costa Rica, Nicaragua, and Panama Slrehla boogstraati Wenzel Tonal in nicaragnae, Panama Sirehta kotilsi Wenzel Tonal hi sylvkohn Colombia and Panama Strehlu inaehadoi Wenzel MitronycU’fis niitinkt, V^enezuela Strebla mirabilis (Waterhouse) Carol}ui perxpicilktta, Brazil. Panama, and I rinidad DextntHlus rot ami as, Peru and Trinidad Dipiiylla erauilaia Glosstiphaga sin k inn, Trinidad Phyilosioinns np., Brazil, Panama, and Peru Phyliostotntis disaolor, Trinidad Phyiloxiotniis i'knigatiis Phylloxtofnax hnsinius, Colombia, Panama, Peru, and Trinidad Tonaiin sp., Colombia Tonal in hiden.s Strebla lonatiae (Kessel) Tonalia hkh-nx Tonal in hmsHiense, Ecuador and Panama [BIOLOGY OF THE F^HYLLOSTOMATIDAE 93 SIrcblii uiedeituiiini Kolenaii Afioiira aiUiiif'er, Brazil Anauni ^‘coffroyi A riiheux jamtiicensis, Panama Chiotopferus auriius, Brazil fyiwmoilu.s ftffuntins, Colombia^ Ecuador, El Salvador. Guaiemala, Dcs/iuhIhs nimtiduA, Colombia. Ecuador, El Salvador, Guatemala, Honduras, Mexico, Panama. Peru, Surinam, 7 rinidad, and Venezuela Vatiipyfopx li/u'iitus., Brazil Tecoiinitluna sandovali Hoffmann ITrombk ulioae) A rs ihcM pluu otLs, M ex ico* Desftitnfti.'i rofnf!eIi(\ Mexico"* Miicrofuji califoniicus^ Arizona Tecoiiiadana wiilkinxi Vercammen-Grandjean Mavrotii'i fadfornkus, Arizona*, California, and Mexico* Trkhobioides perspicilhitiis ( Pessoa and Galvao) (Strfhi idafJ CoroKhi pvrxpk ilhftii, Brazil, Mexico, and Peril DiXfniHlux nxiouiti.s, Panama and Trinidad P/iyllt}xttff}ii{.'< tlixcolor, Colombia, Panama, and Trinidad PhyUDXiiXXiis elongatttx, Colombia PitylUfMoniiis luixtiUifs, Surinam Sttii nint lifiifftu Panama Trichobius adatnsi Augustson (Streiii.idae) Mint (XUS cal {torn fcii.w Arizona, California, and Mexico Tricliobiu-S bequaerti Wenzel Tomukt hidexs, Panama Trtchobiux brentiiini Wenzel Siurnxd (sidovici, Panama Trichobius cernyi Peterson and Hurka At !iheus jxxuticexsis, Cuba jV/(jnojt)/;y//n,s' redxiaxi. Cuba F/iyUo/jycU'ris povyi, Cuba Trichiihius costal iniai Guimaraes Artiheus }ifuraSi{S, Panama Candiki perspk iUuUi, Panama Desxiodiis roimulus, Panama Pityllostomus discolor, Colombia, El Salvador. Guatemala, Panama, Peru, Puerto Rico. Trinidad, and Venezuela Urodertxa hilohdixnh Panama Trichobius diphyllae Wenzel Diphylhi I'caudaiu, Guatemala, Mexico, and Venezuela Trichobius doiiiinicanus Peterson and Hurka Miuxrphylhis sp., Dominican Republic Trichobius dugcsii Townsend Auourts ijeojfroyi, Trinidad Anibeus jiixuiicexsis, Cuba and Panama Curollki pcrspk ifhxx, Costa Rica, Nicaragua, Panama, and Trinidad Dcs/nodus rorundiis, Trinidad Diaexius yoxtigii, Trinidad Enchisihcxcs hurl Ik Trinidad OItJssophiiga soriciftit, Colombia, EJ Salvador, Guatemala, Mexico, Panama, Peru, and Trinidad M i crony Cl er is b rut / ly o (is, Tr i n i d ad Fiiyllostonuis hastcxus, Trinidad Tiiichops cirrhosus, Panama 94 SPECIAL PUBL]CAT[ONS MUSEUM TEXAS TECH UNIVERSETY Trk'hcjhiu.'i dii^esiidide.s Wenzel OtroUsa pcr.vpk illain, Panama Chroiopii'i iis tiKridi.s, i’anama Litnchofhina nuritu. Panama Tiachtips ciirhtKsns, Panama Trkholiiu-s tijbasi Wenzel Ttitutiia sylvfcohh Panama Trichobius frcqiiens Peierson and Hurka Arfiheus jaiiuticL'nxiA, Cuba and Dominican Republic Brachyphyfid tutiiu, Cuba Hravhyphyihi pu/iiihi, Dominican Republic Entphyllii Cuba Mi>iu)phyt(u\ reihminiy Cuba PhyUimycterL'^ poeyi, Cuba and Dominican Republic Trk'hobius ftirniani Wenzel DcmhoiIux rota/aiiis, Peru Dipluylht I'caiHiattu Colombia Gto,sM>phu,i>u ,'itn'ici/uu Paraguay Trictiobiu.s iiiterinedtus PeieritOn and Htirka Artiheus sp.. Guaiemala and El Salvador Arrihen\ hirsiitns, Mexico Aitifn’usJatnaici’iixi.s, Bahamas, Cuba, Dominican Republic, Jamaica, Mexico, Puerio Rico, and Virgin Islands Artihfiis!ittinKus, Mexico ErtiphyUii Cuba .Vfiicnflit.\ waterhousii. Jamaica XftWiiphyllHs rethmtni, Dominican Republic I’hylionyi tcris porvi, Cuba and Dominican Republic Trichobius Jobiiii^t Wenzel ArriheiiSju/iiaici‘/t.\is, Panama A rt ih eus lift! ro! n s. Pan a m a Corttifiu t tiMla/tcu. Panama Ciiroffia pcr.\pk idafu. Brazil, Belize, Colombia. Cosia Rica. El Salvador. Ciuatemala, Panama, Peru, Surinam. Tobago, Trinidad, and Venezuela Carol fid Affbr/(/lY. Panama Chiroder/na villoMi/a^ Panama Dc^tfUHiKx rofii/nius. Panama and Trinidad Gfo.\Mtpfuiga soriciim, Panama and Trinidad /.(>jik‘/iop/j\7/(7 rahnslti, Panama I.ofti horhina (luritid Panama MdcrophyUiifti /naiTophyihim, Panama M itrns liiurtiitix, Panama IjfnihophyUti rohuMa, Panama I'riclinbrns Jonf»ipe?» (Rudow) Atio imi p coffroy i. T r i n id ad Ardhcuy jamalcensbi Cuba and Panama Carol!u! pcrspk iffaiii. Panama ClioeroiiyctiTix ow.xicanth Arizona PhylfoMiiomix sp„ Panama Phyllojiioffius diwolor, Trinidad PhylhfMontiis iuixfatus, Bolivia, Costa Rica, Colombia, Guatemala, Panama, Peru. Surinam, Trinidad, and Venezuela Irichobius inacropliylli Wenzel Cdfollki perspicilhiftu Panama Lonchorhhid imriia. Panama Mai rophylhon inatrophyllum, Panama Trkhobius macroti Peterson and Hurka Mai I c/iiis M'uU'rhouxii, Bahamas and Cuba Tridiobius niendezi Wenzel Toaatia nkanipuai\ Panama Trichobius iicotropieiis Peterson and Hurka Mittroiux ivaieriiousil, Dominican Republic Tridiobius parasitkiis Gervais Caroilki perspk'illuta, Trinidad Desfnoiias runtfHlu.y Brazil, Colombia, Costa Rica, El Salvador, Guatemala, Mexico, Panama, Peru, Surinam. Trinidad, and Venezuela Dkietitax youat^ik Panama Diphylla ci aaiiafa, Mexico G1 1 M'xopl lapa xoricf/ia MoMop/ty/las ri'i/ataak Jamaica P/i yl/o /lycU'ris poryt P/iyllosio/nux haxtafus, Panama lomitia sylvicofa, Brazil Vaaipyrita! spt'dnau Triehobiuji pbyllostomae Kessel Phyilastotuux sp., Brazil PhylloMo/tiNx hnxmiux Trichobiiis pveiidotriincatus Jobling Arliheiixjitatai ix nxix Tridiobius robynae Peterson and Hurka Artihi‘ux jamaicenxix, Puerto Rico Eropfiylht sezi'korni, Puerto Rico Monophylfm reihuunk Puerto Rico Tridiobius sparsus Kessel Carollia pcrspicilhtiih Panama Tridiob ins sphaeronot us J obi ing Li ptonycieris Texas Li'pto/iycteris sarihorni, Arizona, Mexico, and New Mex ico Triehobius truneatus Kessel AftihI'lix ju/fUik eaxix Bracbyphylki cinxtuarntti, Puerto Rico Erophylht homhifrons, Puerto Rico 96 SPECIAl- PU BMC'AT IONS MUSEUM TEXAS TECH UNIVERSIIY Miu ratus \viiterhonsH, Cuba MouophylUis rednutni, Puerio Rico FhyUonyt leris poeyi Tricliobiiis iitiiformis Curran A rt the tts ja i! ui it v nx is. 1^ a n a ni a Dcsfiioiltfs rtuundits. Panama C!osst>phapti sof iciiui. Costa Rica. Gualcmala, Guyana, Mexico, Panama, IX^ru, and Venc/aicla Ltinviiophylia n>husia. Panama Tridiobiiis iirodcrmne Wenzel Uft.)dt‘riiui hiiohtUifrti. Panama and Venezuela Trifhobiii.s vainp,vr(i[iis W^enzel Arlihi’us litarunis. Panama {‘'(unpyyops viifutuis, Panama and Venezuela Trtcliobiiis juiikeri Wenzel Anihnts UtunPus. Panama Ctirttllia perspiciilafa, Panama I.tnicfunhhiti Panama Still t}ini imfovici. Panama Troinbicula diimii Ewing (TartMint Ui id vil) Ti^/n/jyrt pusiilti. Panama W'agennaria siintlis Brennan (Trombiculidae) Gltisstipfuif^u si>rici/ia, Mexico* W Jiarlonia glenivi cnlifoniicii Vercammen-Grandjean, Watkins, and Beck (TftoxtBirL'i idae) ChtH'nntytfi'ii'i im'xifiitia. Mexico* Muc nitiis citlifoniiciis, Arizona*. California, and Mexico* Whartonia gutTrerensis Hoffmann Eynphyffti screktirni, Bahamas W'liartonia nudosetosa (W'arton) Anihi’iis jufiuiict'iisis, \tcxico Ctirollia sp.. Costa Rica* Ciiroliia perspicilliitii. Costa Rica*, Guatemala, Mexico, Nicaragua*, and 'I'rinidad CiirnUia siihnifit. Mexico* Dfstittiilifs rot (Indus. Mexico and Trinidad G!os.\opiiti^>u soririiui, Mexico and Nicaragua* Mw rot ns sp.. Jamaica Miinon ctKitnu'kie, Mexico Wbartonia paebywhsirtoiiii Vercammen-Grandjean Xliironyt teris oifptilons, Brazil Xeiiodontacaras serrafvis Loomis and GofT (TrombicuiHue} Ar/iheiis litnrutns. Mexico Host-Parasite List Ectoparasites known from each host species are listed alphabetically. A single a.sterisk indicates an unpublished record from the Chigger Laboratory, California State University, Long Beach; two asterisks indicate an unpublished record from The Museum, Texas Tech University, Lubbock, Geographic origin of records is given if knowai. Since this list was prepared, four publications (Brennan and Bronswijk, 1975; Brennan and Reed, 1975; Herrin and Yunker, 1975; Reed and Brennan, 1975) have appeared that should be consulted for additional records. BfOLOGY OF- THE PHYLLOSTOMATIDAE 97 Anoura sp. Aniisirchhi tfUintnh'/ii (Streblidac), Venezuela (reporied by Wenzel (/ nl., 1966, as from A. ucuiedUi, possibly a manuscript name, but in any event unknown to us) Excnf/ffhw r/fO'f.vi (Streblidae), Venezuela (same as above) M'c/ rn/iM'fe.v /) fTromb ic u I idae), Venez uel a PcWA'/fvc/jrn.v vuri'a^i (Spiniurnicidae). Guaiemala Anciura ciiJtrata Handley A/Mi/ri*/Atf (Streblidae), Panama i/orni (Strcblidae). Panama Eii'r/ff//.Kc /rnn n/Z/ji'/n (Strebh'dae), Venezuela (Sireblidae), Panama and Venezuela Anoura caudifer (E. Geoffroy St,-Hilaire) /l/fi/Vr/ounpizi;yhrffia/i/' (Labidocarpidae). Venezuela Am.\f/c'/f/ii (Streblidae), Colombia '^crfe/i'Of/e.v /ouj^/pi'A (Streblidae), Brazil E/tnm'o.u /iiu'fif’iJi’rut urvu/dun/ fTrombieulklae), Venezuela Et'/iA’/Ec //rus ru/krtzs (Spinturnicidae), V'enezuela Err/jf/ivc/fruv (Spiniurnicidae), Venezuela n (Streblidae), Brazil Anoura geoffrojlGray Labidocarpidae), Nicaragua and Trinidad Ayu/xfr( />/i/ ///////iu/r/if (Streblidac), Panama and Venezuela A/u/.Uff/)//j //iiH/tsf/f// (Streblidac), Guaiemala, Mexico, Panama, and Trinidad A yp/iiiffp/i‘f'a pAy//(Jxffjr/tyNt7Ni6/f/ (Chirorhynchobiidac), Venezuela Ixoth's tiou-fi.si (Ixodidae). Trinidad LifOftiisici dcxmoiifi.Y (Trombiculidae), Venezuela Ny (■{ cr imist ci' p/*/n t /c v (Trom b icu I idae), Ven ez ue I a yVvc/r'wuzv/f V (Trombiculidae), Costa Rica* and Venezuela PartiJyscf! irici parv uloidcs (Strebi idae) Pitraciicfemhlex iofipipcs (Strebl idae) Pi’n'ftltschnis pifimiffistcnius (Spinturnicidae), Venezuela /Vng/(ji‘t /jn(A K’di^fasi (Spinturnicidae), Colombia, Guatemala, Mexico, Panama, and Venezuela RdilfonlieKci a/iaumv (Macronyssidae), Mexico Rmljhrdkdhi (Macronyssidae), Mexico Speist’riit iitnhi\dHi (Streblidae), Trinidad SpcktHhir fu/Ae/ff (Erey net idae). Trinidad Sirchlii n'iciictiwnfii (Streblidae) I'rkhohiii.s ihipesii (Streblidae), Trinidad 1 rk hohitis fattpipes (Streblidae), Trinidad Artibeus sp. AspiiiopU’ni hiiscki (Streblidae). Puerto Rico A sp ido pHni piiyUosivnuitis (StrcbViddc) Perififischnis /Viefoipj (Spinturnicidae), Panama Spekii'fn hynchuA prtii'i nrsor (Spelaeorhynchidae), Mexico Trichohius inier/nedius (StTCbiidac), Guatemala and El Salvador Artibeus aziecus Anderson Eninmihh idci tilfrcddtipt^si (Trombiculidae). Mexico* \fiii ro/iyssokh's kiK hi (Macronyssidae), Panama 98 SPECIAL PUBLICATIONS MUSEUM I'EXAS TECH UNIVERSITY Offiifhintoi’o.'i yufitaiensis (Argasidae), Mexico P(inttrkii(thiu-s (SirebJidae). Panama Parichoronyfi.sussp. (Macronyssidae), Panama Pcrau-s (i/niphihtilftui ( ITombicuIidae), Mexico* Ao/e/f\«7j'fj'f.v/Va'rof,!?/(Spinturnicidae), Mexico, Panama, and Venezuela Pc risst }pii I la hcf (ni ; o (Tro m b i c u I i d a e). M e x i co * Arfibeu-s chitTeus (Gervais} Atahkh/airptis f’ltya/icnsix (Labidocarpidae), Surinam Chirnysxakics captii fi (Sarcoptidae). Panama Etiinwihu ala (Trombiculidae). Trinidad McykMoptHlii sp. (Strebiklaeb Trinidad i\'cafrichohiu.s (Slreblidae), Panama and Trinidad Pcfii^ffischrux theii/iiii(Splnturnicidac), Panama, Paraguay, and Venezuela Artibeii*^ toiicolor Peiers Pcfiyliwhnti ciciitisicrnus (Spintumicidae), Venezuela PcfifiHschriix iherin^i (Spiniurnicidae), Venezuela Artihi^us Itir.sutux Andcritcn PcrixMipalla hcftrufii (Trombiculidae), Mexico* Tt k htfhinx iniernialinx (Streblidae), Mexico Artibeus jamaicensis Leach Aftihlyotnma sp. (Ixodidae), Venezuela Axpitiapierti (Streblidae), Colombia, Cuba, Guatemala, Mexico, and Panama Baxifiii tycltaniii (Nycieribiidae), Brazil Htixiiki roiiilanii (Nycieribiidae) iiaxiliu ire/jce//(Nycieribiidae), Panama C/n>nyjf.vo/fi'('.v ru/jf/rj'/(Sarcoptidae), Mexico** and Panama Chirtypicila (Trombiculidae), Mexico Eitiiitxhtihekki Myobiidae), Cubadsla de Pinos) and Mexico** Entroftthk iila u//ref/(//d'^u'.V ( Trombiculidae), Costa Rica* Etitrotiihii'nlo u chhi (Trombiculidae), Venezuela Hoopcrclhi saciopicry.\ (Trombiculidae), Costa Rica* Ifooperclia n.'ipenti.'itth (Trombiculidae), Mexico, Nicaragua*, and Trinidad /Af)i/e,vsp. (Ixodidae), Venezuela lA'ptotronihidiuni htuntixuiiu/n (Trombiculidae), Costa Rica* and Panama XfiicrofiysMikh’x kochi (Macronyssidae), Cuba, Panama, and Trinidad Mc^>ixiopo(la urui-jcu (Streblidae), Brazil, Costa Rica, Colombia, El Salvador, Guatemala, Mexico. Panama, Puerto Rico, Surinam, Trinidad, and Venezuela \ttichixmtLs pseitdop/erux (Streblidae), Panama XfkrotroathicuUi t'urtitcnae (Trombiculidae), Panama .\'coirichyhius del teat nx (Streblidae), Panama j\'ycfcr<}pfii({a coxahi (Streblidae) Oi niihodinox azrofi (Argasidae), Cuba. Mexico, and Venezuela Offuthoiktntx hrodyi (Argasidae), Mexico Or/tillu>iit)rox iluxhabcki (Argasidae), Cuba (Isla de Pinos) Or/iitfuulonix /iuac) (Argasidae), Venezuela Pu™/ubk/oZu'u.v/r( t/Neu5(Streblidae). Cuba and Dominican Republic Trk hobiit'i hitcrmcdiits [SirebWdae)^ Bahamas. Cuba, Dominican Republic. Jamaica, Mexico. Puerto Rico, and Virgin Islands Trk htihinx Joh!i/igi (Streblidae), Panama Ti k ht>hiu.\ (Streblidae), Cuba and Panama Trk hohiux pxeudoiruiiciitiis (Strcbl idae) Trkhohfux roZiVfpi^’(Streblidae). Puerto Rico Trkh(fhiti.\ tfummax (Streblidae) Trkhohiux ;/p'iif/bnii;A (Streblidae). Panama Wh(tr!(}fikt fuuktxeioxa (Trombiculidae). Mexico Artibeus lituratus (Olfers) Anihlyofttnia Rp. (Ixodidae), Venezuela Anihlyaoitiui longiroxirt' {lxod\dac)^ Venezuela A 'ipidtiptera hiisc ki (Streblidae), Panama Aspidoptera phyllosionititix (Streblidae) fkitopereflu vesperuginix (Trombiculidae). Mexico Mtu ronyssokiex kochi {Macronyssidae), Brazil, Colombia, and Trinidad Mi'gixiopoda (imiii'ii (Streblidae), Colombia, Panama, and Trinidad Maelaxfnttx pxeiidopierHs (Streblidac), Panama and Paraguay Or/ikhodtHos huxei (Argasidae), Costa Rica Orniihodorox yuuuiienxix (.Argasidae), Mexico IkirahihidiK-arpiiX arfibci (Labidocarpidae), Trinidad Pantirk hohiits sp. (Streblidae), Colombia and Panama Pi/rufrfWiob/ns(Streblidae), El Salvador and Trinidad Feriglixchrtis (Spinturnicidae), Brazil, Colombia, Guatemala. Honduras, Panama. Paraguay, Surinam, Trinidad, and Venezuela PhyifossofNony.xxiix conradyutikt'rl (Gastronyssidae), Surinam and V'enezuela Pxorergtiioidcs uriihei (Psorergatidae), Surinam Trsc/iohiifx coxtiilimai (Streblidae), Panama Trkhohiux (Streblidae), Mexico TrichohiusJohli/igi (Streblidae), Panama Trkhohius lofniiophylhie (Streblidae), Panama Trkhohiux (Streblidae), Panama r/’(V/iob;V/.v.vfi//A:£'ri (Streblidae). Panama Xcnitdontiicurtts xerraiux (Trombiculidae), .Mexico Artibcii-S phaeotis (Miller) fecomatiami xandoYtdi (Trombiculidae), Mexico* Artibeus toltecus (Saussure) C/i/r«y,woiWex £Yj{tsii (Spiniurnicidae), Mexico** Artibeifs u atsciiij Thomas PaniH-k hobiifs towei (Sirchlidae). Panama Kraeliyplivlla eavemariiiii Gray UinremctHatpHs w/(fvo7ir^s(l.abidocarpidae). Puerto Rico Ltiwrciu i tu tifpit?! ptn'rti>r(ct'n.'ro.\ (lusei f.ArgasidaeX Guadeloupe** and .Martinique RiuHofiiicUa inu!(’nuin.\! (Macronyssidae), Puerto Rico Trk fiohius !ruf!< iiiu\ (Sireblidae), Puerto Rico BraehypJiylhi nana Miller liiitiHsbohi'kin ci'nn'f (Myobiidae). Cuba s\Itu t<}>iys.\ok(c.'i ktx'hi (Macronyssidae), Cuba ()rf!ifh.\ a:feci (.Argasidae), Cuba ()r/tiilu)d(x <)x vipneru.'ii ( Argasidae), Cuba Pcriy'fisvhrus cifhattu.s (Spinturnicidae), Cuba Trichtthinsjieiinens (Strcblidae). Cuba Braehypliylia puniila Miller Ti ichithius freijuens (Streblidae), Dominican Republic Carol Mu sp. licciiiierellti lietda.'ieufa (Trombiculidac), Costa Rica* Ciiirfiyswules eiiridiktv (Sarcopiidae), Panama //oiiperelUt vesperupinis (Trombiculidae), Panama Ltxxijtskt afeiflume {Trombiculidae). Venezuela ijm/uiski ilextiKnitix (Trombictil idae), Venezuela /.oo/nf.v/n yitnkcri (Trombiculidae). V'enezuela Ornithodoros oc/et / (Argasidae), V'enezuela OrniduHlortn hrtxiyi (Argasidac), Venezuela OriiiiSttfdoros /noe/(Argasidae), Venezuela Sindila (Streblidae), Surinam Whdrfoitiii /auht.u’tosa (Trombiculidae), Costa Rica* Carcdlia Itrevicauda (Schinz) A /f/ A n/f n orp7/b r/nf/ /u (L a b i d rx' a rp 1 d a e). V e n ez. u e I a Afiihtyonuiia sp, (Ixodidae). Venezuela Eufrotnhictifa ^x>eidii (Trombiculidae), V'enezuela Eiffnfffthictda paaie, (Trombiculidae), Venezuela Ptimkoxii ftidtiriild (luibidocarpidae), Venezuela Carolina eastanea H. Allen Httoperella vespempittis (Trombiculidae), Nicaragua* Lootiii.siii i/cj.vjna/j'o (Trombiculidae), Costa Rica* Ltit>/ni.\iii spitH ssi (Trombiculidae), Nicaragua* Riidfhrdiefhi edFdlikte (Macronyssidae), Panama Speiseria ont/nX'no (Sireblidae), Panama SpeleiH ida xeeifuda (Trombiculidae), Nicaragua* Spekieorhynehus prm’enrMtr (Spelaeorhynchidac). Mexico Snehlit eoro///(/£'(Streblidae), Panama Tr ichoh i n ,v ji )h / / /if> ((SI re b I i dae), Pa n a m a Carollia perspicillala (I.tnnaeus) AfahidiHarpus furnttifii (Labidocarpidae), Venezuela Alcxftiinki eiiihinyeferis (Trombiculidae), Panama litOKOGY OK THE PHVIJ.OSTOMATIDAE JOl Afuhlyofniiw .santut/nvtni.'i (Sarcoptidae), Surinam Chirny.s.sikclcs zn/tderyensis (Sarcoptidae), Surinam Dc/titHh'x c-i[(ro///uc (Demodicidac), Surinam Di intHiex ixsimus (Dumoiikiddc), Surinam Hooperelia (Trombiculidae), Nicaragua*. Surinam, and Trinidad Lun rctti ecKttrpfis /obn.v (Labidocaipidae). Nicaragua Ijxtmisid (Trombiculidae). Colombia. Nicaragua*, Trinidad, and Venezuela Musttfptera yuifiwraesi (Slreblidae), Panama Mei^istoptnla aranea (Streblidae), Panam:i Mcielasmus pseiuk>pierus (St rebi idae), Panama .^yctet iucistcs priruus (Trombiculidae), Venezuela i\ycteroph(iia paraelli (Slreblidae), Panama Orui/ht)di>rox azieci (Argasidae), Panama (Canal Zone) Or/jiihodaros hrodyi (Argasidae), Panama and V'enezuela Orahhodaros /in.vcPf Argasidae), Venezuela Ontithadaros yu/tHUi'/ssis (Argasidae), Venezuela Faraiahidoearpux carol!sae (Labidocarpidae), Surinam r£/if 'o.i( 7i o £7ie £rvf /u //t c.t'UAVA'ru A ( T rom h i c u H dae), Panama Farasrk liohias loapicnts (Slreblidae) Parichoronyssifs £ ra.¥.v//7Ci: (M acronyssidae), Panama Pcraicx atittphtkahua (Trombiculidae), Trinidad P£7'/!,'//ir (Spelaeorhynchidae), Brazil, Colombia, Mexico, and Venezuela Sp£d£7>t7N> ££i/-£j///£i£'(Erey net idae), Surinam SpeU'oeofa secitttdu (Trombiculidae), Surinam Strehla alnnani (Slreblidae), Panama Sirehla rara/Z/rn'(Slreblidae), Brazil. Colombia, Panama, Trinidad, and Venezuela Sirehla comocias (Slreblidae), Trinidad S/rehIa mirahifis (Slreblidae), Brazil, Panama, and Trinidad T(ichobioides per.vpkvV/u/io (Slreblidae), Brazil, Mexico, and Peru Trtcliohitis costaliniai (Slreblidae), Panama Trk iiohitis depesli (Streblidae). Costa Rica, Nicaragua, Panama, and Trinidad Tricliohias dttpesiokUs (Streblidae), Panama 102 SPECIAL PUlUJCA'l JONS MUSEUM I EX AS TECH UNIVERSIIY TtjohiifiaHSyvM'ul'A^), Brazil, Belize. Colombia, Costa Rica, El Salvador. Guatemala, Panama, Peru, Surinam, Tobago. Trinidad, and Venezuela Tr ivht>h itt v joh n m >/uic (S I rebI idae), Pa nama Trk hohmx iofi}>ipi‘s (Streblidae), Panama Trichohiu.s /}UirFi)phyHi (Streblidae), Panama Irk'hohisix paraxifiais (Streblidae). Trinidad Trkhohtiix ,vpfif.17/v (StreblidaeI, Panama Trfi/u/bfia ynnAt rf (Streblidae). Panama WharfdHia timiosctom (Trombiculidae), Costa Rica*, Guatemala, Mexico, Nicaragua, and Trinidad Caredlia subrufa (J-Iahn) Chin}ysx<>kit’i airoHitic (Sarcoptidae), Panama HtHfpirelfa vespentginis (Trombiculidae). Mexico* and Nicaragua* Lttofuixia dcMnotliis (Trombiculidae), Nicaragua*, Panama, and Venezuela Spi isi’riu umhi^oui (Streblidae), Panama Spe!eoc<>ht seattnhi (Trombiculidae), Nicaragua* Strehia curoZ/wt’(Slrebltdae), Panama Trk'hohuixjohlinf^i (Strebl idae), Panama hart on in /iinhiseiOM (Trombiculidae), Mexico* Centurio seiiex Gray Bu.siiiti sp. (Nycteribiidae) ChirofftTina sp, Pioii/fisthms ihcfin^i (Spin turn icidae), Venezuela Chiroderina salviiii Dobson Cfiiniys.sihdes vapnrii (Sarcoptidae), Panama Or/i!thoilort>.s hnwi (Argasidae), Venezuela Piirtkrk ludfiifs (Streblidae), Panama PefiiiHsc hrttx iherin^i (Spinturnicidae), Panama and Venezuela Chiroderma vitlosiim Peters Afiihlyootfiia sp. (Jxodidae). Venezuela Aspidoparn huseki (Streblidae), Panama Piinifrk hohii(.\ sp, (Streblidae), Panama Trichohiu.s jt/hlinpi [SircMid^iCh Panama Clioeroiiiscus itiiiior (Peters) At)}h/yi>/ijn}(i sp. (Ixodidae), Venezuela Choeronycteris inexk'una Tschudi Paniirk hohftt.s (murk anux (Streblidae), Arizona Trkdudfiiix hmyipex (Streblidae), Arizona IVhtiffiinifi plcnni califor/ik ux (Trombiculidae), Mexico* Chrotopteruv aurihis Peters Btixilkt huphxiotfi (Nycteribiidae). Brazil ITiopendla vespernpinix (Trombiculidae). Nicaragua* Or/siihodonix hrodyi (Argasidae). Mexico Sfri’hla (Streblidae), Brazil Trkhohiitx iisfpi’xiofdcx (Streblidae), Panama Desmodus rotundits E. GeofTroy St.-Hilaire titisilki ferrixi (Nycteribiidae), Venezuela Chiniyxxoidcx caporti (Sarcoptidae), Panama Enduxhtihekkt arpanai (Myobiidae), Mexico Httopendfa stu copscry.\ (Trombiculidae), Trinidad Moopendit! vixpentpinix (Trombiculidae), Nicaragua* and Trinidad Loo/uow di'.v^norki'.v (Trombiculidae), Guatemala, Nicaragua*, and Venezuela Macronyxsaklex km hi (Macronyssidae). Brazil and Trinidad H[OLOGV OF THE PHYLLOSTOMATIDAE \ 0 } itm/iea (StrebliUae), Panama Micron o/nhit iiht hiinai (Trombiciilitiae), Mexico* Nycterimtstcs primux (Trombiculidaejf, Venezuela OrniiluHioros ciztcci (Argasitlac), Mexico, Panama, and Trinidad Oniifhotioros pernvkitins (Argasidae), Peru OrnithifJorox ytt/tmic/isis (Argasidae), Mexico PitraftihkhK'arpus dexmacliis (Labidocarpidac), Surinam Fiirusc(>sc!uH‘ftg(cs[i(i mcpasiyrax (Trombiculidae), Trinidad Pdi'asci'ia Ifotgiailcar (Trombiculidae), Trinidad Pcriite.H nnophtludota (Trombiculidae), Mexico and Panama Pi'rigfh(hru\ hcrrerai (Spinlurnicidac), Panama. Trinidad, and Venezuela Pcriglhcht us iheriiigi (Spinlurnicidac), Mexico and Panama Pcris.wptilla cxfutnuitifs (Trombiculidae), Trinidad Pc r/.v,wpu //u pre rur n; (T rom b i c u I i dae), Tr i n i d ad RDiifoniicifd iicstinnli (Macronyssidae), Panama and Trinidad RaiffordicKu oHdemanxi (Macronyssidae). Brazil Speixeriii a/uhignu (Streblidae). Panama Spclcocola dni’rvi(Trombiculidae), Mexico Speieocohi scattido (Trombiculidae), Trinidad Strchid (Streblidae), Panama Sindda diphylluc (Streblidae), Guatemala Strchia itcrrigi (Streblidae), El Salvador and Panama Sirchhf /nirubdis (Streblidae), Peru and Trinidad Strchla vciedeoHsntu (Streblidae), Colombia, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Panama, Peru, Surinam, Trinidad, and Venezuela. Tea) mat how sandovidt (Trombiculidae), Mexico* Triciufhioidc.s pcrspicilhiitis (SirtbUdsLO), Panama and Trinidad Trichohios vosudiftHii (Streblidae), Panama Trk'hohiuxdtigesii (Streblidae), Trinidad TricItiddus furnHi/ii (Streblidae), Peru Streblidae), Panama and Trinidad Trkhohius parasiik'ifs Brazil, Colombia. Costa Rica. El Salvador, Guatemala, Mexico, Panama, Peru, Surinam, Trinidad, and Venezuela. Trkhohinx Nuf/umj/j (Streblidae), Panama IVlwrtioiiu /todoxeioMi (Trombiculidae), Mexico and Trinidad Diaeinu-s youngii (Jentink) /Vytvmoj'y.v'rj'/.v desoiodics (Macronyssidae), Venezuela Pcrissopidla cxfuioutius (Trombiculidae), Trinidad Rodfordielki imdetminsi (Macronyssidae), Trinidad Strehia sp. (Streblidae), Trinidad Strehia dkuoni (Streblidae), Colombia and Panama Trichohinx dugesii (Streblidae), Trinidad Triefiohittxpiirasitfcux (Streblidae), Panama Diptiylla ecaudata (Spix) Sirehht i///j/iy//ue (Streblidae). Guatemala and Mexico Strchht minihiilx (Streblidae) Trichohius diphyilae (SlrebWdAeh Guatemala. Mexico, and Venezuela Trk'hohhis fiirnuoii (Streblidae), Colombia Trickohins punisiticHx^ Me%\co Enchistliieiies hartii (Thomas) Macrofiysxoidcx sp. (Macronyssidae), Panama i04 SPFXIAJ- PUtilJCAI lONS MUSEUM TEXAS rECH UNIVERSITY Fumkosii (l.abidocarpidae). Venezuela Farutrichohuis siiuvhezi (Streblidae), Panama and Venezuela Fvrsglisihnts (/u'n/j'.y/(Spinturnicidae). Panama and Venezuela Ti'k hohitis cUijiifsU (Sircblidac), Trinidad Erupliyllii bombirruiis (Millerl Ornithifdifros (Argasidae). Puerto Rico Ti k hohuis (Sirebliduc), Puerto Rico Eropliylla sezekorni Gundlach lAHKuhki tUwtniHius (Trombiculidae). Hahamas MicrotrofHhk siUi houefi (Trombiculidae), Bahamaii Fi‘nucs atu>ph!hal ;('I rom bicuI idae). Bahamas Fvripti.H'hnts vithanus (Spinturnicidae). Cuba Ti' k 'hohiti s fn- q mils {Si re b I i dae). Cub a Trk hohiifs inicnuediiis (Strehlidae). Cuba Trk hohiiis (Streblidae). Puerto Rico Wfidfiitniu tfiti'rrcrt'fisis (Trcmbicul idae), Bahamas Glovsopliaga sp. Oniithiuhiros pt^ruviauus (Argasidae), Peru Pt/fg/f.v(dou^.\ (Spinuirnicidac). Mexico and Panama Clossophaga ulticola Davis HiHipi ri flii vcspi’i tipiiiis (Trombiculidae), Mexico and Nicaragua Glossopliaga comiiiissarisi Gardner Jioopi'rdfa sui ctipferxw (Trombiculidae), Mexico’*' f!tH)pi‘i el!C(ifpiisJ'!fr/U(i/ti (l.abidocarpidae), Venezuela Anihiyoiittuii sp, (Ixodidae). Venezuela liiinuunhinilii (Trombiculidae), Venezuela I.iuifiiiskt ik'.\nuniti.\ (Trombiculidae), Venezuela Oi iiifhtHiorns iizti'i i (Argasidae), Venezuela Onik/hHkiros /iuac/ (Argasidae). Venezuela Onii{/nHli»i}s nnvHArgasidae), Venezuela Fiirakosa nuiximu (l.abidocarpidae), Venezuela f^amkosa huittritiii {l.abidocarpidaeJ. Venezuela Fe ri^ i isi-ii r us i v t lip //.v (S p i n t u r n ic i d ae), Venezuela Glovsophaga sorleina (Pallas) Alahicit/curpus ftuuitiui {] iibk\oi:iup\&Ao/n/.vjVf (Trombiculidae), Mexico, Nicaragua*, Panama. Surinam, and Venezuela Looinisiu sprocssi (Trombiculidae). Mexico* LoDiuiski u/iivuri (Trombiculidae), Mexico* Xfiu ronysstikiis Macronyssidae). Trinidad Xfii nXfoinhk ulii hatteti (Trombiculidae), Mexico* iSiycrei ifitisU’s prifiitis (Trombiculidae), Venezuela Oniitiiddortjs tizUTi (Argasidae). Venezuela Fifnaiysi'/iirki pun uloith's (Strebi idae) Fcirivhor<}nys.sus it /em.i (Macronyssidae), Panama BJOI.OGY OF THE FHYLLOSTOMATIDAE ]05 PfriiitiM hniii caUfiu.s iSp'mluTnlc\diie:), Brazil. Panama, Surinam, Trinidad, and Venezuela Prri.sA'opulla heitratii (Tronibiculidae), Mexico* Pfri.vsitpdlhi iwliNtnastts ('rrombiculidaei, Trinidad Pi'ri\sitpullu pn'iunit (Trombiculidae), Mexico Fsarer^aioides (Psorergaiidae), Surinam Speifii'i ioe (Slreblidae), Trinidad Spidai’orliynrfuis praeci/rsof (Spclaeorhynchidae), Aniazon(?) SpeletHohi duvisi {Trombiculidae). Mexico Spi'leoctiiu scrdfuld ('rrombiculidae), Nicaragua SfL’iifottyssttsjikiijuiifii (Macronyssidae), Brazil Stri'hia cfiTcd/iVa'(Streblidae), El Salvador. Panama, and Venezuela Sinhld itiirahili.s (Streblidae), Trinidad Tfic/iohius (hipcsti (Slreblidac), Colombia. El Salvador, Guatemala. Mexico, Panama, Peru, and Trinidad Tr/f/aJ /? / fzv/ r# ut7 /H (Si re b 1 i d a e), P a rag u a y 7/Streblidae), Panama and Trinidad Ti ichohius (dnpipts (Streblidae) TiicSud^ius (Streblidae), Costa Rica, Guatemala, Guyana, Mexico, Panama, Peru, and Venezuela Wdptdumrui .dniiHs (Trombiculidae), Mexico* /luiSi/si'toya (Trombiculidae), Mexico and Nicaragua* Hyloiiyctcris luulerwoodi Thomas iieLsilia rofiilufiii (Nycteribiidae) l.eplouycieris ciiraxoae Miller Anif wolti sp, (Argasidae), Venezuela ()i/iif(wiIort>s sp. (Argasidae), V'enezuela Leptonycleris itivalis Saussure c'orv/ror/H’nf (Nycteribiidae). Texas £//U£/e/i.v(Macronyssidae), Arizona Ontiihoii(tr<)s rtKsxi (Argasidae), Arizona /Vr/>/m7fno viupasi {Spinturnjcidae), Mexico and Texas RdJpiniii’ihi orictdii (Macronyssidae), Mexico Tr/r/fohhij; (Streblidae), Mexico and Texas LeptojiycterLs sanborni HofTmeisler Basilia tin(ri>zi>i (Nycteribiidae) Peripli.H hru.s varpusi (Spinturnicidae), Mexico Speleocoid Ji/v/j./(Trombiculidae). Mexico Tr/t /fohm.v xp/iuero/jofw.v (Streblidae), Arizona and New Mexico IJonycferis spurrelli Thomas M t fer/>i!(rv(e.v prinui.s (Trombiculidae), Venezuela Pi‘riplix( hrtis hopkifisi (Spinturnicidae), Venezuela Tra /nj/uV/.v//f^mertTfV/h (Streblidae). Panama and Peru Loitebophylla coneava Goldman Heddtt'relfd avdidM dfa (Trombiculidae), Costa Rica* LiH)/ui.\!(! xproc-v.v/(Trombiculidae), Costa Rica* LoncJioptiylhi rohustii Miller Addstrehlu dyeferhiix (Streblidae), Panama Aiuitricfutfyids xcorzui (Streblidae), Panama Chiroeiefex /cJ/n7ifjp/iy//« (Macronyssidae). Venezuela Eldu/ttiia /jfci'/t ep.GStreblidae), Panama Loodtixid Je.vmoi/jzi (Trombiculidae). Costa Rica* and Venezuela i\'ycierhmxtes .veriz/n/f;* (Trombiculidae), Costa Rica* PL>riplisi fin/.s (Spinturnicidae). Panama SrEC[Al. PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Speixi'riii anihi^’ua (Streblidae), Panama Strehla t'(iri>l!iiu‘ (Streblidae), Panama Ti U'hohiuxJi)hli/!!ii (Streblidae), Panama Trichohiusjohnsittuie (Streblidae). Panama Trichohius lo/tiiuiphyllm' (Streblidae), Panama Trichoh ins tin ifortit is (StrebI idae), Panama Lonehoriiina atirita Tometi Biisilia (Nycteribiidae). Venezuela /V.v t / tr f; i £ ,v p; j f/A (T r om b ic u I id a e), V e n ez u el a Onfir/WoroA ££.:/£'£ /(Argasidae), Cuba, Jamaica, Trinidad, and Venezuela Ornifiifulonts hrodyi (Argasidae), Venezuela Or/iilinHloros luisei (Argasidae). V'enezuda Ferif^'lischi tis ffntneroi (Spinlurnicidae), Venezuela PsorergtiiOkies lo/ichorltiiuie (Psorergalidae). Venezuela Speisi’fiii iifnhif>nti (Streblidae). Panama Sfrehici (ilituiini (Streblidae), Panama and Venezuela 5/ /■£7i /££ £ £mf j// f££ f (St re b I i d a e I. P a na m a Trk hohtiis (iiisesiokies (Streblidae), Panama 7r/£7j£j/f/f£A(Streblidae), Panama Trichohiits iiUicrophytli (Streblidae), Panama Ti k hohitis yunkeri (Streblidae), Panama Luiidiorhiita oriiioceusi.s Linares and Ojasii Otniihodoros sp. (Argasidae), Venezuela Ornititodoros heisei (Argasidae), Venezuela Ontithoitoros jwa/(A rgasidae), Venezuela \lacTophyllum macruphyllliim (Schinz) Hits ilia eofisrrictii (Nycteribiidae) Eairomhivtila variahilis (Trombiculidae), Venezuela OtnithoJoriKs azieci ( Argasidae), Venezuela Fitrnlahkioeaypns tnarrophyUifin (Labtdocarpidae), Surinam /Vn^7f.vr/rr;rv sp, (Spinturnicidae), Panama Strehla altniani (Streblidae), Panama Strehia tY£r£7//£££'(Strebl idae), Panama Trichohius johiift}>i {SlTt^bU&dch Panama Trichohius mticrophylii (Streblidae), Panama Macrotus sp. Whurtonia liiHht.setosa (Trombiculidae), Jamaica Maerotus caUrorniciis Baird LtHMiiski sprtH'ssi (Trombiculidae), California /V.vf r£'ro/)/ji!7af toAYjr/v (Streblidae), Arizona and California Otnitlunhiros sp. (Argasidae), Arizona Oniitluxloros r p li y 11 us c 11 ba n u.s Miller Eiidn.\luihi'kia ro^h kyi (Myobiidae), Cuba Fi riislixchni.'t yur^’tisi (Spinturnicidae), Cuba MoncipIiylJus rediiiani l^each RiulftudieUa nuitiapitylli (Macronyssidae), Cuba Speiacorhym htix /nmiophylU (Spelaeorhynchidae), Puerto Rico Tfkhohius n r/iyi {Streblidae). Cuba Trifhtfhiifx freqtii’itx (Streblidae), Cuba Ti'k hohiifx //j/c/’«n'(/hf.s (Streblidae). Dominican Republic Ti it luihiifs paraxidcu.s (Streblidae), Jamaica 7V/(7n'/tm.v roMvjfu'(Streblidae), Puerio Rieo Trk'hohiitx rnonar/i'S (Streblidae), Puerto Rico Ptiylloderma steiiops Peters SV r(7> fa t h rix t itut c (S t re b I i d ae), Pa n a m a PhylJonycteris apliylla (Miller) OrifiihtHktros hti.u’i (Argasidae), Jamaica*’'^ Pbyllonycteri-s peteyi Gundlach AfUricoki ttHiriiimuui (Argasidae). Cuba Autricoli! ,v//vni(Argasidae), Cuba Einhishithckki datticli (Myobiidae), Cuba Xhicfotiys.stfhlex Aor/ri (Macronyssidae), Cuba OrukhtHloros viptierusi (Argasidae), Cuba and Haiti"** Pcri}flixch/ ti\ ctihiiniix {Spinturnicidae), Cuba Trk hohiiix cer/iyi (Streblidae), Cuba 7Vn7fi(ts Huiudfitx (Streblidae) Pliyllostomu,s sp. Axpidopiiru phyIkKsio/uatis (Streblidae), Brazil Htisilki hi‘!liii‘dii (Nycieribiidae), Brazil Piisilk: speixeri (Nycteribiidac) Muxfoptt ra ffuiimtruesi (.Streblidae), Panama Mi'yixh.>pf>ilci arutH'ii (Streblidae), Brazil and Cuba Purichortffiyx.sus .v<(Macronyssidae), Costa Rica Strchhi i (Streblidae), Peru and Surinam Sfrehiii /Ji/m/n7;.v (Streblidae), Eirazil, Panama, and Peru lifUf^ipi'x (Streblidae), Panama 7>jf7io7)ira.v p/iyZ/f/v/o/nuc{Streblidae), Brazil BIOLOGY OK IHE PHYl.l.OSTOMATIDAE 109 Pliytloiitcmus discolor fV/e.v per-v/jk-y/^um.'i: (Streblidae), Colombia. Panama, and Trinidad Trh'hohkis coxnili/iitii (Slreblidae). Colombia, El Salvador. Guatemala, Panama, Peru. Puerto Rico, Trinidad, and Venezuela Tf ichohkis lo/iptpes (Slreblidae), Trinidad Ph>llostomu.s elongatus (E, Geoffroy St.’Hilaire) Liin remToirirpus phyllostoiiifis (Labidocarpidae), Venezuela PcripHschms uouj'/iveni/fv(Spinturniddae), Trinidad and Venezuela FscKdoakihidovarpus scais (Labidocarpidae), Venezuela Sir eh Id ti 1 1 ni h il is (St re bl i d a e) Trichifhlikdes perspk iihkus (Streblidae), Colombia Phyllosforims hast at iis (Pallas) Aldhklocitrpits phytloskiifii (Labid(,>carpidae), Surinam Blaukaiirthi sitituiitturyi (Trombiculidae), Panama Didiuiilex phyllosionuiiis (Demodicidae). Surinam Ealnimhicithi poeldii (Trombiculidae). Venezuela Maxtopteris piikiniraesi (Slreblidae), Colombia and Panama MdslopU ra mkiHia (Slreblidae), Colombia Mkroiici/iihicHlii bouefi (Trombiculidae), Panama Mkrotniinhk iiiu varwemic (Trombiculidae), Panama Mi u u c r eu n.v / {T ro m b i c u ] i d a e), V e n ezu c I a A/eofr/t7 j o/i i/iJi f/e/ic’i(r/iA (S treb 1 idae), Su rinam Ornkhodonis it zt it I {Arga.sidae), Venezuela Ot nithodoros hasei (Argasidae), Venezuela Fiircu'ucienodes lonpipcs (Slreblidae), Brazil Peripiisi hrus iictfiisu’t nifs (Spinturn'Kid^e)^ Colombia, Panama, Trinidad, and Venezuela Fi'riplischrus /o/TCfi/Au: (Spinturnicidae), Panama, Trinidad, and Venezuela Speistria atnhipiici (Streblidae), Panama Spi leoi hii phykosio/iii (Ereynetidae), Colombia Sirehlti aindliae (Streblidae), Panama Stri’bfd couior/ii.v (Streblidae), Surinam. Trinidad, and Venezuela SiiTbkt Iwriigi (Slreblidae), Costa Rica, Nicaragua, and Panama Strebki iitirubilis (Streblidae), Colombia, Panama, Peru, and Trinidad 7Vif7N)/>ioi£/e.v piTspiciUaius (Streblidae). Surinam Trichobiiis diigesii (Streblidae), Trinidad TrkiiobiiisJohlitipHStrchhduei}. Panama and Trinidad Trichobius (o/igi/fcj (Streblidae), Bolivia, Costa Rica, Colombia, Guatemala, Panamk Peru, Surinam, Trinidad, and Venezuela SPHCtAl. PU HI. [CATIONS MUSEUM I' EX AS TECH UNIVERSITY I 10 Trichahius paidstiicus (StrebI idae), Panama Ti ichohiiis phyllomondw (Streblidae) RJiinophylJa puiniMo Peters Periplisvhrtis hopkiftsi (SpintutnicidaeK Braj^il and Venezuela Fcripli.H'hrits rdi’rfni (Labidocarpidae), Puerto Rico Fi-ripHschrus (Spiniurnicidae). Puerto Rico Slurtiira liliiim (E. Geoffroy St.-Hilaire) Aspuiopiem (Streblidae), Guatemala and Panama A'spiclopU'ni phyllostomafis (Streblidae), Paraguay ChirnyxsoHh’x fSarcopiidae), Brazil Eiiilusbiihekid Icpkiosetu (Macronyssidac), Nicaragua Eiitiotnfyit tih; ptn'liiii (Trombiculidae), Venezuela Exasiittiati r/ortY/(StrebIidae) Hoopen'Ita veipet upiais CPrombiculidae), Nicaragua"* /-ViWe.v sp. Ilxodidae), Venezuela Lu e (Rose n s t e i n i id a e), B ra z i I Ornkhoilfxos iip. (Argasidae). Venezuela Ornithiniorox hasei (Argasidae). Venezuela Farakosd {acidritid (Labidocarpidae), Venezuela Fdruitihkhfcarpdx drtihei (Labidocarpidae), Nicaragua FcripUschrds then tipi (Spinturnicidae), Mexico Fcriplixi hrus ojd.sti (Spinturnicidae), Panama, Trinidad, and Venezuela Fdnpiixchrtix vargtt.si (Spinturnicidae), Mexico Tfiehifbioidex perxpii ilia lux (Streblidae), Panama Sfurnira luduviei Anthony IX Oil ex up. (Ixodidae), Venezuela Mepistopodd pmrbnu {Streblidae), Costa Rica \fepi.siopoda //teix/fu/(Streblidae), Panama Microirotuhiculd curdtetme (Trombiculidae), Costa Rica and Panama ;V/krotrom/jfcfdu srifrmrae (Trombiculidae). Costa Rica and Panama Orftithitdftrox /ru.iei (Argasidae), Venezuela Fttrasecid xduranyatui (Trombiculidae). Panama Parichoro/tys.xtix eiiihyxter/tum (Macronyssidae), Panama Feriplixchntx //ler/ni?/(Spinturnicidae), Colombia and Venezuela Feriplixchrttx ojdxti (Spinturnicidae), Panama Fxeudoxchoeftpaxitd huihifem (Trombiculidae), Panama Trk'hohiiis hretida/ti (Streblidae). Panama 7r ff/i o /u' f/ i y »tj A e r I {S t re b 11 d ae), Pa n a m a Sturnira niordax (Goodwin) Mieroindtihictdd cur/nefuie (Trombiculidae), Costa Rica Microirofnhk tda i7/ir;7t;>ue (Trombiculidae), Costa Rica Sturnira tildae de la Torre A/uhfyodimd sp. (Ixodidae), Venezuela Tonal ia sp. Mmtapterd (Streblidae), Bolivia, Colombia. Ecuador. Peru, and Surinam BIOLOGY OF THE PHYLLOSTOMATtDAE in Sfiz-ostrehla lougirostris (Streblidae), Brazil and Colombia Ssrehfa szalifuloi (Streblidae), Trinidad Sireh/t! >Nira/nVn (Streblidae), Colombia Tonatla hidens (Spix) Spi‘isi‘i ia umhiiiua (Streblidae) Strehfii fzafi/uioi iSteb\idm]y Panama Sfrehlti ifiifdhilis (Streblidae) S/n^hhi iDtuiiiae (Streblidae) Trichohius (Streblidae), Panama Toiiatia brasiliense (Peters) SovWa row;/fw (Streblidae), Ecuador and Panama Tonatia iiicaraKiiae Goodwin Masioptcra (Streblidae), Panama I^M'iidosirehlii gi ecnwetli (Streblidae), Panama Ssrehht hoogstraulHSirtbli&dc), Panama Trichohius /Henc/e-ci (Streblidae), Panama Tonatia sylvicola (D'Orbigny) Basilia coftsn k ta (Nycteribiidae) Mastoptera m//mm (Streblidae), Brazil and Panama Omiihodoros (Argasidae), Panama P\etid(fsirehla nTie/ro/(Streblidae), Brazil and Panama Strehia kohixi (Streblidae), Colombia and Panama Trkhohias dyha.si (Streblidae), Panama Trichohiusjohiiugi (Streblidae). Panama Trichohius parasiiicus (Streblidae), Brazil Tonatia veneziielae (Robinson and Lyon) Chirnyssoides venezuciac (Sarcoptidae), Venezuela Partdahidocarpus tonufiue (Labidocarpidae), Venezuela Trachops sp. Strehhi co/tjornw (Streblidae), Peru Traehops cirrhosus (Spix) Loofuisiu desmodus (Trombictilidae), Mexico* Orniihodoras^p, (Argasidae), Venezuela Oruifhodoros azteci (Argasidae), V'enezuela Orukhodoros hrodyi (Argasidae), Panama Paralahkiocarpus tnichi/ps (Labidocarpidae), Surinam Periglischrus parucutisternus (Spinturnicidae), Venezuela Periglischrus vurgasi (Spinturnicidae), Panama Speiserki amhigua (Streblidae), Panama Strehhi alt muni (Streblidae), Panama Strehia raro/Z/V/e (Streblidae), Panama Strehhi diphylliie (Streblidae), Guatemala Trichohius dugesii (Streblidae), Panama Trichohius dugesioides (Streblidae), Panama Trichohius johlingt (Streblidae), Panama Uroderma bilobatnm Peters Aluhtdocurpus nicaraguae (Labidocarpidae), Nicaragua sp. (Ixodidae), Venezuela BasilUi consirichi (Nycteribiidae) Btisida niyotis (Nycteribiidae) Chirorhynchohki urodermae (Chirorhynchobiidae), Panama £///rof/i/j/c7/(£j fr/j/u/nji (Trombiculidae), Trinidad Macronyssoides sp. (Macronyssidae), Panama Neotrichohius deUcuius (Streblidae), Panama 112 SJ’ECJAl. JHJUl.lCA'nONS MUSEUM TEXAS TECM UNIVERSITY Oinis/tiHloros hitsei {ArgasKlaeK Paiiami'i Ftirasccui /imtmi’li (TrunibiculidacK Costa Rica* Fiti-tttriclufhi{t.\ ilmtiti (SirebSidae), Panama Ftirtih iclnthitis foni^’ifrtis iStrcblidac ! Ft’ri^i>li.stiir;f\ ihctint^’i (Spintiii nicidae), Guatemala. Panama, Paraguay, and Venezuela Pliyltti\tnit!innys.sii\ conradyunkcri (Gasironyssidae). Surinam 7) i( iiohiii.s nntiiliinai (St rebliiiae), Panama 'rfivhtihiusjolylinyi (Streblidae). Panama I'livhohiux L('i‘/}(uti (Sireblidae), J’anama Tricht}h!iL\ //rodenni/cMStreblidae}. Panama and Venezuela IJrodtTiiia miigniroslnim Davis AhihiiitH'arpiiM tuihidocarpidac). Venezuela EiHliishaht kltt /rroi/enut^iMMyobiidae). Brazil OrniihiHhfros /niM’i (Argasidae), Venezuela Vainpyressa iiyinpliiiea Thomas Aspiii(tp!cni husvki (Streblidae). Panama .XU'ii’hiMtiifs p\vtHtop(cfn\ (Streblidae). Panama Vampyressa pusilta (Wagner) Cliirny.ssoidi s i npartt (SareopI idae), Panama Xhu rtynyssoiihx sp, (Macronyssidae), Panama Af'o/ne/nj/juo (/e/hfij'/rv (Streblidae). Panama and Venezuela Pi'iifytisi hrfLt>uU'.s (npiii ti (Sarcoptidae), Panama hirichi>nnty.\su\ sp. ( Macronyssidae). Panama Pc riiiH.schnLS ihi'ri/ipi (Spinturnieidae). Panama Spt'kun hh /)r(j.v/7/>n.vA (Ereynetidae), Brazil Vainpyrops sp. Pcf ipiiM hnis ihetiu^'i (Spinturnieidae). Paraguay Vaiupyrops dorsalis Thomas p£'n,e//\(7jnc-v //le/m.ei (Spinturnieidae). Venezuela Vampyrops lielkri Peters Ahihiiirt}x /!££«'/ (Argasidae), Panama Puruirh fiohins ^p. (Streblidae), Panama P('ri}*!iM hrns (Spinturnieidae). Mexico and Panama / Vj y // f ).iV£ j / >/j y v.v (£ A to // ru dy £/ £ j A er / (G ast r 0 n y ss i d a e), S u r i n a m Vampyn>ps liiveatus E, Geoffroy St.-Hilaire Mt’pisiopditt! pihitei (Streblidae), Brazil. Cuba, Mexico, and U.S.A. Ptiranichohiu.s (Streblidae) Pcriglisch nis ih enXe/ (Spi n lu rn icidae), B raz il Sirehlft n ii‘di‘fnatini (Streblidae), Brazil Vampyrops vittaliis Peters C/nnrvvAc>n/e-v cupidii (Sarcoptidae), Panama LtK>/}ii.sin f/t'.o/jot/iv.v (Trombiculidac), Costa Rica* M(i< r(iny.'i\t>Uh‘\ ronsiliaitis (Macronyssidae). Panama Paruirk hohii(\ %p. (Streblidae), Panama Sp t' ;t7 (oj t h w f/ fM SI re b I i d a e). Pa n a m a Trichohiu.\ T’tt/upvTfJ/jf.v ( Streblidae), Panama and Venezuela BIOLOGY OF THE PHYLLOSTOMATIDAE 113 Vinnpyruin spcclruin (Linnaeus) HiH>pen>iii( (Trombiculidae), Panama and Trinklad Panisi chi ion^ikaU ar (Trombicul idae), Panama Tyk hohiua purusiiicns (Streblidae) AcKNOVV'LEDGMENTS Our thanks are given to the many acarologists aiioniisia gen. n. with descriptions of three new^ Venezuelan species (Acarina: Trombiculidae). J. ParasitoL, 58:796-800. -. 1973. More new' genera and species of chiggers (Acarina; Trombiculidae) from Venezuela. J, ParasitoL, 59:706-710. SPECIAL PUHL!CAT[()NS MUSEUM I EX AS TECH UN[VERS1TY J 14 --. 1974. The genu?; Eini-Dnjhivnia in Veneiiuela (Acarina: Trombiculidae). J. Parasitol., 60:699-7H. __. 197S. A list of Venezuela chiggers. particularly of small mammalian hosts (Acarina: TromhicuHdae). Brigham Young Univ. Sci. Bull., Biol. Ser., 20:45“?.^, Bbe-nnan, j. M.. and C. E. Yunker. 1966. The chiggers of Panama (Acarina: Trombi- culidae). Pp. 221-226, in Ectoparasites of Panama (R. L.. Wenzel and V. J. Tipton, eds,). Field Mus. Nat, Hisi., Chicago, .\ii+ 861 pp. Cl ARK, G. M. 1967. New Speleognathinae from Central and South American mammals lAcarina, Tromhidiformes). Proc. Helminth. Soc. Washington. 34:240-243. Ci.ieeord, C. M., G. M, Kohls, and D, E. Sonensmink. 1964, The systematics of the sub¬ family Ornithodorinae (Acarina: Argasidae). I. The genera and subgenera. Ann. Enlomol. Soc, Amer.. .^7:429-437. C(x>i EY, R. A., AND G. M. Kohls. 1944. fhe Argasidae of North America, Central America and Cuba. Amer. Midland Nat. Monog., 1:1-1.^2. Cracraft, j. 1974. Continental drift and vertebrate distribution. Ann. Rev. EcoL SysL, 5:215-261. Cruz, J, de la. Noias sobre las garrapatas del genero Anirivohi Cooley y Kohls, 1942 ([xodiformcs, Argasidae) con la descripcion de una nueva especie. Acad. Cienc, Cuba, Ser. Espel. Carsolog., 44:1-13. Cruz, J. de la, J. R. Iamsitt, and D. Valdivieso, 1974, Three new species of labldo- carpine mites (Listrophoroidea. Chirodiscidae) from Puerto Rican bats. Life Sci, Occas. Papers, Roy. Ontario Mus., 23:1-13. Desch, C., R. R, Llbel, W. B, NuniNu, and F. S, Lukoschus, 1971, Parasitic mites of Surinam. I. Dt'/tiodex cttroiliue sp. nov. (Acari: Demodicidae) from the fruit bat enrollki perspicillntti. Parasitology, 62:303-308. Desch, C, W. 8. Nu n iNG, and F. S. Lukoschus. 1972, Parasitic mites of Surinam. VII. Di’/tiodi'X lo/igL\si/t}us n. sp. from Cnndlla perspicHlata and D. moiossi n. sp. from Moiossus ntohxxns (Demodicidae: Trombidiformes); Meibomian complex inhabitants of Neotropical bats (Chiroptera). Acarologia, 14:35-53. DusiuitEK, F. 1967«. Jiimesonhi. a new' genus (Acarina: Myohiidae) with seven new species from Cuban bats. Folia Parasitol., (Praha), 14:247-261. -—. 19676, To the knowledge of miles of the subfamily Nycieriglyphinae (Acarina: Rosensieiniidae) from Cuba. Folia Parasit. (Praha), 14:239-246. -, (969. Generic revision of the myohiid miles (Acarina: Myohiidae) parasites on bats. Folia Parasit. (Praha), 16:1-17, -. 1970. New records of parasitic mites (Acarina) from Cuba and Mexico. Mitt, Zool. Mus. Berlin, 46:273-276. Dlsbabek, F,. and F. S. I.ukosc hus. 1971 Mites of the genera Eyvinga/in and Ugiouiohia (Acarina: Myohiidae) from Surinam bats. Parasitic mites of Surinam. XVL Folia Parasit. (Praha), 18:337-345. -, 19716. Some Spinturnicidae (Acarina: Mesostigmata) from Surinam bats. Parasitic mites of Surinam. IX, Folia Parasit. (Praha), 18:149-1.54. -. 1973. Parasitic mites of Surinam. XX111. kfanndti tnarhu\ new genus and new species (Acarina: Myohiidae) from a phyllostomid bat. Zool. Anz., Leipzig, 191:108-113. Fain, A. 1956, Une nouvelle famille d'acariens endoparasites des chauves-souris: Gastro- nyssidae Fani. nov. (Acarina: Sarcoptiformes), Ann. Soc. Bel. Med. Trop„ 36:87-98. —--, 1959. Les acariens psoriques parasites des chauves-souris. X, Le Genre Chirnys- .u)ides g. n. chez les chauves-souris sub-americaines (Sarcoptiformes: Sarcoptidae). Bull. Inst, Roy. Sci. Nat. Belgium, 35:1-19. --, 1960. Les acariens psoriques parasites des chauves-souris. XIlI. I^a famille Demodicidae Nicolet. Acarologia, 2:80-87. -. 1962. Les acariens psorique parasites des chauves-souris. XXIL Especes des genres ChirnyssoUies et Noutedres. Bull. Ann. Soc. Roy. Entomol. Belgium, 98:392-403. ElfOLOGY OF THE PHYLLOSLOMAUDAE 115 -. 1963. Les (yroglyphldes commensaux des chauves-souris insect ivoiX’s-De' script ion de cinq espcces nouvelles. Rev. Zool, Hot. Africa, 67:33-51!!. -. 1966. L’acariase des fosses nasales chez les mammiferes. Description d'une nouveile espece chez une chauve-souris de Trinidad (Ereyneiidae: Trombidi- formes). Acta Zool. Pathol. Aniverpiensia. 41:103-108. ■-. 1967. Diagnoses d'acariens Sarcoptiformes nouveaiix. Rev, Zool. Hot. Africa, 75:378-382, -, 1970n. Notes sur les speleognathines parasites nasicoles des mammiferes (Ereynetidae: Trombidiformes). Acarologta, 12:509-521. -1970/n Parasitic mites of Suriname. 111. Diagnosis of new lisirophorids. Bull. Ann. Soc. Roy. Entomol, Belgium, 106:175-180, --—. 1970c. Un nouveau genre et une nouveile espece dans la sous-famille Rod- hainyssinae (Sarcoptiformes: Gastronyssidae). Acaroiogia, 12:160-163. -. 1972, Diagnoses de nouveaux Myobiidae (Acarina; Trombidiformes). Rev. Zool. Bot, Africa, 86:148-157. Fain, A., and T. H. G. Aitken. 1969. Acariens nasicoles d’oiseaux et de mammiferes du Bresil, 11. Ereynetidae de la region de Belem (nord Bresil). Bull. Ann. Soc. Roy. Entomol. Belgium, 10.5:33-44, -. 1970. Acariens nasicoles d'oiseaux el de mammiferes du Bresil. IV. Nouveaux Ereynetidae (Trombidiformes) et Turhinoptidae (Sarcoptiformes) de la region de Belem (nord Bresil). Acaroiogia, 12:326-338, Fain, A,, AND F. S. Lukoschus. 197Hn Parastic mites of Surinam. XV. Nasal ereynetid mites of bats with a key to the known species (Trombidiformes). Bull. Ann. Soc. Roy. Entomol. Belgium, 107:284-297. -. 19716. Parasitic mites of Surinam. XVIII. Mites of the genera Vofoct/rc.v and ChirnysMtideM from bats (Sarcoptiformes; .Sarcoplidae). Bull. Ann. Soc. Roy. Entomol. Belgium, 107:298-313, -. 1972. Parasitic mites of Surinam. XXVll. Gastronyssidae. Rhinonyssidae and Ereynetidae from the nasal cavities of bats or birds. Stud. Fauna Surinam and other Guyanas, 13:84-94. Fain, A., and P. H. Vercammen-Grandjean. 1953. Les fosses nasales. nouveile localisa¬ tion parasitaire des larves de Thrombidions chez les rongeurs (Note preliminaire). Ann. Soc. Belg. Trop. Med., 33:41-42. Fain, A., G. Anastos, J. Camin, and D. Johns ion. 1967, Notes on the genus Spidat'o- rhytu hns. Description of S. pmecursor Neumann and of two new species. Acar¬ oiogia, 9:535-556. Faircmii.d, G. B., G. M. Kohls, and V. J. Tipton. 1966. The ticks of Panama (Acarina: Ixodoidea). Pp. 167-219, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.) Field Mus. Nat. Hist., Chicago, xii+ 861 pp. Ferkis, G. F. 1924. The New World Nycteribiidae (Dipiera Pupipara). Entomol. News, 35:191-199. Gekberg, E. j., and F. C, Goble. 1941. Tw'o unusual records of Mallophaga from bats. J. Mamm., 22:454. Greenhaum, I. F„ AND C, J. Phili ips. 1974. Comparative anatomy and general histology of tongues of long-nosed bats {Ltpionycierix Sitfihorni and L. nivalis) with ref¬ erence to infestation of oral mites. J. Mamm., 55:489-504, Guimaraes, L. R. 1944. Sobre os primeiros estadios de alguns Dipteros Pupiparos. Pap. Avul. Dept. Zool, Sao Paulo, 6:181-192. -. 1946. Revisao das especies Sul-Americanas do genero Basil ia (Diptera- Nycteribiidae). Arq. Zool. Esi. Sao Paulo, 5:1-88, -. 1966. Nycieribiid baiflies from Panama (Dipiera: Nycteribiidae). Pp. 393- 404, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.). Field Mus. Nat. Hist., Chicago, xii-l-861 pp, -. 1968. A catalogue of the Diptera of the Americas south of the United Slates. Family Nycteribiidae. Dept. Zool,, Sec. Agri., Sao Paulo, 101:1-7. SPECIAE- PURLICAT[(>NS MUSEUM TEXAS TECH UNIVERSITY I 16 -. 1972. Venezuelan nycteribiid badlies (DipiLTa: Nycieribiidac). Brigham Young Univ. Sci. Bull., Biol. Ser.. 17:1-1 I. Glimaraes, L, R„ and .VI. A. V. D'Andretta. 1956. Sinopse do.s Nycteribiidae (Diptera) do Novo Mondo. Arq. 7.ool. Esl. Sao Paulo, 10:1-184. Ham , E. R., and K, R, Kei son, 1959, fhe mammals of North America. The Ronald Press Co., New York, 1:xxx+ 1-546 4* 79 pp. Herrin, C, S.. and E. J. Radovskv, 1974. Vcne/iielan Macronyssidae. 3. Chirovcae.s loiH liopiiylhu n. g., n. sp,. from a long-tongued bat, J. Med. Entomol., 11:747- 351. Herrin, C. S., and C. E. Yunker. 1975. Systematics of Neotropical Hintionyssns mites with special emphasis on Venezuela (Acarina; Mesostigmata). Brigham Young Univ, Sci. Bull., Biol. Ser.. 20:97-127. HtiFfMANN, A. 1972. Dos ectoparasitos de murcielagos poco frecucnles en Mexico. Rev, Lat.-Amer. Microbiol., 14:191-195. HortMANN, A„ AND 1. B. DE Bakrera. 1970. Acaros de la Tamilia Spelaeorhynchidae Rev, Lat.-Amer. Microbiol., 12:145-149, Hot r MANN. A., I. B. DE Barrera, and C. Mendez. 1972. Nuevos hallazgos tie acaros en Mexico. Rev. Soc, Mex. Hist. Nat,. .77:151-159. Jameson, E. W., Jr. 1971. Comments on Etulif.sht/hi'khi, new name Dusbabek, 1967. preoccupied), with two new species (Acarina: Myobiidae). J. Med, Entomol. 8:517-518, Jepsen, G. L. 1966. Early Eocene bat from Wyoming. Science, 154:1777-1779. JoiiMNG, B. 1949. Host-parasite relationship between the American Streblidae and the bats, with a new key to the American genera and a record of the Streblidae from Trinidad, British We.st Indies (Diptera), Parasitology, 39:315-729. Jones. E, K., C. M. Clif tord, J. E. Keirans, and G. M. Kohls. 1972. The ticks of Vene¬ zuela (Acarina: Ixodoidea) with a key to the .species of Atuhivinnttm in the Western Hemisphere. Brigham Young Univ. Sen. Bull,, Biol. Ser., 17:1-40. KiNtjstON, N., B. Viii.a-R,, and W. Lorez-Formem . 1971. New ho.st and locality records for species of the genera Pi riyfi.schni.'i and ( Acarina: Spinturnicidac) on bats from Mexico, J. Parasitol., 57:927-928. Koiu s, G. M. 1957. ImhE'x douni.v/, a new species of tick from a cave in Trinidad, British We.st indies {Acarina-Ixodidae). Proc. Entomol. Soc. Washington, 59:257-264. Kimi.s, G. M,, and C. M. Ci.ifeord. 1964. Orniihiniorox iAU'cforohiiix) h(fiiriensis sp. n. (Acarina: Argasidae) from bats and hou.scs in Bolivia. J. Parasitol., 50:792-796. KOI I IS, G. M., D. E. Sonensuine, and C. M. Clifford. 1965. The sy sternal ics of the subfamily Ornithodorinac (Acarina: Argasidae). IL Identification of the larvae of the Western Hemisphere and descriptions of three new species. Ann. Entomol. Soc. Amer., 58:331-364. 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A new species of Jamesofua Dusbabek 1967 (Acarina, Trombidiformes, Myobiidae) parasitic on the vampire bat, Dismotlus nHn/nins, with a description of the nymphal and larval stages. Accad. Naz. Lincei, 171:157-173. Webb, j. P.. Jr., and R, B. [jx>mis, 197L Trombiculid mites of the genus Micrtfiro / rj - hiatlii (Acarina) from Costa Rica, Contrib. Sci., Los Angeles Co. Mus., 207:1-1.*), Wenzel, R, L. 1970. A catalogue of the Diptera of the Americas south of the United States. Mus. Zool. Univ. Sao Paulo, 100:1-25. -. 1972. 3. Some ob.ser vat ions on the zoogeography of Middle and .South Amer¬ ican ectoparasites. J. Med. Eniomol., 9:589. Wen/ei, R. L., and V. J. Tipton. 1966. Some relationships between mammal hosts and their ectoparasites. Pp. 677-723, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.). Field Mus. Nat, Hist., Chicago, xii+ 861 pp. Wenzel. R. L., V. J. Tipton, and A. Kiewlicz. 1966. The streblid batflies of Panama (Diptera Calypierae: Streblidae). Pp. 405-675, in Ectopara.siies of Panama (R, L, Wenzel and V. J. Tipton, eds,), Field Mus. Nat. Hist., Chicago, xii + 861 pp. Wharion, G. W. 1938. Acarina of Yucatan caves. Carnegie Inst. Washington Publ., 491:137-152. WuiTAKFR, J. O., jR.. AND D, A. Easteri A. 1975. Ectoparasites on bats from Big Bend National Park. Texas. Southwestern Nat., 20:241-254. BIOLOGY OF THE PHYLLOSTOMATIDAE 119 WfiirAKEft, J. O., Jr„ and N. Wilson. 1974. Host and distribution lists of mites (Acari), parasitic and phoretic, in the hair of wild mammals of North America, north of Mexico. Amer. Midland Nat.. 9:1-67, WRF.NN, W. J., and R. B, LCK3MIS. 1967, OforhinophUa, a new' genus of ehiggers (Acar- ina, Trombiculidae) from western North America, Acarologia, 9:152-178. Yunkfr, C. E. 1970, A second species of the unique family Chirorhynchobiidae Fain, 1967 LAcarina: Sarcoptiformes), J. Parasilol., 56:151-153. ORAL BIOLOGY Car IF: TON J. Phi],lips, Gary W. Grimes, ani> G. Lawrence Forman Mammalian dentitions have attracted considerable attention from paleon¬ tologists and taxonomists. This mainly is because jaws and associated teeth are the most commonly found remains of mammals in fossil beds and, thus, the materials most readily available for comparison w ith extant species. The degree to w'hich occlusal patterns or coronal shapes have differentiated, even at the specific or subspecitlc levels, has been the subject of many investi¬ gations. How'ever, relatively little is known about other aspects of oral anatomy and biology of the ora] environment in most mammals. Dental researchers, on the other hand, generally have approached the study of mammalian dentitions from a medical or clinical point of view at the tissue, cellular, and subcellular levels. Until recently, researchers have been inclined to study only selected lab¬ oratory rodents, a few' species of primates, and a variety of domestic forms. The blending together of these two basic orientations results in a much broader, interdisciplinary approach that can be termed '"oral biology,” in view' of the wealth of information available about teeth and associated structures and the wdde variety of sophisticated techniques now at hand, it no longer suffices to undertake only highly specialized, traditional investigations. Thus, in this paper we have attempted to utilize a broader biological approach to a subject that a few- years ago w'ould have been limited to a discussion of coro¬ nal patterns and their taxonomic implications. Until now', no efforts have been made to study comparatively the oral biology of a group of w'ild, free-living species of mammals. It is our contention, however, that such studies are ne¬ cessary if we arc to overcome the artificiality of investigating only selected com¬ ponents of a system as if they exist individually in nature. We agree with Romer’s classic comment that many of our colleagues seem to view- teeth as though they in themselves act as species. Among the Chiropiera, the Phyllostomatidac are perhaps the best suited of all families for comparative analysis of oral biology. Within this one family there are species that have extremely diverse feeding habits, w'hich if not obligate, certainly are restricted. Diets are known to include essentially carnivorous, frugivorous, omnivorous, nectarivorous, and sanguivorous modes. Indeed, the adaptive radiation within these Neotropical bats is extensive even though they have a relatively close genetic affinity and a common ancestry, it is especially important, of course, to underscore the significant relationship between this divergence and the biology and stryctural characteristics of the oral systems. It is unlikely that any other natural assemblage of mammals could provide a more suitable or potentially rewarding source for an evolutionarily oriented study of the biology of oral sy stems. Ideally the present report would follow the traditional format of a review paper. Unfortunately, however, the opportunity to write this particular report 121 122 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY came a few years too soon for us to use such a format exclusively. Much impor¬ tant information has not been published elsewhere so as to be available for re¬ view. One possibility was for us to write a shorter version of this chapter on a restricted aspect w'hile knowing that considerable salient additional data would be published shortly, making the present paper almost immediately obsolete, or at least seriously inadequate. We have chosen instead to wTite a chapter with a somew'hat variable format that ranges from portions which are a typical review to portions composed entirely of our unpublished data. Consequently, some sections and subsections w'ill seem to the reader to be somewhat disproportionate in length and detail. In our view, most of w hat can be said at this time about dental gross anatomy, abnormalities, and taxonomic arrangements based on coronal patterns has been published elsewhere by Slaughter (1970) and Phillips (1971). Although these areas lend themselves to promising additional work, relatively little has been published since these earlier studies. W'e have chosen, therefore, to summarize in a review' style the available information on these matters. Consequently, our emphasis is on the more poorly known aspects of chiropteran oral biology such as evolutionary mechanisms, general dental microanatomy, and compara¬ tive studies of salivary glands. Sections of this paper covering these topics report much previously unpublished information or ideas essential to an understanding of the oral biology of phyllostornatid bats and are, therefore, more detailed. l.astly, a word about the authors is relevant. The senior author initiated the research program and is responsible for most of the interpretations presented herein. The section on transmission electron microscopy of the parotid and sub¬ mandibular salivary glands of Aniheus plumuis, which is the first such published information for phyllostomatids, was written by Carleton J. Phillips and Gary W'. Grimes. The summary of phyllostornatid masticatory apparatus (tongues, neck, and throat musculature) was wTitten by G. Lawrence Forman. Materials and Me i hods Specimens used for portions of this paper that report previously unpublished information about phyllostornatid oral biology were collected in the Mexican states of Nuevo Leon, Hidalgo, Nayarit, and Jalisco, in 1972 and 1973 or in Jamaica, in 1974. In addition to specialized preparative techniques, which are detailed below, many specimens w'ere preserved either as typical museum study specimens (skins and skulls) or as flu id-preserved specimens. Samples from each of the species collections have been deposited as voucher specimens in The Museum, Texas Tech University. Slides, in the case of histological prepara¬ tions, have been deposited in the Department of Biology, Hofstra University. Gene ml Hi st ohfgy Specimens selected for general histological studies with the light microscope (LM) were killed in the field; tissues were removed, placed tn individual con¬ tainers, and fixed in one of the following solutions: 10 per cent nonbuffered or buffered formalin; Camoy's fixative; or aicohol-formalin-acetic acid. Some BIOLOGY 01- THE PHYl.LOSTOMATIDAE 123 specimens were fixed in roio. AIJ materials subsequently were stored in 70 per cent ethyl alcohol until studied. Calcified tissues were prepared for embedding by one of the following two methods: tissues were placed for at least 20 days in 10 per cent ethylenediamine tetra-acetic acid (versene) adjusted to pH 7 or they were placed in Decal (Scientific Products) for tw^o to six hours depending on outcome of tests for calcium (Lillie, 1965). Following decalcification, the speci¬ mens were washed, dehydrated, and cleared in xylene for at least 30 minutes. Vacuum infiltration for 30 to 45 minutes at 25 inches mercury was followed by embedding in paraplasi. Sections were cut at five to seven micrometers. Selected slides were prepared for study with the light microscope by staining with a variety of general histological procedures as follows: Harris’ hematoxylin and eosin-Y (H&E), the periodic acid-Schiff (PAS) reaction, azure-A and cosin- B (pH 4.8-5.0), Masson’s trichrome stains (using Harris’ hematoxylin for two minutes), Mallory triple connective tissues stains (Humason, 1972), aldehyde- fuchsin following oxidation for 30 minutes in peracetic acid (Fullmer and Lillie, 1958), and silver impregnation. Unless otherwise noted, formulae, times and interpretations of tinctorial results of these techniques were based on those of Lillie (1965). Transmission Elec iron Microscopy Transmission electron micrographs are included in this report. Materials for this technique were partially prepared in the field. Glands were removed from bats at lime of death, minced, and fixed in 2 per cent glutaraldehyde in 0.1 M PO 4 buffer. Although not wiiolly desirable, some specimens were stored in this fixative for as long as three weeks prior to embedding. The materials were washed in 0.1 M PO 4 buffer for two hours, post-fixed in one percent OSO 4 in 0.05 M PO 4 buffer for one hour, and embedded in Epon. Additional details of technique can be found in Pease (1964) and Hayat (1972). Sections w-ere made on a Porier-Blum MT-2 ultramicrotome and stained with uranyl acetate and lead citrate. Observation and photographs were made on an RCA EMU 3-G electron microscope (TEM) operated at 50 kilovolts (KV). Scanning Electron Microscopy Critical point drying w'as used for preparation of soft tissues for the scanning electron microscope (SEM). Specimens were embedded (subsequent to decal¬ cification in the case of jaws) in the manner described above (General Histology) and the blocks w'ere sectioned to the desired plane. In this way, it w-as possible to retain prepared slides that could be stained with general histological pro¬ cedures for comparison to the three-dimensional view obtained with the scan¬ ning electron microscope. The blocks then were deparafinized in xylene (usual¬ ly two hours in two or three changes), dehydrated in ethyl alcohol, placed in successive changes of 50, 60, 70, 90 and 100 per cent amyl acetate: ethanol and, finally, dried in a CO^ critical point drying apparatus. The dried specimens were mounted on aluminum stubs and coated lightly with carbon and gold:pal¬ ladium (60:40). With exception of decalcification, embedding, and critical point 124 St^EClAL PUHLJCA't IONS MUSEUM I EX AS JECH UNIVERSITY drying, the Siime procedures were used to prepare hard materials such as teeth. The materials w-ere studied with an Hitachi HHS 2-R scanning electron micro¬ scope at 10 or 20 KV and photographed with either Polaroid PN/55 or Kodak 4127 film. Excellent detailed techniques can be found in Anderson (1951), Boyde and Wood (1969), and Hayes (1973). Tcrminolo^fy Dental nomenclature is highly complex; the terminology employed here for descriptions of coronal patterns essentially is that of Van Valen (1966fd and Herskovitz (1971); an additional, fairly detailed explanation can be found in Phillips (1971). Basic terminology for dental microanatomy is that used in the standard textbook edited by Sicher and Bhaskar (1972). Dental formulae are used only sparingly in this chapter, but w'hen presented they do have phylogenetic implications. By convention, lower-case letters have been used for lower teeth, and upper-case for upper teeth. Thus, the last upper premolar is labeled with an uppercase P and the number 4; the latter suggesting an evolutionary status tor the tooth. We have followed Handley (1959) and Phillips (1971) in regard¬ ing the 13 as the missing incisor in those species having only tw-o upper incisors. A lower-case d denotes deciduous teeth. With regard to the salivary glands, wc have not follovved Wimsatt (1955) and DiSanto (1960), w'ho also have studied these structures in phyllostomatids. These authors used the names parotid, prin¬ ciple sub max illary, and accessory submax illary for the major salivary glands of bats. In so doing, they cited Robin (1881). Herein, w'C have used the names parotid, submandibular, and sublingual for these same glands because the term submaxil¬ lary is not descriptive for a gland generally located near the angle of the mandible, and the name sublingual is used most often in literature for the pair of large salivary glands positioned between the dentaries at the base of the tongue. The nomen¬ clature for throat and cervical musculature is based on that used by Wille (1954). Histologists traditionally have used the terms serous and mucous to describe cells comprising secretory acini of salivary glands. These terms have been val¬ uable for easily communicating the general appearance of cells. Mucous cells, for example, have a clear, almost achromatic cytoplasm when fixed w ith formalin and stained with hematoxylin and eosin-Y, whereas serous cells generally have a relatively dense, chromatic cytoplasm and a large concentration of basophilic material in the basal ergastoplasm (Sicher and Bhaskar, 1972). Numerous studies have indicated, however, that this classification scheme is inadequate re¬ garding nomenclature of secretory products (see Junqueira ei af., 1951; Lebiond, 1950; Wimsatt, 1956; DiSanto, I960). Cells having the appearance of mucous type cytoplasm in the parotid gland of Anibeits jamiJkcnxix, for example^ ap¬ parently do not secrete mucins and, thus, have been termed pseudomucous by Wimsatt (1956). We agree wdth the comments of Shackleford and Wilborn (1968) in not following Wimsatt in using this term. For the purposes of this report we have chosen to use the traditional terms mucous and serous in the descriptive way stated by Sicher and Bhaskar (1972) w-ithout implication of knowledge of the chemistry of the secretory products. Additionally, we have followed others BJOLOGV OF THE PHYLLOSTOMATIDAE \25 (see Shackleford and Witborn, 1968) in referring to those secretory cells having combined morphological or tinctorial characteristics (or both) of classical serous and mucous cells as being seromucoid. Deciduous Dentitions The deciduous (primary) dentitions of phyllostomatid bats are highly special¬ ized and strikingly different from the permanent dentitions. Unlike the deciduous teeth of most mammals, those in bats are not directly functional in comminiitton of food material Instead, chiropteran deciduous teeth generally are regarded as modified for use as instruments for clinging to the female (see Allen, 1939; Reeder, 1953; Friant, 1963). In this connection, it is interesting that in some rhinolophid species the deciduous teeth are resorbed prior to birth (Grasse, 1955; Spillmann, 1927; Dorst, 1953), and in at least one molossid (Mops) some, but not all of the deciduous teeth are resorbed prenatally (Dorst, I957fd‘ Each species of phyllostomatid thus far studied apparently possesses a full comple¬ ment of deciduous teeth that are retained after birth. However, the lower first deciduous premolar (dp2) has not been observed and possibly either is lost short¬ ly after birth or is resorbed prenatally, if it forms at all The deciduous teeth of phyllosiomatids generally are smaller and morpholog¬ ically less complex than those of vespertiljonids and molossids (see Phillips, 1971, for a summary). "I’he simplicity of deciduous teeth in the phyilostomatids is consistent with their comparatively reduced permanent dentitions; only the Phyllostomatinae have permanent teeth with fairly complex coronal patterns (see next section). From an evolutionary and systematic point of view, there are two especially noteworthy features in the known phyllostomatid deciduous dentitions. As de¬ scribed below', morphological differences in the upper deciduous incisors sug¬ gest different systematic relationships among the phyilostomatids than are im¬ plied by the current scheme of classification. It is important to note that such discontinuities have been indicated by a variety of other investigations based on such diverse approaches as comparative serology, chromosomal morphology, and osteology (Gerber and Leone, 1971; Baker, 1967; Walton and Walton, 1970). Secondly, the presence in many phyllostomatid species of three upper deciduous premolars (dP2, dP3, and dP4) is of evolutionary' significance be¬ cause the normal permanent dentitions include only two upper premolars. The only know'n exceptions to this configuration apparently result from abnormal¬ ities such as double initiation and atavism (Phillips, 1971). The presence of a small and unreplaccd upper deciduous premolar directly posterior to the canine provides strong evidence, in the absence of an adequate fossil record, that the two remaining permanent prcmolars can, in fact, be regarded as P3 and P4, in the evolutionary sense. The follow'ing paragraphs summarize current knowledge about the deciduous dentitions in the various phyllostomatid subfamilies. Among the Phyllostomatinae, the know'n deciduous dental formula is 2/3; l/I; 3/2-37. This subfamily is of special interest because of differences in shape I2fj Si^BCJAL PUHLICATIONS MUSEUM TEXAS TECH UNIVERSITY of the first upper deciduous incisors in Macroius in comparison to those of Ton- atkt, MimoHy Chrotoptems, and Phylkfsfo/futs, In Phyllostomus, the second upper deciduous incisor is longer and more greatly curved than the inner one^ which is thin and tapers to a fine, recurved point at the tip (Miller, 1907). In Mimon (Phillips, 1971), Totunia (Dorst, 1957/?), and Chrotoptents (Leehe, 1878), the inner upper incisors resemble those in Fhylhhsftmni.s in being smaller than the outer ones and in having a fine, recurved point. In Macroius, on the other hand, the first upper deciduous incisor is as large as the outer one and is forked, with the mesial lobe being somewhat larger than the lateral one (Nelson, 1966). It also is of interest that in Macroitis the three lower incisors have only two permanent replacements {Phillips, 1971). The first and second lower de¬ ciduous incisors arc trilobed, like their permanent replacements, whereas the third, w'hich is shed but not replaced, is a small, simple spicule. This disparity between numbers of deciduous and permanent teeth is yet another example of the value of the analysis of deciduous teeth toward deciphering evolutionary sequences. The deciduous dental formula in the Glossophaginae is 2/2; 1/1; 3/2-3. Among the 13 genera of glossophaginc bats, the deciduous dentitions of only Glossophaga, Lcpumycteris, and Choeronyeferis have been studied and de¬ scribed (Phillips, 1971; Stains and Baker, 1954). All three apparently have at least 22 such teeth; the first lower deciduous premolar (dp2) has not been found and possibly is either resorbed or shed early in life. The major difference wdthin these genera is in the shape of the upper deciduous incisors. In Ghssophaga and Li’ptonycten's (Fig. 1) these teeth are forked, whereas in Choeronyctais they are pointed and recurved. Within the Carolliinae, only the genus Carollia has been studied; the decidu¬ ous dental formula is 2/2; 1/1; 2/2-3. In Caroliia, the first upper deciduous incisor is the most notew'orthy component because it is thin and the apical end comes to a fine, recur\'ed point (Lcche, 1878; Miller, 1907). This deciduous tooth thus resembles the inner upper deciduous incisor in some species of Phyl- lostomatinae as well as at least Choenmyaeris in the Glossophaginae. The deciduous dental formula in the Slenoderminae is 2/2; 1/1; 2/2. In the two genera of this subfamily (Arfiheus and Anietrida) for which data are avail¬ able, the first upper deciduous incisor is forked, as in Mactofus (Lechc, 1878; Miller, 1907). The deciduous teeth in species of Phyllonycterinae have not yet been investi¬ gated. In the Desmodontinae, both Dewufdus and Diphylkt have a deciduous dental formula of 2/2; 1/1; 2/2. d’he teeth are small and greatly simplified; apparently the two upper deciduous incisors, both of which are the same size and are simple hooklike spicules, are functional (Miller, 1896; Birney and Timm, 1975). The remaining deciduous teeth are extremely small, barely penetrating the gingivum, and apparently are shed rather soon after birth (Birney and Timm, 1975). BIOLOGY OF THE PHYLLOSTOMATIDAE 127 Fifi. I.—An example of deciduous teeth in the Mexican long-nosed bat, Lepiouycieris ttivitih. The tooth marked with a ? has not actually been seen in available specimens. From Phillips, I97L Permanent Dentitions Development Dental development is a complex process that is almost usistudied in phyl- lostomatid bats. This is unfortunate because full understanding of comparative dental ontogeny probably would be of considerable value to interpretation of evolutionary mechanisms and relationships. This is especially true regarding interpretation of such abnormalities as double initiation, incomplete dichotomy, and atavism. Furthermore, the direct relationship between morphogenetic in¬ tegration and the process of dental ontogeny is readily apparent (Phillips, 1971). Bats remain almost unknown in this regard, although development of molars in relationship to integration of coronal configurations has been investigated in a horseshoe bat, Hipposideros bra/n.if (Marshall and Butler, 1966). Studies on other mammals, particularly insectivores and marsupials, further underscore the im¬ portance of an understanding of dental ontogenesis to the determination of dental evolutionary mechanisms (Ziegler, 1972a, 1972/); Kindhal, 1963; Berkovitz, 1967, 1972; Osborn, 1970, 1973). Three readily recognizable formative stages of dental development generally can be used to delineate aspects of mammalian tooth development (Sicher and Bhaskar, 1972). It must be remembered, however, that ontogenesis is a con¬ tinuous process rather than stepwise, as might be implied by common use of the term stage. Dental lamina stage .—Initiation of the teeth results from cellular pro¬ liferation within the epithelial dental lamina, which is of ectodermal origin. Dental buds, which are the priniordia of individual teeth, develop simultaneously with differentiation of the dental lamina. 128 SPECIAL, PUBLICATIONS MUSEUM lEXAS TECH UNLVBRSITY Demal cap This developmenial phase is characterized by uneven cellular proliferation resulting in formation of an outer and inner enamel epithe¬ lium. It is especially important to note that development of the dental cap in- nuences the mesodermal mesenchyme, which in turn condenses to form the dental papilla. The dental papilla provides the primordium for both the dentin and pulp; it is this mesodermal component, through the process of dentinogenesis, that actually sets the size and occlusal pattern of the finished tooth (Tonge, 1971; Osborn, 1973). Denitd bcil stage .—This stage is characterized by both histodifferentiation and morphodifferentiation that results in formation and alignment of amelo- blasts and odontoblasts, which in turn will form the matrices of enamel and dentin, respectively. A remarkable SEM micrograph of a developing permanent upper premolar in a near-term fetus of a specimen of the Jamaican fig-eating bat, Ariteus flave- scens, is shown in Fig. 2. In this instance, enamel and dentin formation has reached the cement-enamel junction, and root formation is well underw'ay. A distinct, smooth-surfaced epithelial diaphragm can be seen at the root apex. A dense band representing the columnar cnamel-producing ameloblasts and stratum intermedium also can be distinguished easily in this electron micrograph. The developing tooth is cushioned by the stellate reticulum, also w'ell illustrated by this figure. The long processes that connect the component cells, together w'ith the fact that the stellate reticulum is fluid rich, gives this layer a “lacy” appearance. Within the Phyllostomatidae, dental development in relationship to age has been investigated histologically only in the Jamaican fruit bat, Aritbeus jamaken- sis. The following summar>' comments are based on this study {Farney, 1975). Because ages of individual bats cannot be determined w'ilh precision, Farney (1975) used the standard measurenieni of crown-rump length in his report. At some future date, it might be possible to relate these measurements to actual age; at best, they currently provide chronological indications. In the 9-milli- meler embryo, a dental lamina was evident; tooth buds could be distinguished anteriorly. In an embryo that measured 13.5 millimeters Farney (1975) w'as able to identify typical bell stage primordia, again representing only anterior teeth. At this time in grow th of the embryo, the low'er first molar was in the cap stage, suggesting early initiation of the molar field. It probably is true that in phylloslomalids in general the upper and lower first molars develop and erupt early. This possibility is strongly supported by earlier studies (Phillips, 1971) of tooth eruption sequences in three glossophagine genera {Glossophaga, Lx’pto- nycteris, and Choerotiyctem). The importance of this finding is reflected in tenns of morphogenetic fields and, consequently, dental evolution. The upper and lower first molars in phyllostomatids can be regarded as molar determinants as discussed by Osborn (1973) and, thus, arc pivotal in additional studies of developmental interrelationships of permanent teeth in these bats. An embryo of Artiheus jatnaicensis that measured 20.5 millimeters was found BIOLOGY OF THE PHYLLOSTOMATIDAE 129 Fig. 2* —Scantling electron micrograph of a developing upper premolar in Arifens flavescenx. Abbreviations are: D, dentin; E, enamel; EE, enamel epithelium; ED, epithelial diaphram; O. odontoblasts; P, pulp; PZ, proliferative zone; SR. stellate reticulum. 109 X. lo have deciduous teeth in a late bell stage; buds representing permanent teeth were detected anterior to the first molar (Farney, 1975). In a 3L5 millimeter embryo, the deciduous incisors and canines were fully formed; the permanent incisors were in the bell stage and the permanent canines were in the cap stage. The first permanent premolars w'ere reported to be in an early bud phase, whereas the second premolars were somewhat more developed; this latter finding strongly suggests that the last premolar (P4) can be regarded as a premolar determinanL SPECIAI. PUBLICAI IONS MUSEUM TEXAS TECH UNIVERSITY L^O Eruption and Shedding Among the phyllostomatids, eruption and shedding has been studied only in three genera of glossophagines {Glossophaga, Leptonycteris, and Choenmycteris) and, therefore, is a topic still largely unknown (Phillips, 1971). In general, eruption is a continuous process that can be divided into three stages (Sicher and Bhaskar, 1972): a pre-emptive phase during which the dental organ completes development and enamel and dentin arc Ibrmed; a prefundional eruptive phase during which the root(s) forms and the new tooth moves to the occlusal plane; and the functional eruptive phase, which begins after the tooth reaches the occlusal plane. The first two of these phases arc of primary con¬ cern to the present discussion; the third phase, which involves the complex move¬ ments of mature, functional teeth is of interest in investigations of changes that take place in the dental arcade as responses to stresses and attrition. For example, in the eommon vampire bat {Desmodns rotimdns)^ the mature permanent teeth continue to move into the occlusal plane as considerable attrition due to thegosis reduces the crowns of the enamcl-less teeth (Phillips and Steinberg, 1976). Regarding developmental problems of the first two eruptive phases, there pre¬ sently are three main topics of considerable interest. How' and w'hy do developing teeth undergo the initial process of eruption, w hat is the relationship of erupting permanent teeth to the shedding of the deciduous teeth, w'hat is the process of passage of the permanent teeth through the oral epithelium and the mechanism of epithelial attachment to the surface of the tooth? Many mechanisms of eruption have been suggested and studied experimentally but the problem is far from resolved (see Sicher and Bhaskar, 1972; Phillips and Oxberry, 1972). Studies of the glossophaginc long-tongued bat, Choeronye- leris mexieatuu provide the only information on this subject for phyllostomatids (Phillips, 1971). Analysis of these materials reveals no trace of a hammock ligament, as reported by Sicher (1942) for certain rodents. It is of further interest that there is no indication of a vascular role in eruption because the pulp and connective tissue adjacent to the developing apical foramen are not highly vas¬ cularized, at least in the studied specimens of Choeronyaeris. The relationship of the newly erupting permanent teeth to the deciduous teeth in Choeronycieris is more straight forward. In conjunction with enamel maturation, the stratum intermedium and amclobiasts, and perhaps the reduced outer enamel epithelium, become indistinguishable due to changes in cellular morphology (Figs. 3, 4). Both cell types appear to develop large, clear (formalin fixed and stained with hematoxylin and eosin-Y) vesicles within the cytoplasm. In most of the cells, the vesicle is so large that the heterochromatic nucleus is basally restricted and crescent shaped (Figs. 3, 4). Additionally, in these speci¬ mens there is a distinct, thick proliferative zone of cells at the coronal apex of the developing permanent tooth (Figs. 3, 5). The mesodemial connective tissue that surrounds the developing tooth following the loss of the stellate reti¬ culum and reduction of the outer enamel epithelium shows considerable altera¬ tion in those areas adjacent to the proliferative zone (see Phillips, 1971). It has been suggested elsewhere (Sicher and Bhaskar, 1972) that cells of the prolifera- (UOLOGY OF THE PHYLLOSTOMATIDAE LH live zone produce an enzyme (possibly hyaluronidase) that leads to a loss of the ground substance within the collagenous fibers that comprise the principle fiber bundles. The previously dense connective tissue thus becomes a loose, fluid-rich tissue with fine argyrophilic fibers. This presumed process of proteo¬ lysis allows for passage of the new' tooth tow-ard the oral cavity. Although lymph¬ ocytes have been reported from regions of connective tissue degradation in other species (Fullmer, 1967), no intlammatory cells have been observed in such areas in young of Choeronycteris (Phillips, 1971) and are not generally regarded to be a factor in this developmental process. In sections from a new-born Choero- nyaeris stained w-ith the periodic acid-Schiff reaction (PAS), the region of con¬ nective tissue undergoing degradation is PAS negative or, at most, only UKxler- ately PAS positive. This is in contrast to the unaffected adjacent tissue, which generally is strongly to moderately PAS positive. Following Spicer cf al. (1965), it can be said that connective tissue undergoing degradation is low in mucosub- stancc. Based on studies of Choeronycteris (Phillips, 1971), it also can be suggested that the process of initial eruption of permanent teeth in phyllostoniatids is fairly rapid. Unlike the process in man, there are no histological indications of periods during which areas of resorption are partially repaired. The eruptive process of permanent teeth directly affects shedding of deciduous teeth (Sicher and Bhaskar, 1972). Although the mechanisms have not yet been established, it is clear that in young phyllostomatids the proximity of a perman¬ ent tooth results in resorption of the root of the deciduous tooth (Phillips, 1971). As is visible in Fig. 5, the medial surface of the root of a deciduous upper pre- molar w-as undergoing resorption; it is of additional interest that multinucleated cells that morphologically and tinctorially resemble osteoclasts can be seen w-ithin the resorbed area. It has been suggested, but not confirmed, that pressure exerted by the permanent teeth causes osteoclasts to differentiate from the surrounding mesodermal connective tissue (Sicher and Bhaskar, 1972). Coronal Con ftgurat ion s The coronal configurations of secondary teeth of many phyllostomatid species have been described, figured, and discussed tn a wide variety of publications (Hall and Kelson, 1959; Slaughter, 1970; Miller, 1971; Phillips, 1971; Winkel- mann, 1971; Farney, 1975). Consequently, the following generalized comments are not descriptively detailed but instead are intended as a review' and back¬ ground. Traditionally, know'ledge about dental morphology of bats has had great practical value because of the use of dental characteristics in taxonomy. This aspect is underscored by an examination of books such as Hall and Kelson (1959), in which keys to families, genera, and species frequently are based mostly on dental characters. How-ever, as we learn more about various inherited char¬ acters of phyllostomatids, it is likely that features of secondary dentitions taken alone will fall short of providing adequate and accurate presentations of real genetic relationships. 1.^2 SPFCtAL PUm.lCATIONS NfUSEUM TEXAS TECH UNIVERSITY Fig. T—Reduced enamel eplihclium of developing permanent premolar in Chocro/iycfcrix nu'xitatui. Abbreviations are; D. dentin; E, enamel space; PZA, proliferative zone of amelo- blasts; V, vesicles. Hematoxylin and eosin-Y. 662 X . Phyllask)t}Ui{i}nu '.-—Dental formula; I 2/1-2; Cl/I; P 2/2-3; M 3/3. The teeth of all species in this subfamily are robust and relatively primitive. The inner upper and lower incisors typically are larger and more developed than arc the outer ones (when the latter are present). The canine teeth tend to be thick based and have notable cingula. The height of the canines is not appreciably greater than is the height of the premolars and molars. The upper molars are nearly square; the ectoloph is primitively W-shaped and is considerably higher than the remainder of the tooth. In occlusal aspect, the ectoloph occupies ap¬ proximately one-half of the tooth. The low^er molars also are primitive; the irigonid has the typical triangular shape as does the talonid, giving these molars a W-shaped appearance. The cusps and commissures of both upper and lower molars normally are relatively sharp, regardless of the individual’s age. Sharp- BIOLOGY OF THE BHVIXDSTOMATIDAE 133 Fit;. 4.—Details of reduced enamel epithelium in Ciioi'ronyclerh mexicanu. Top: near coronal apex. Botio/n: junction of crown and root surface. Abbreviations are: A, amelo- blasls; D. dentin; E. enamel spaces; EC. enamel cuticle: N, euchromaiic nucleus of amelo^ blast; N2, heterochromatic nucleus of vesiculated cell; S, stellate reticulum; V. cytoplasmic vesicle. Hematoxylin and eosin-Y. 1299 X. ness is maintained by ihegosis, which in turn is a consequence of the occlusal pattern, Glossophagitiae. —Dental formula: 1 2/0-2; Cl/I; P 2-3/2-3; M 2-3/2-3. The genera comprising this subfamily can be divided into at least three distinct groups based on dental characteristics; if all aspects of dental morphology are considered equally, the pattern is even more complex (see Phillips, 1971). In all studied species, the teeth are relatively small and in some they actually are minute in comparison to those of the Phyllostomatinae. Among the glossophagiiies (Fig. 6), the upper inner incisors are either large to moderate in size (for instance, Glossophaga, LepionycteriSy and Plaialina) or much reduced and separated by a distinct median gap (for instance, Anoura, Hylonyaeris^ and Cfweronycteris). The canines in all of the species are high and slender. Three or four groupings of 134 SPECIAL PUBIJCAIIONS MUSEUM TEXAS TECH UNJVERSHY Fic. Developing permaneni premolar and resorbing adjacent deciduous tooth (arrow) in Clitierofiyt/i’ris fnexiaiiin. Abbreviations are: A, ameloblasts; D, dentin; DC., altered con¬ nective tissue; E, enamel space; OC\ “osteoclast;" P, pulp; S, stratum intermedium. Hema¬ toxylin and eosin-Y, 304 X . molars can be distinguished within the subfamily, as currently defined. Within the first group of genera, Anoura, Lionycteris, and Lcftwhophylla have the most primitive configuration. A relatively high, W-shaped ectoloph is present, and the metastyle is prominent. In Glossophaga, Monophyilus, and Leptonyaeris^ the ectoloph also is W-shaped (although considerably elongated in the last genus) but is low er, and the metastyle is much reduced in comparison to the first group of genera. In yet another grouping (Lichonycieris^ Scleronycreris, and Hylonycteris), the ectoloph is low and has been modified considerably, especially in the anterior elements; the paracone apparently has been lost, leaving a distinct parastyle on the anterior clement. Determination of whether or not the paracone or parastyle BiOLOGV OF THE PHY[.LOSTOMAT[DAE A B Ft£i. 6.^—An example of the extremes in arrangement of permanent upper and lower in¬ cisors in glossophagines. A: Lom hftphyUa rohuxtu. B: A/iotira fieoffroyi. From Phillips, 1971. was lost (in absence of a fossil record) was based primarily on a remarkable series of Lichimycferis that has been described elsewhere (Phillips, 1971). It should be noted, however, that Winkelmann (1971) apparently disagreed with this interpretation, although he has not offered other evidence or an explanation for his opinion. The posterior element of the ectoloph in these genera still is nearly triangular. The genera Pkiialina, Musonycieris^ Choenmycieris, and Choeroniscus all have highly modified upper molars, in which the labial edge consists of a raised lip with an anterior parastyle and posteriorly positioned metacone. In contrast to the extremely complex evolutionary pattern found in the upper molars of glossophagines, the lower molars are remarkably uniform. Variation in configuration mostly is in size and height of the cusps and is relatable to the degree of modification from the primitive pattern found in the upper molars. Carollii/uie. —Dental formula: I 2/2; Cl/I; P 2/2; M 3/3. The dentitions in species of Ccirollia and Rhinophylla have been influenced by shortening of the upper and lower jaw'S. The upper inner incisor is large and somew'hat pro¬ cumbent, whereas the second incisor is much reduced and almost peglike. The lower incisors also are small. The upper and low'er premolars are robust and high-crowmed. The upper molars are considerably modified from w'hat must have been the primitive configuration. The W-shaped ectoloph usually is re¬ duced, or indistinguishable. The stylar shelf is high; the paracone and metacone often are almost linearly arranged along the labial border (depending on species). The protocone is absent, and, in fact, the entire lingual portion of the molars is anteroposteriorly narrowed. The lower molars also are highly modified; in Carollki, the lingual cusps (metaconid and entoconid) are much reduced, and in Rhinophylla they have been lost altogether. we.—Dental formula: I 2/2; C 1/1; P 2/2; M 2-3/2-3- The dentitions of the large and variable number of species in this subfamily have been greatly influenced by both widening and shortening of the upper and lower 1^6 SPECIAL PUBL[CATlONS MUSEUM TEXAS [ ECH UN1VERS[TY jaws. The basic phyllostoniatid pattern of large inner and small outer upper incisors and relatively small lower incisors has been maintained. The canines usually are extremely robust and broad based as also are the upper and lower premolars. The upper molars in these species typically have a low profile and lack the primitive W-shaped ectoloph. Instead, the teeth either are narrow and long or nearly cuboidal (as in Sriinum). The paracone and metacone have been shifted to the labial margin, forming a longitudinal Hp. The lower molars also are broad and clearly designed for crushing fruit; the trigonid, however, generally has been maintained and is easily distinguished. Thegosis is insignificant, probably because of the occlusal pattern, and, thus, the molar surfaces of steno- dermines have a rounded appearance rather than the distinct, sharpened cutting edges characteristic of the Phyllostomatinae. PhyilonycH’/inae .—Dental formula: 1 2/2; C 1/1; P 2/2; M 3/3. The three genera {Brachyphyiki, E/t>pfiylUi, and Phyil(?fjycferis) tentatively classified in this subfamily (Jones and Carter, 1976) present two basically different dental patterns. In Brachyphyiiay the upper and lower jaws have been shortened and widened, as is typical of the stenodermines. The dental arcade has been modified considerably but in overview' also is similar to those found among species of this subfamily. The major difference, visible in the upper molars, is that the protocone and a relatively large metaconule are found in the basin between the lingual and labial borders. The other two phyllonycterines, Emphylla and Fhyllonyaeris, have narrow, elongate upper and lower jaw'S, and the teeth are greatly reduced in size in ac¬ cordance with an evolutionary trend possibly toward nectar feeding. The upper molars in these bats are relatively broad and exhibit, in reduced form, the pat¬ tern found in the stenodermines; the teeth have a distinct labial edge and broad concave basin. Desmodontimte .-—Dental formula: 1 1-2/2; C 1/1; P 1/2; M I-2/1-2. The three genera of vampire bats obviously have the most strikingly modified denti¬ tions among the phyllostomatids. The cheek teeth are much reduced in number and size because selective emphasis has been placed on the upper inner incisors (which are procumbent) and upper and low'er canines. These teeth are large and have extremely sharp cutting edges maintained by thegosis (see Phillips and Stein¬ berg, 1976). Evolutionary Mechanisms A survey of the highly variable dental configurations found in extant species of phyllostomatids produces a striking example of adaptive radiation of coronal dental anatomy in at least partial relationship to modifications in diet. Evolution¬ ary patterns are relatively easy to decipher, even in the absence of a useful fossil record. Likewise, development of possible evolutionary' scenarios and logical arguments for various genetic relationships among the species, based on dentitions, are reasonably straight forw-ard, even if incomplete. The most sig¬ nificant problem, as yet essentially unanswered, is; how' did the teeth evolve, or, what are the mechanisms of dental evolution? The following comments are BIOLOGY OF THE PHYLLOSTOMATIDAE 137 intended ta supply some ideas on, and review some of the conceptual aspects of, this subject, 1 wo questions of special interest regarding evolutionary mechanisms are: how are teeth lost (keeping tn mind that many phyllostomatids have lost one or more permanent teeth or have variable numbers of teeth); and how' are coronal configurations modified? Loss of certain permanent teeth is a characteristic shared by all phyllosto- 111 at ids. Several mechanisms potentially are involved and although some mammal¬ ian taxonomists seem to believe that teeth simply become smaller and eventual¬ ly disappear, the actual factors arc more interesting and certainly more complex. As pointed out in the section on deciduous teeth, the pattern of loss of permanent teeth can in some instances be deciphered by the presence of unreplaced decidu¬ ous teeth. Relatively few workers have emphasized the fact that species of several chiropteran genera have unreplaced deciduous teeth. Such inconsistencies be¬ tween number of deciduous teeth and number of permanent teeth are not re¬ stricted to the Chiroptera (Berkovitz:, 1968). Among the glossophagines, which have been studied most intensively, Choeronyaeris mexicana usually retains some of its four lower deciduous incisors in adulthmid, and, furthermore, histo¬ logical studies have revealed that permanent incisors form but do not erupt (Phillips, 1971). Thus, in this species at least, the permanent lower incisors have not really been lost. The mechanism involved is unknown but possibly a mutation has caused a localized biochemical alteration that results in destruction of par¬ tially developed permanent teeth at a time when they are in an advanced stage of morphogenesis. Grewal (1962) reported a similar instance in certain strains of laboratory mice (Miis mu.sc id us). Extension of the mcchanism(s) that cause permanent teeth to fail to develop fully and erupt, even though their morphogenesis is normally initiated, raises the question of whether other missing permanent teeth in phyllostomatids have been lost completely in the sense that morphogenesis is not even initiated. The high incidence of an atavistic P2 in some glossophagines (Phillips, 1971) in¬ dicates that the potential for development and eruption of this tooth still is present, Krutzsch (1953), Johnson (1952), and Sheppe (1964) debated the evolutionary importance of supernumerary and atavistic teeth in rodents, but the situation does not appear totally analagous to that found in phyllostomatids. In phyllostomatids, supernumerary teeth (Fig. 7) resulting from double initiation or from some abnormality during differentiation can be recognized because they are morpho¬ logical duplicates of another tooth or because they actually are a part of an¬ other tooth. The theory that the small, single-rooted, permanent tooth formed between the P3 and canine in many glossophagine species most likely repre¬ sents the permanent P2, and thus is atavistic, also is supported by the fact that in at least three genera (Glos.suphagu, Lepionycieris, and Choemnycieris) there is a deciduous P2. Sheppe (1964:35) stated that "in the evolutionary history of a tooth there is a time when it occurs in almost all individuals. If for some reason the genes necessary for its development begin to be lost by the population. . , . eventually SPECIAI. PUHI.ICA I IONS MUSEUM I EXAS TECH UNIVERSIT Y DECIDUOUS CONFIGURATION DOUBLE INITIATION ATAVISM incomplete DICHOTOMY p4 Fi(i. 7.—A comparison of normal deciduous configurations and abnormal or unusual permanent configurations of potential evolutionary importance. From Phillips, 1971. all geneiic basis for the tooth will be lost. . . [and] if a tooth later appears in the same place it will be because of either a new mutation or some developmental accident without genetic basis.” In fact, however, the factors controlling initia¬ tion, development, and eruption of teeth are considerably more complicated than this statement w-ould suggest. Loss of various permanent teeth in the evolu* lion of phyllostomatids clearly has been a complex process and not simply a matter of losing necessary genes. Among other mammals, Kurten (1963) discus¬ sed the loss and return of ni2 in the evolution of certain felids; he pointed out that the return of a tooth could have been the product of activation of the field of molarizalion, which presupposed that the genotype for m2 had never been lost although the tooth had been lacking. This topic is brought into sharper focus W'hen viewed in light of data from current developmental studies. The role of mesenchymal papillae in detennina- BIOLOGY OF THE PHYLLOSTOMATIDAE 139 tion of tooth size, shape, and presence or absence (Kollar, 1972, 1975) is so considerable that in our view models of the evolutionary process can use this single component as a cornerstone. For example, Glasstone (1965) has shown that mechanical division of a papilla into halves will result in development of two half-sized but morphologically normal, teeth. Reduction of a mesenchymal papilla into fragments of less than one-half its normal size will cause an abortion of the developmental process (Glasstone, 1965; Kollar, 1975), From experi¬ mental data such as these, it can be theorized that intrinsic or extrinsic factors limiting mesenchymal cell division (during proliferation) to a level below' a critical mass w ill result in the phenotypic absence of a given tooth. Such intrinsic or extrinsic factors presumably are reversible; the model thus derived is theoret¬ ically sound and easily applicable to descriptive data such as those presented by Kurten (1963), Additionally, a model based on Glasstone's (1965) studies also enables us to offer a solution to the puzzle of how' teeth disappear following an apparent trend tow'ard reduction. Teeth can become phenotypically absent when proliferation of mesenchyme is limited to production of a mass of cells below threshold level. Most importantly, when developmental data are integrated into studies of evolutionary mechanisms, it becomes clear that loss of genes is not necessarily a factor. Teeth can be phenotypically absent even though develop¬ ment of their ectodermal (epithelial) component is initiated in the dental lamina. When the above model is considered with regard to atavistic teeth in glos- sophagines (in conjunction with the presence of a dP2), the case for atavism is greatly strengthened. It also is applicable to instances where teeth apparently have been crow'ded out by narrowing of the jaw bones. Any intrinsic factor (such as DNA mutation within specific mesenchymal cells) or extrinsic factor (such as mechanical limitation, innervation, circulatory pattern, or influencing product released by adjacent cells) that causes limitation of mesenchymal pro¬ liferation can have a major influence on size and even presence or absence of teeth in the phyllostomatids. Our model of mesenchymal proliferation also can be applied to problems in geographic variation. For example, Jones and Phillips (1976) recently presented a taxonomic review of the genus StHrnira in the Antilles. In this study, coronal differences between samples of Smrnira lilium were consistent enough to sug¬ gest limited interbreeding between insular populations. Consequently, subspecific designations were w-arranted. Differences in size, occlusal patterns, and even presence or absence of the last molars clearly are nonrandom in this phyllosto- matid species. Indeed, consideration of data in terms of evolutionary mechanisms allows us to elaborate on the taxonomicalJy important patterns reported by Jones and Phillips (1976). Thus, we now can delineate the levels to which such varia¬ tion can be traced. Variation in upper and lower molars in Antillean Sturnira lilium probably has the following sources: 1) Variation in occlusal pattern (size and shape of cusps) in ml and Ml results from variation in arrangement of the mesenchymal papillae of these teeth. Size, and to some extent shape, can be modified by a simple, slight increase or decrease in number of cells; that is, the exact cut-off point of mitotic division during proliferation of the mesenchymal papilla. What- 140 SPEC]At. PUBLfCATlONS MUSEUM TEXAS TECH UNIVERSITY ever the factors determining cessation of division, their effect varies between populations. 2) The upper and lower third molars are smaller than the other molars. Geographically, specimens from Dominica and Martinique have smab ler third molars than do specimens from more southerly islands, such as St. Lucia (Jones and Phillips, 1976). Additionally, some specimens from Dominica and Martinique bilaterally or unilaterally lack m3. Again, these patterns of variation can be traced initially to the mesodermal component. Loss of cusps might partially be a function of reduction in size (number of cells) of the mesenchy¬ mal papilla due to early cessation of mitotic division. In vitro studies w-ith mechanically reduced mesenchyme would suggest, however, that more is involved than a severe reduction in number of cells (Glasstone, 1965). Consequently, the answer might be that an additional component such as a lessening in organiza¬ tional control is the cause in loss of cusps, Osborn and Crompton (1973) refer¬ red to this process as an aging of the dental lamina. Actual absence of one or both m3 quite possibly rctlects an inadequate mass of cells in the mesenchymal papilla of the tooth germ. Again this wotdd be attributed by us to intrinsic or extrinsic factors causing early ce.ssation of mitotic division within the proliferat¬ ing mesenchymal papillae. Overall, the basic model explained here is especially important relative to the phylloslomatids. It seems clear, furthermore, that loss of a particular tooth in¬ volves much more than loss of genes. Pilbeam and Gould (1974) recently pointed out certain pitfalls to interpreta¬ tion of apparent evolutionary changes in tooth size. Their mathematical analysis was applied to human evolution but clearly has important implications for the present discussion. These authors presented data that, although somewhat in¬ conclusive (by their own judgement) in terms of statistical significance, never¬ theless, clearly supported their contention that positive allometry' between tooth size and overall body size can be an important factor and certainly w-orlliy of consideration in making an evolutionary analysis. If these authors are correct, one would expect an increase in tooth size and possibly modification in shape in conjunction with selection for increase in body size. In absence of useful data on body weight, Pilbeam and Gould (1974) used length of skull as one criterion of size in determination of geometric scaling of teeth. We regard this as especially noteworthy in view of the common role of mesoderm in both bone formation and determination of tooth size and shape. This developmental com¬ monality reflects the potential for a common controlling, or intluencing, factor in terms of size, as outlined in our model discussed above. Although w'C have not yet run regression analyses on any of the phyllosto- matids, it is reasonably apparent that the role of positive allometry must be considered in our effort to understand dental evolution in these species. The stenodermines perhaps will provide the best example of positive allometry. In this group of species, there is a fairly wdde range in size, and it is quite probable that many of the dental differences are more nearly a consequence of selection for size than they are the product of direct scleclional pressure on the teeth. Additionally, the loss of third molars in the largest species and in small species BIOLOGY OF THE YLLOSTOMATIDAE 141 having the broadest rostra might paradoxically be coupled with geometric scaling. In most (probably all) phyllostoniatids the first molars are the determinant teeth in the permanent dental arcade. That is, these teeth develop first and directly or indirectly influence the other cheekteeth and maybe even the anterior teeth, although the latter seems doubtful (this interdental relationship is discussed more fully in following paragraphs on morphogenetic fields). The role of deter¬ minants w'as not explored by Pilbeam and Gould (1974), but clearly their analysis of positive allometry is directly applicable. The paradox is that although positive allometry can result in larger and more robust teeth in conjunction w-ith increased body size, it probably also can cause reduction and even loss of third molars. It apparently is typical for Ml and ml to form first (Osborn, 1970) and in- tluence the others; the reduction of coronal pattern in the posterior-most molars has been related by us to alterations in the mesenchymal papilla and referred to as an aging of the dental lamina by Osborn and Crompton (1973). Extension of these various data clearly suggest that apparent loss of third molars can be presumed to be a secondary impact of selection for increased body size or in¬ creased rostral length or width simply because their development is retarded, retrogressed, or even eliminated through resulting lack of space or even mechan¬ ical or biochemical influences from the preceding molars. This model seems especially applicable in studies of the genus Anibeus, in w'hich species of various sizes currently are classified. Additionally, it probably is noteworthy that the third molars have been lost in species of the glossophagine genus Leptonyderis (Fig. 8). An evolutionary trend tow'ard increase in body size (in comparison to other species in the subfamily) is readily apparent in these species. The most intriguing outcome of this logic is that we are presented w ith a sound model for explaining loss of posterior molars and once again reminded that teeth can be altered in size and shape as well as in presence or absence without any direct selective pressure on their own genetic complement. Reduction in size of incisors for certain of the glossophagine species probably reflects what Pilbeam and Gould (1974) regard as “absolutely small” in contrast to geometric scaling. We have not tested our hypothesis mathematically for purposes of the present discussion but, nevertheless, would suggest that the evolu¬ tionary trends in these teeth for those species in which they are small or missing reflect direct selective pressure. In terms of evolutionary mechanisms, this scenario contrasts sharply w ith that presented for the trend tow'ard loss of third molars. The subject of Morphogenetic integration of developmental fields is another major component of a discussion on evolutionary mechanisms involved w'ith dental coronal patterns. The phyllostomalid bats have been little studied in this regard, except for an earlier analysis of several glossophagine species by the senior author (Phillips, 1971), Many w'orkers have pointed out that interpretation of dental evolution requires analysts of coronal patterns in terms of complex inter¬ relationships between developing permanent teeth rather than in a simplistic fashion whereby genes are tied to particular characters by direct, complete causa¬ tion (see Gould and Garwood, 1969; Olson and Miller, 1958; Kurten, 1963; Van Valen, 1965, 1970). At the same time, however, the mechanisms involved In 142 S1>ECIA1- PUBLICA'MONS MU-SEUM TEXAS TECH UNIVERSITY GLOSSOPHAGA {Upper) dP 4 dP 3 dc dP2 dh \ t \ f / Ml \ [ 1^2 P3 p 4 c h 7 dl2 f 12 ] * (Lower) / / p2 dp3y ^dp4 di2 fMl m2 \ 2 ] /p3 P47 ^ /C M37 dil LEPTONYCTERIS (Upper) dP 4 dP 3 dll dP2 dC dl2 ( f M1 W P4 P 3 C ti I27 M2 dp2? di2? dp3dp4dc dil (Lower) / / \ f / \ * P2 /^Ml i27 ^'^'^/‘P3P47 /"m 2 c il7 CHOERONYCTERIS (Upper) M1 dP4 dP3 dP2 / / I ^ [M2 P3 P 47 [CM3J d C d 1 1 d 1 2 / f / ' /ii 12 ;/ dp2? r (Lower) / Miy dp4 dp3 / t M2 /'P3 p4 m37 dc 1 c Fio. B.—Comparative eruption sequences for three genera of glossophagines. Note the early eruption of the Ml, which apparently is the molar determinate, and the relatively late eruption of M2 in Li-piofiycterix. From Phillips, 1971. developmental interrelationships remain far from being established. As Van Valen (1970) indicated, current models requiring gradients might not be widely applicable but, nevertheless, are useful in helping to explain some developmental patterns that in themselves certainly do exist. Three aspects of morphogenetic integration have been demonstrated in gtos- sophagine bats (Phillips, 1971): at least some significant correlations arc found between coronal features of different molars; interpretation of the correlation patterns is facilitated by inferential knowledge of developmental sequences (Fig. 8), suggesting that certain teeth could act as determinants; and in evolution of longer Jaws and decreased complexity of occlusion, one also finds a decrease in significant correlations between dimensions of upper and lower molars. HIOLOGY GF THE PHYU OSTOMATIDAE 143 The factors controlling relationships suggested by significant correlations within and betw een mature teeth are uncertain. It must be remembered, of course, that significant positive or negative correlation coefficients suggest, but do not prove, a causal relationship. Various workers have cited “intraembryonic com¬ pensation" (Van Valcn, 1962) or “reciprocal variation in odontogenesis" (Cam a aL, 1966J as possible mechanical factors and enzymatic, hormonal, or neural controls as organizers and indirect factors (Schour, 1934; White, 1959; Van Valen, 1966/?). Although it once was widely assumed that directional selection would result in a decrease in genetic and phenotypic variation, Guthrie (1965) and Bader and Lehmann (1965) obtained data that strongly suggested an increase in varia¬ tion in dental characters w'ithin species undergoing rapid evolution. Application of this finding to analysis of dimensional variation in coronal components in several species of glossophagines led the senior author (Phillips, 1971) to hy¬ pothesize that the paracone rather than the parastyle has been lost from the upper molars in some of these species. The arguments for this opinion are based mainly on two findings: a definite pattern of high coefficients of variation in the para- cone-parastyle length in those species having both elements; and individual variation in specimens of Licfiouyctcris obscuni in w'hich all conditions, includ¬ ing intermediates, w'ere found and studied (Fig. 9). In this regard, it should be remembered that Winkelmann (1971) has disagreed, or at least has not followed this hypothesis. He has not, how'ever, offered a counter argument. Future comparative statistical and morphological studies of coronal features undoubtedly w-ill be valuable in efforts to determine evolutionary pathw'ays and mechanisms. Presently, however, limitations in knowledge of developmental biology and tissues pose the greatest problems to delineation of evolutionary' mechanisms. One apparent solution is to combine histological and histochemical information w'ith available developmental data for mammalian dentitions. This method w'as employed, for example, in preparation of a model for evolution of ever-growing molars in certain microtine rodents (Phillips and Oxberry, 1972). In studies of phyllostomatid dentitions, this approach has offered some insights into the evolution of enamel-less, self-sharpening permanent teeth in the common vampire bat, Desniodus rotundus (Phillips and Steinberg, 1976). In this instance, histological and scanning electron microscopic analyses were compared with basic developmental information derived from studies on other mammals. Certain patterns of developmental biology could thus be shown to have been of preadap- tive value in evolution of enamcLless but cement-covered permanent teeth. Consequently, it should be possible in the near future to delineate an evolutionary' model incorporating the actual mechanisms that in a sense set the stage for the process of natural selection. In the case of Demiodus, it can be show-n that the preadapiive features that facilitated the evolution of a highly specialized dentition very likely included the following: 1) Mammalian teeth are not dependent upon enamel, or, more strictly, ectoderm for determination of tooth shape (Sicher and Bhaskar, 1972; Kollar, 1975; Osborn, 1973). Thus, even in absence of the enamel-producing 144 SPECIAI. PUB].[CAT[ONS MUSEUM TEXAS TECH UNIVERSU Y Fic. 9,—Va rial ion in the first upper molar in four difTerent specimens of Lkho/tycteris ohMitrn, From Phillips, 1971. ectodermal component, a mesodermal tooth germ will still result in an essentially normal tooth, insofar as the crown is concerned. This factor, which probably is common to mammalian dental morphogenesis, obviously allow's for selection of enamel-less, but otherwise normal, teeth in any given species. 2) Cementoblasts w'ill differentiate and produce a cementoid layer if enamel or dentin come into contact with periodontal tissue in the course of dental morphogenesis (Sicher and Bhaskar, 1972; Phillips and Oxberry, 1972). The coronal cementoid layer on permanent teeth in Desmodns apparently results because of a lack of protec¬ tive, reduced enamel epithelium. 3) Active attrition of coronal surfaces eventually would result in loss of a crown, w-ere it not for continuing eruption and con¬ sequent replacement of tissue. Based on studies of laboratory species, it generally is agreed that even non-ever-growing mammalian teeth normally continue to erupt throughout life (see Glickman, 1972). Teeth continue to shift position in response to a w'ide variety of factors subsequent to termination of the active erup¬ tive phase, during w'hich, teeth move into the occlusal eruptive plane. Cement added onto roots of teeth has the effect of maintaining the length of the rooted portion. In the common vampire bat, the anatomical root becomes crown as the original anatomical crown is worn away by thegosis, w'hich is the most sig¬ nificant component of bimodal attrition in this species (Phillips and Steinberg, 1976), The continuing search for solutions to evolutionary puzzles clearly requires analysis of phenotypic data within a framew'ork of developmental biology. In this regard w'e are fortunate that the needs of medical science have produced a relatively large body of supportive information, particularly in the area of oral anatomy. Although lack of information about controlling factors in morpho¬ genetic “fields” or gradients remains a major problem w'ith interpretation of apparently real relationships betw'cen individual teeth wdthin an arcade, it never¬ theless is apparent that combination of phenotypic realities with applicable data on developmental systems is a valuable trend in research. Dental Microanatomy Denial histology has been investigated in considerable detail in a variety of mammalian species, with exception of the Chiroptera. In view of the variety BIOLOGY OF THE PHVLUOSTOMAT[DAE US Fig. 10.—Compostte microanaiomical overview of maxillary leerh in a specimen of SfKniint ludoviii. Abbreviations are; AC, acellular cement: AF, apical foramen; C, crown; CC, cellular cement; CX, cervix; D, dentin; EA, epithelial attachment; F. fat cells; G, gingivum; ID. interdental septum: IGD, inlerglobular dentin; IR, interradkular septum; M, maxillary bone; PL. periodontal ligament; R. root; SG, minor salivary glands. Hematoxylin and eosin-Y. 19 X . of dental configurations found in the Phyllostomalidae, careful comparative histological studies not only are warranted but likely will prove valuable in ex- panding our knowledge about dental evolution and comparative oral biology. Fig. 10 presents an overview of the basic microanatom tea I features of the upper Jaw of a specimen of Sturnira. In this composite photograph one can study the spatial relationships and general histological characteristics of the tissues. Enamel Enamel typically provides the outer protective covering for mammalian teeth. Among the phyllostomatids, only the common vampire bat, De.smodus has been show-n either to lack enamel or at most to have an enantel covering only over the apical portion of newly formed permanent teeth (Phillips and Steinberg, 1976). In this species, continuing eruption and concomitant attrition due to ihcgosis, quickly eliminate the original coronal apices. Although its sister genera, Diaemus and Diphylla, have yet to be studied in this regard, it seems likely that they too have at least a reduced enamel layer. 146 -SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fiti. 11.— Lcfi: Scanning electron micrograph of occlusal surface of Ml in Artihvus Jitiniticfii'ii.'i. Note the enamel ridges and small surface inconsistencies or fractures (EF). 2000 X. Riiihn Scanning electron micrograph of fractured enamel in an upper canine from Gloxsopluiiid xork inti showing the interrow sheets (arrow s). I4S0 X . UitKhemicaJly, human enamel is known to consist mainly of inorganic mater¬ ials similar to apatite and to be scmipermeable (Sicher and Bhaskar, 1972; Zimmerman, 1968), As of this writing, essentially nothing is known about the biochemistry of enamel of the Phyllostomatidae or other chiropterans. Among the phyllostomatids, specific differences in feeding habits (and, thus, availability of calcium, phosphorous, and Vitamin D) and the general oral environments would suggest that this particular subject could provide an intriguing oppor¬ tunity for future comparative study. Relatively little is known about the detail of enamel structure in most kinds of mammals although a few' species are notable exceptions to this statement; in these kinds the structural arrangement of enamel has been investigated in¬ tensively by means of X-ray diffraction, histochemistry, and electron microscopy (for example, see Gustafson and Gustafson, 1967; Helmcke, 1967), General correlative studies (ordinal level) of cross-sectional configurations of enamel have been undertaken by Shobusaw'a (1952) and Boyde (1964, 1971). Accord¬ ing to Boyde (1971), enamel prisms in the Chiroptera are essentially the same as those usually found in the Cetacea, Sirenia, and Insectivora, He (Boyde, 1964, 1971) has described the enamel prisms as hexagonally packed, nearly straight in course, and having distinct cylindrical boundaries that allow tor easy recognition of interprism at ic regions. In the course of our investigations on phyl- lostomatids, only the long-tongued bat, OlifSsopha^Hi wrict/ia^ has been examined (Fig, 11), The portion of enamel shown w'as expo.sed by fracturing of an upper BJOLOGY OF THE PHYLLOSTOMATJDAE 147 Fk:. 12.— Left: Scanning electron micrograph of the anterior surface of an upper canine from Ariihi'its Jaftmice/iais showing ridgelike perikymata (arrows). The longitudinal fracture line at the left resulted from vacuum prtx:essing. 1 11 X, Right: Scanning electron micrograph of perikymata, surface scratches (A) and a groovelike inconsistency in the enamel surface (F). 370 X . canine, which occurred in a vacuum chamber during preparation for study with the scanning electron microscope. The course of the enamel prisms is apparent¬ ly nearly straight. The in ter prismatic material was found to be in the form of inter-row' sheets, some of which wxre left standing and clearly visible when the canine fractured (Fig. 11). This structural feature is at variance with what Boyde (1971) reported for the Chiroptera, how'ever, structural variation could easily cxicur between species wathin an order. Ordinal comparisons (Shobusawa, 1952; Boyde, 1971) have resulted in groupings that most certainly do not reflect phylo¬ genetic relationships. Thus, at least at the ordinal level, structural differences or similarities arc difficult to interpret and a given species probably cannot be considered indicative of an order without a considerable survey of species. Although the structural characteristics of enamel remain essentially unknown, the subject has considerable promise. Farney (1975) undertook a detailed scan¬ ning electron microscopic analysis of enamel of the J amaican fruit bat, Artiheus janiatcetisis. Surface features of enamel in phyltostomatids are poorly known. Prior to widespread use of the scanning electron microscope, teeth were studied by means of dissecting microscopes. This instrument is inadequate, however, because of the extremely small size and reflective qualities of the teeth. Examination of the outer enamel covering with the scanning electron microscope typically re¬ veals a coronal surface that is far from smooth (Figs. 11, 12). In adult sped- 148 SI'KX’IAI. PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY mens, one usually can itlenlify the per iky mat a, particularly on the sides of the crowns (Fig. 12). Perikymata, which have the appearance of being transverse, wavelike grooves, are regarded as external indications of the striae of Retzius (Sicher and Bhaskar, 1972). The ends of enamel rods and edges of lamellae arc visible in human teeth at various states of erosion (Scott and Wyckoff, 1949; Scott tT u/., 1949) but have not yet been observed in any of the phyllostomalids examined with scanning electron microscope. Numerous scratches and irregular grooves arc common, however, and easily visible (Fig. 12). Some phyilosto- matids, particularly Ariiheus, have notable surface modifications in the form of high interconnected ridges of enamel between the cusps (Fig, 11). These ridges become flattened due to abrasion and probably serve as treads that increase friction for holding hard rinded fruits in place when they are being crushed by the molars. Demiti Dentin is the major component of teeth in phyllostomatids. Although its chemical composition in these bats is unknown, in man it consists of approxi- mately 30 per cent organic material and water and 70 per cent inorganic material (Sicher and Bhaskar, 1972). The shape of the dentin is maintained by the organic constituents of decalcified teeth in histological preparations. The presence of mucosubstanccs is indicated by a PAS-po.sitivc staining reaction, which ranges from moderate to intense with phyllostomatids. Collagenous fibrils also are present in the organic dentin, as is indicated by green or blue staining of dentin with Masson’s trichronie or Mallory triple, respectively. In all phyHostomatids studied histologically thus far {Aniheus, Smrnira, Fhylh^.sNmnts^ Des/thHltLs, C/oi.vo/j/irigu, Leplotiycieris^ Attoum, and Ouyeronyc- ter/s), the coronal portion of the dentin is characterized by distinct dentinal tubules that follow a general S-shaped path from the pulpal chamber to the dentino- cnamel junction (Fig. 13). These tubules tend to be highly arborized, especially near the enamel junction. I bis especially is (rue in vampire bats {Desniodu-s) although there is an absence of a definite layer of enamel. In this particular spe¬ cies, the tubules begin to branch at approximately one-half the distance between the pulpal chamber and outer coronal surface. In contrast to the coronal dentin, dentin comprising the roots of teeth in phyllostomatids does not contain so reg¬ ular a series of tubules. Instead, the tubules are sparse and usually have a twisted and irregular course. In the healthy teeth from phyllosiomatid bats, the dentinal tubules contain a cytoplasmic process from an odontoblastic cell, positioned at the lining of the pulpal chamber (Fig. 13). Inlcrglobular dentin, which results from incomplete fusion during the process of dentinal mineralization (Sicher and Bhaskar, 1972), is found commonly in coronal dentin in phyllostomatids (Figs, 10, 14). Hypomineralized and liyper- mincralized globules stain differently due to the differing amounts of organic ground substance left after decalcificalion. It is perhaps noteworthy that differ¬ ences in amounts of interglobular dentin cannot readily be related to diet, at least based on comparison of the nectarivorous Lcpionycleris to frugivorous Aniheus or insectivorous vesperiMionid species. BjOLOGV oy THE PHYLLOSTOMATJDAE 149 Fig. 13.— Lcfi: Histological view of dentin and associated odontoblasts in Sturnira !ii(h)vic!. Abbreviations are; D, dentin; DT, dentinal tubules; FB, fibroblasts; OP, odonto¬ blastic process; PD, pre-dentin. Hematoxylin and eosin-Y. 942 X. Riyht: Scanning electron micrograph of fractured dentin in Glossopluif^’u suricitKi. Arrows denote dentinal tubules. 1900 X. Irregular, or reparative, dentin is found in instances where attrition is high due either to thegosis or abrasion. This type of dentin is formed quickly and serves to block the pulp chamber and, thus, prevent exposure of the pulp (Phil¬ lips and Oxberry, 1972). Among the various phyllostomatids that have been studied histologically, reparative dentin is found most commonly in the vampire bat (Desniodus). This undoubtedly is the result of relatively rapid, continuing eruption into the (x;eliisal plane and a reflection of the considerable thegosis that planes the permanent dentition (Phillips and Steinberg, 1976). Christian (1956) reported that he could determine age in the vespertilionid big brown bat {Eptesk us fusctis) by counting annular rings in dentin. According to Christian (1956), “a wide band of dentin is deposited during summer months followed by a dense zone during periods of hibernation," Although secondary dentin can be demonstrated histologically in most phyllostomatids, there is no evidence that it conforms to age except in a most general w'ay. Bands, or areas, of secondary dentin vary in number and w'idth both wdthin a given tooth and among different teeth. Overall, it can be stated that secondary' dentin, which usually is found adjacent to the coronal pulp chamber, seems most common in older individuals. Absence in the phyllostomatids of a marked annual physiolog¬ ical change such as hibernation essentially eliminates the possibility of using annual dentinal changes for age determination. 150 SPECIAI. PUBt.lCATiONS MUSEUM I EXAS TECH UNIVERSITY Fki. 14.— Lcfr. Histological view of inlerglobular dentin in Siurniru hidovici. Abbrevia¬ tions arc: CA. hypercalcified area; H, hypocalcified area. Hematoxylin and eosin-Y, 388 X. Rii^hn Inconsistencies in dentin mineralization in Desnitidus rottaniits. Pale areas (arrow) are hypomineralized; other abbreviations are: P, pulp; C, cement. Masson's trichrome. 232 X. Ccnientiuri The ccnientujTi is the softest of the three types of hard dental tissue. It is perme¬ able and in man at least, is comprised of about 55 per cent organic material and water and 45 per cent mineral (Sicher and Bhaskar, 1972). A cemenloid layer of variable thickness covers the roots of the teeth in the phyllostomatids. With the exception of Desmodtis, the pattern is essentially the same In all other species that have been examined. The cenientoid layer (along the sides of the root) is thin and acellular. Cementoblasls, characterized by small, ovoid, and somewhat hetcrochromatic nuclei, are found aligned along the root surface, even in adults (Figs. 15, 16). Typically, high magnification with the light microscope (1500X) reveals the elaboration of collagen into the cementoid tissue betw een the cemento- blasts and cementoid layer (Fig. 15). The weak, or negative, PAS response in the cementoid tissue apparently reflects the polymerization of connective tissue muco- substances (see Sicher and Bhaskar, 1972). Among the phyllostomatids studied thus far, the cementoid layer appears thin in the glosstjphagincs (specifically, Glossapha^ay Anoura, Lvptonyctvris^ and Chocronyaeris) and of moderate thickness in PhyUoKfomus, Anibeus, and Sturmni. A strikingly different situation is found in Desnuulus (Phillips and Steinberg, 1976). A thin layer of cement extends onto the coronal surface, where in ab.scncc of enamel it covers the dentin. The cementum covering the H]OL(.)GY OF THE PHYLLOSTOMATIDAE IM Fig. 15.— Tufr. Histological view of periodontal ligaments investing root surface iSsununt iiidovk i). Abbreviations are: C, cement: CB, nuclei of cementoblasts; FB, fibroblasts; PL. periodontal ligament. Hematoxylin and eosjn-Y. lllOX, Bottom: Histological view of periodontal ligaments investing alveolar bone (S. iudovkt)- Abbreviations are: A, alveolar bone; FB, fibroblasts; SF, Sharpey‘s fibers. Hematoxylin and eosin-Y. 1110 X. roots lends to be thick and is characterized by definite incremental lines (Fig. 16) that reflect periodic deposition in response to both stress and eruption. Al¬ though numerous workers have suggested (and, in some cases, actually tested) the value of incremental lines in cementum as a means of age determination (Klevezal and Kleinenberg, 1966), such a technique cannot be employed with DesmoditsiPhlWips and Steinberg, 1976). The numbers of incremental lines in the cementum varies both whthin teeth and, especially, among different teeth. Resorption and remolding of cementum, in addition to deposition of new ce- menium, can be related to dental drift by comparison to changes in adjacent atevolar bone. Dentai Pulp The pulpal cavity in mammalian teeth typically is filled with a complex soft tissue that is continuous with the periapical tissue through the apical fora- 152 SPECIAL PUBl 1CA [ ICJNS NfUSEUM TEXAS TECH UNJVERSJTY Fig. 16.— Top: Histological view of cementoid covering on upper incisors in Dcsinotiux Cememum (arrow) extends well onto the medial surface of the crown. Abbreviations are: A, alveolar bone; AB, alveolar bone resorption; C, cement: D. dentin: P, pulp. Hematoxylin and eosin-Y. 41 X, Bofro/n: Higher magnification of irregular incremental lines in cement (arrows). Abbreviations are: A, alveolar btjne: BC, blood vessels; CB, cementohlasts; [C, incremental cellular cement: P, peritnlontal ligaments. Hematoxylin and eostn-Y. I.S7X. Both photographs from Phillips and Steinberg (1976). men of the root. Thus, the pulp is a continuation of the soft tissues that surrountJ teeth and is histologically and, to some extent, biochemically similar to that connective tissue. It generally is agreed (Ogilvie and Ingle, 1965; Sicher and Bhaskar, 1972) that the pulp, which is histologically simple, but nevertheless dynamic, serves four basic functions; formative, defensive, nutritive, and sensory. The formative function of the dental pulp probably should be regarded as the principle one among the four. Basically, the pulp develops from a mesodermal aggregation of cells that gives rise to fibroblasts, which elaborate the collagen¬ ous pulpal matrix, and to odontoblasts, which are directly involved with for¬ mation of dentin. The latter process is initialed by interaction between odonto- BIOLOGV OF THE PHYLLOSTOMATIDAE 1,-^3 Fu.. 17— Left: Pulpal pathology with displaced odontohlasi nuclei (N) in a specimen of Gioxjufphiigii xoricino. Abbrevations are: D, dentin; H, histiocyte {macrophage); P, pulp; r, dentinal tubules. Hematoxylin and eosin^Y. I^44X. Righc Normal (healthy) pulp in Sturtiini htdovki. Abbreviations arc: C, capillaries; D, dentin; OB. odontoblasts. Hema¬ toxylin and eosin-Y. 760 X, blaslic cells and ectodermal elements during tooth development (Stanley and Ranney, 1962). The highly specialized odontoblasts line the walls of the pulpal cavity in the normal (healthy) stale and, thus, are a major histological feature (Fig, 17). In the phyllostomatid species studied, there were no interspecific differences apparent with light microscopy. As can be seen in Fig. 17, active odontoblasts are columnar in shape and have ovoid, heterochromatic nuclei. In specimens of bats displaying severe pulpal pathology of variable etiology, the odontoblasts frequently are flattened along their long axes. In fruit-eating phyllostomatids having relatively robust teeth (such as Ariiheus and Sturnira), the pulp most often is healthy and odontoblastic cells essentially normal. Based on the genera of Glossophagiiiae that have been studied {Lepioftycieris, Anouni, Glossopfmga, and Choen^nycteris), it is apparent that the pulp frequently is pathologic in wild individuals of these species. Each of these species is char¬ acterized by small teeth that are not especially important for feeding and, there¬ fore, not under considerable coronal stress. Nuclei of ixlontoblasts frequently are displaced and, in fact, located within adjacent dentinal tubules (Fig. 17). Such a displacement (or, perhaps, migration) is thought to result from an increase in intrapulpal pressure, due to inflammation or pulpitis and is an initial pulpal 154 SPECIAI. PUBLJCATIONS MUSEUM ] EXAS TECH UNIVERSITY Fi(i. 18.— Lefr. Scanning electron micrograph of pulp chamber in an upper third premolar of Lt>pionycteris nivaiU 1242X, R{!^>f)t: histological section 5 micrometers away from view shown on left. Hematoxylin and eosin-Y. I242X. Abbreviations for both are: D. dentin; OP. odontoblastic processes; OV, occluded vessel; N, nuclei of odontoblasts: T. dentinal tubule. reaction to injury (Ogilvie and Ingle, 1965; Beveridge and Brown, 1965; Ostrom, 1963). Although the body of the odontoblastic cells is most prominent histO' logically, a moderate amount of cytoplasm is incorporated within the dentin in the form of processes that extend from the wall of the pulpal chamber to the dentinoenaniel junction. In phyllostomatids, the processes were found to ar¬ borize considerably, especially near their termini. This feature apparently is typical in mammalian teeth (see Ogilvie and Ingle, 1965). The pulp of one species {Leptofiycteris nivalis) has been studied wdth the scanning electron micro¬ scope, following decalcificalion of the teeth, standard sectioning dowm to the desired level, and critical point drying. This material affords a remarkable view' of the odontoblastic cells and their processes (pig. 18). Additionally, it is pos¬ sible to compare the SEM photographs with a histological view, made from directly adjacent tissue. The odontoblasts in this instance w'ere flattened, pos¬ sibly as a result of certain pathologic features (thrombosis) within the pulp or adjacent to the rooted portion of the tooth (lesions in the oral mucosa). Among the odontoblastic prtxresses exposed by the retraction of the cells from the wall of the pulp chamber, w'hich is a common artifact of nonperfused histological preparations, at least one clearly is divided at its base. The process of this partic¬ ular cell apparently matched the paired dentinal tracts, also visible whth the SEM- In the SEM photograph, the i>dontob]asts appear to rest on a distinct, continuous membrane like layer that w'outd serve to separate the main portion of the cells from the wall of the puipal cavity. Such a structure has not been BIOLOGY OF THE PHYLLOSTOMATIDAE reported previously even in recent, detailed TEM and autoradiographic studies (see Weinstock and Leblond, 1974), and it thus is possible that what appears to be a continuous membrane actually is either the apical portion of the cell mem¬ branes of adjacent cells or, perhaps, a layer of predentin. Fibroblasts were found to be the second-most numerous cell type in the healthy pulp in phyllostomatids (Fig. 19). Such also is the case in man and rodents and, presumably, other mammals as well (Ogiivie and Ingle, 1965; Phillips and Ox- berry, 1972), Numbers of fibroblasts clearly w-ere greatest in developing teeth and least numerous in permanent teeth of old bats. Again, this pattern is essential¬ ly the same as in man and other mammals {Sicher and Bhaskar, 1972; Phillips and Oxberry, 1972). Histologically, fibroblasts are easily identified within the dental pulp of bats. I he nucleus is large, ovoid, and euchromatic. Additionally, the nucleus to cyto¬ plasm ratio is high and, therefore, the latter is difficult to distinguish. Collagenous fibers, w^hich are produced by the pulpal fibroblasts (Avery and Man, 1961), are found throughout the healthy pulpal stroma in phyllostomatids. Such fibers, which can be seen clearly with the PAS procedure, vary in abundance from in¬ dividual to individual, as well as from tooth to tooth within a given specimen. It seems likely that both age and position of individual teeth within the dental arcade have an influence on abundance of fibers (Sicher and Bhaskar, 1972; Stanley and Ranney, 1962). Staining of the pulp in Anibeus, Snirninu and Lcptonycietis with aldehyde- fuchsin alone or with aldehyde-fuchsin following oxidation with peracetic acid failed to reveal either elastic or oxytalan fibers. Neither type of fiber has been demonstrated in the dental pulp of other species of mammals {Ogiivie and Ingle, 1965). Korffs fibers, wliich are reticular and apparently become part of the collagen¬ ous matrix of the dentin (Bevelander, 1941), probably are present in the normal pulp of all phyllostomatids, especially in instances where new dentin is being formed. We have studied them only in Leptonycteris nivalis, w'hcre they were revealed by silver impregnation. Histiocytes, which can become macrophages, and lymphoid w'andering cells, are found regularly in apparently normal (healthy) pulp in the phyllostomatids (Fig. 19). Neither of these types of cells (especially the second) is so common as the fibroblasts, but, nevertheless, they are easily identifiable. Both the histiocytes and lymphoid cells have defensive roles; the former remove dead cells, bacteria, and foreign materials by phagocytosis and the latter possibly are a source of antibodies and plasma cells (Zander, 1946; Bloom and Fawcett, 1968). The dental pulp and dentin are nourished by a complex netw'ork of blood vessels, some of which are extremely smalt (Figs. 18, 19). The details of pulpal circulation in phyllostomatids are as yet unstudied but it is likely that, as in man, the pulpal capillaries are contractile with numerous arteriovenous con¬ nections (A-V shunts) as well (Han and Avery, 1963). In contrast to man (Ogiivie and Ingle, 1965; Provenza, 1958), the phyllostomatids apparently lack a distinct subodontoblastic plexus and zone of Weil. Such a plexus and cell-free zone are thought to reflect the nutritive requirements of the odontoblastic cells. Possibly, i56 SPECIAL PUBI.ICAI IONS MUSEUM TEXAS TECH UNIVEKSI TY Pic. 19.—Hisitologlcal examples of typical componems of healthy pulp in a molar from an adult Siurriini !ndovki\ small collagenous fibrils are labeled with an F. Hematoxylin and eobin Y, Both are 3300 X. the extremely small size of the puJp chamber in these bats negates the impor¬ tance of a subodontoblastic plexus. On the other hand, a distinct zone of Weil is found in the pulp of the molossid genus Tadarkla, which also has an extremely small pulpal chamber (Phillips, unpublished data). Two circulatory abnormalities frequently are noted in the phyllostonialids. Many specimens contain a high percentage of dental pulpal blood vessels that BIOLOGY OK THE PHYLLOSTOMATIDAE 157 are either totalJy or partially occluded by a thrombus or hyaline material (Fig. 18). Extensive hyperemia (dilation of capillaries) is commonly found in speci¬ mens of Phy!lost(mu(.% Siurnira, and Lepronycieris. This condition has been shown to be either transitory or an indication of early pulpitis (Ogilvie and Ingle, 1965). Which is the case in a given specimen is not always clear. I he sensory aspect of the dental pulp in phyllosiomatids is as yet unstudied. The pulp undoubtedly is richly innervated and it is likely that, as in man, sensory nerve endings extend peripherally into the layer of odontoblastic cells and, pos¬ sibly, into the predentin (Ogilvie and Ingle, 1965; Sicher and Bhaskar, 1972). PeriodonffUfn The periodontal membrane consists of connective tissue fiber bundles that connect teeth to surrounding alveolar bone. Connective tissue cells, blood and lymphatic vessels, nerves, and a variety of intercellular substances form the prin¬ ciple constituents of this membrane. In previously studied mammals, collagen, oxytalan and elastic fibers, and acid mucopolysaccharides have been found extra- cellularly (Fullmer, 1967). Embryologically, the periodontal ligaments are de¬ rived from the dental sac that envelops the developing tooth. The fibrous con¬ nective tissue of the sac differentiates into three layers; the outer is adjacent to alveolar bone, the inner lies adjacent to the cemenium, and the intermediate layer of unorganized fibers becomes rearranged and thickened to periodontal ligaments (Orban, 1927; Glickman, 1972), Groups of collagenous fibers that form the prin* cipal fiber bundles can be grouped according to orientation (Sicher and Bhaskar, 1972). Gingival ligaments attach the gingtvum to cementum; trans-septal (inter¬ dental) ligaments are found between adjacent teeth; and a group of alveolodenial ligaments connect cementum directly to alveolar bone. Principle fiber bundles representative of each of these categories, have been demonstrated in all species of phyllostomatids thus far studied. It is apparent, however, that among the alveoiodental group, the oblique fiber bundles are least common. Instead, the principle fiber bundles frequently can be traced from within alveolar bone (Sharpey’s fibers) to their point of attachment in cementum (Figs. 15, 20). Addi¬ tionally, scanning electron microscopy reveals at moderate magnifications (4000 X) that many of the principle fiber bundles are interconnected by a net- w'ork of fine connective tissue fibers (Fig. 21), The exact structural constituents of the fine interlacing fibers of the phyllostomatid periodontium are unknown, and cannot be determined by scanning electron microscopy alone because the SEM docs not reveal the typical periodicity of collagen. Comparison of the SEM micrographs of Lephmycteris to those of other mammals (for example, man, dogs, and armadillos) suggests that these interlacing fibers represent the indif¬ ferent fiber plexus described by Shackleford (1971), who suggested that the den¬ sity of the indifferent fiber plexus is directly related to amount of stress placed on the coronal surfaces. In the specimen of Lx^ptonyaeris illustrated in Fig, 20, staining of the histological material with silver impregnation indicates that at least some of the fine interconnecting fibers might be reticular. Two types of reticulin (precollagenous and basement membrane) usually are present in the periodontium (Fullmer, 1967), Both are found in phyllostomatid bats. The pre- 158 SPECIAL F>U plications MUSEUM I EX AS TECH UNIVERSITY pKi. 20.— Top-. Histological view of components of the periodonlium (in Artiheus jamuicensis). Abbreviations are: EC* endothelial cells; HR, epithelial rest; FB, fibroblast. Hematoxylin and eosin-Y. I284X. HoHom-, Silver impregnation staining of collagenous principle fiber bundles (arrows) in Ixpiioxycteris nivalis. Abbreviations are: A, alveolar bone; C, cement; D, dentin. 375 X. collagenous (or argyrophil) reticular fibers become black with silver impregnation (Fullmer* 1967; Lillie, 1965). Consequently, they can be distinguished fairly easily from the collagenous principle fiber bundles, which are larger and stain pale red-brown or violet. Few species of phyllostomatids have been studied with silver impregnation, and, thus, no specific comparisons of numbers and distribution of periodontal reticular fibers can yet be made. 1J[OLOGY OF THE F^HYLLOSTOMATEDAE [59 Fig. 2[.^L(’ft: Scanning electron micrograph of typical periodontium in a specimen of Lrpfttnycferis nivaHs. Abbreviations are; A, alveolar bone; BV, blood vessel; D, dentin; PL, periodontal ligaments. 700X. Detail of periodontium showing fibroblasts (F), collagenous principle fiber bundles (PF), and "reticular" fibrils of I he indifferent fiber plexus (R). 2800X. Overall, the principal fiber bundles of representatives of the genera Artiheus, PhyUostomHS, Sfi^rfiirciy Desmodus, Lt'pfonyctens, Anoura^ and Chocrofiycteris are essentially the same in basic histological features, staining reaction, density, and distribution. Observed individual differences generally were relatable to unusual or pathological conditions, Fullmer and Lillie (1958) and Fullmer (1967 and elsewhere) have found and studied in detail a type of connective tissue fiber that they termed oxytalan fiber. Although these fibers, which apparently are related to elastic fibers and possibly are their precursors, arc common in the periodontium of several unrelated species of mammals, they have not been found in the periodontium of phyllostomatid bats. Staining with aldehyde fuchsin follow-ing oxidation with peracetic acid has revealed a relatively small number of oxytalan fibers in the submucosa of the oral epithelium in phyllostomatids. Absence of oxytalan fibers in the peri¬ odontium of those species in which the teeth are under considerable stress (for example, Desmodus', Phillips and Steinberg, 1976) is of interest in view of the fact that these fibers reportedly are abundant in high stress regions in other mam¬ mals (Fullmer, 1967). In low stress situations such as developing teeth or adult teeth in such species as the armadillo, the oxytalan fibers, if present, lend to be nonatigned (Shackleford, 1971). The cellular components of the periodontium of phyllostomatids are e.s- sentially the same as in other mammals. Long, slender, fixed fibroblasts com¬ prise the majority cell type directly associated with periodontal ligaments in phyllostomatids having normal (healthy) periodontium. These cells have large, 160 SPEC] A I- PUP] ICATIONS MUSEUM TEXAS TECH UNIVERSITY ovoid nuclei (Figs. 15, 20) and appear, in hisiological preparations, to be inter¬ spersed with principle fiber bundles. The cytoplasm, which is eosinophilic and sparse in comparison to the nucleus, is difficult or impossible to distinguish with standard stains such as hematoxylin and eosin. When studied by scan¬ ning electron microscopy, the stellate shape of fixed fibroblasts can be readily di.stinguished (Fig. 21). Usually these cells are associated with principle fiber bundles; elongate cytoplasmic processes and formative bundles of collagen (re¬ ticular fibers) are visible at 4C)()()X. Cells resembling fibroblasts but having smaller, more elongate nuclei frequently can be found in association with blood vessels, particularly capillaries, in the periodontium of adult (fully grown) phyllo-slomatids. These cells apparently are undifferentiated mesenchymal cells that persist into a bat's adulthood. If they are in fact mesenchymal cells, it is likely that they arc invportant in response to local innammatory or pathological conditions (Bloom and Fawcett, 1968). The periodontal ligament is known to serve as the periosteum for alveolar bone (Fullmer, 1967). It is not surprising, therefore, that osteoblasts and osteoclasts are found in the periodontium adjacent to alveolar bone in phyllostomatid bats. The presence of these cells is related either to apposition of new bone or resorp¬ tion of old bone and, thus, their occurrence frequently rellects degree and direc¬ tionality of stresses on teeth. The general phenomenon of alveolar responses to stresses has been studied intensively in other mammals, particularly with regard to human dental medicine (for example, sec Fullmer, 1967; Sicher and Bhaskar, 1972). Variations in activities of cellular enzymes (certain dehydrogenases and nonspecific esterases) have been related to the process of bone remolding (Glick- man, 1972). Marked alveolar resorption is common, for example, adjacent to teeth that have become nonfunctional due to acute dental caries in PhyUostomus hasKitiiS (Phillips and Jones, 1970). Atrophy of periodontal ligaments also re¬ sults from loss of function and is common in this species, which exhibits an un¬ usually high incidence of dental caries in nature. Resorption of alveolar bone in conjunction with periodontal innammaiion caused by infestations of oral mites also is common in three giossophagine genera {Lepmuyiteris, Anoum, Mouo- phyll{is)\ in these species, large numbers of multinucleate osteoclasts are found adjacent to alveolar bone (Phillips et cv/., 1969; Phillips, 1971), Among the phyllostomatids thus far studied, alveolar bone remolding (Fig. 16) typically is most extensive in the common vampire bat, Desnuxias nyitmdus (Phillips and Steinberg, 1976). In this instance the teeth undergo considerable coronal stresses as a result of thegosis (tooth sharpening) and, therefore, drift and continuing enjptive movements into the occlusal plane have considerable effect on alveolar bone. Cementoblasts are yet another type of cell commonly found in the periodontium of phyllostomatid bats (Figs. 15, 16). They typically are found along the surfaces of roots and are characterized by spherical or slightly ovoid nuclei. As visible in Fig. 15, they are notably smaller than fibroblasts and are easily distinguished. Cemenlogenesis, especially at root apices, apparently con¬ tinues throughout the life of individual bats. Resorption and deposition of cement li to LOGY OF (HE PH YLLOSTOMATIDAE 161 is intluenced by essentially the same factors that influence remolding of bone (Gtickman, 1972). For example, marked deposition by cementobIa.sts was found to be a usual response to extreme coronal stress in PhyiUfsiofnushasnuiLS (PhllWps and Jones^ 1970) and Desmodus rotund us (Phillips and Steinberg, 1976), Alveolar Process The portions of the mandible and maxilla that support the sockets into which teeth are set are termed the alveolar process. In man and other primates for which data are available, the process can be divided into two components; alveolar bone, which surrounds the roots and provides a site for attachment of the peri¬ odontal ligaments; and supporting alveolar bone, which consists of both cortical plates (compact bone) and spongy bone (see Sicher and Bhaskar, 1972). in phyllosiomatid bats, the alveolar process differs somewhat from this model in that spongy bone is minimized, and the typical interdental and intcrradicular septa are characterized by extensive amounts of compact bone (Figs. 10, 16). The only exception among the species that have been studied histologically seems to be the large spear-nosed bat {P/iyilostomus hasuuus). In this species, one can find moderately extensive spongy bone, hut, even so, the specimen of Siuniira illus¬ trated in Fig. 10 is a better example of the typical alveolar process in phyllosto- mat ids. Oral Epitludlum ami Gintfivuni The oral mucous membrane is a structuraily variable lining of this cavity in mammals. The morphological characteristics of the membrane are thought to reflect specific functions or mechanical influences related to position within the oral cavity as well as to feeding habits (Sicher and Bhaskar, 1972). The oral epithelium of phyllostomatid bats has not been studied previously in detail. Structurally, the oral mucous membrane can be divided into three components: an outer layer of stratified squamous epithelium; an intermediate lamina propria; and a variable submucosa layer comprised of connective tissue. In all species of bats studied for this report, the squamous epithelium was found to consist of a basal cell layer, an intermediate cell layer, and either an orthokeratinous (totally keratinous), parakeraiinous, or nonkeralinized outer surface (Figs, 22, 23). The gingivum, which is the portion of the oral mucous membrane that surrounds the teeth and, thus, usually is subjected to a variety of mechanical forces, and the prominent palatal ridges were found to be either orthokeratotic or parakerototic. Orthokeratoiic tissue typically is comprised of scalelike cells apparently lacking nuclei (Fig. 23), whereas the parakeratotic layer is characterized by the presence of naitened, heterochromatic (hematoxylin and eosin) nuclei within the epithelial cells (Fig. 22). In the phylloslomatids, the gingivum Is most commonly para- keratoiic, whereas the palatal ridges are about 50 per cent parakeratotic and otherwise orthokeratotic. As might be predicted, the degree of keratinization of the hard palate in these bats can be related directly to food habits. Consequently, the hard palate of Artibeus jamaiceusis, which feeds on fruits that sometimes have 162 SPECIAL PUPL[CATrONS MUSEUM PEXAS TECH UNIVERSITY Fifi, 22.— Left: Histological overview of gingivum and oral epithelJum of the lip in a specimen of Anouru }*eoffroyi. Abbreviations are: A, alveolar bone; AT, adipose tissue; D, dentin; EA, epithelial attachment; EC, epithelial cuff; E, epithelium; M, muscle; P, pulp; PL, periodontal ligaments; V, bases and shafts of vibrissae. Hematoxylin and eosin-Y. 63 X. Ri^ht: Gingivum in Leptoncyteris nivitUs. Abbreviations are: B, basal cells: K, keratinized surface; P, prickle cells; PA, papilla; N|, euchromatic nuclei of prickle cell; Na, heterochro- malic nuclei of perakeratolic layer; arrow, space between prickle cells. Hematoxylin and eosin-Y. H76X. hard rinds, is considerably more keratinized than is the hard palate of Lep- toiiyctetis, which feeds almost exclusively on pollen and nectar. Ii seems possible that keratinization is a local response to mechanical stress, and, thus, the more mechanical force applied, the thicker and more keratinized the outer layer of epithelium. The basal cell layer of the mucous membrane, which is separated from the lamina propria by a distinct PAS-posittve basement membrane, is structurally variable within a given bat. Generally, the cells are cuboidal and the nuclei rela¬ tively euchromatic (Figs. 22, 23). On the other hand, in some areas (particularly the gingivum), the cells tend to be elongate with relatively little cytoplasm and heterochromatic nuclei. A prickle cell layer also is present, as it is in other mam¬ mals. In man, this layer of stratified epithelium is most distinguishable in areas of orthokeratotic or parakeratotic mucosa. When viewed with oil immersion microscopy, large numbers of intercellular '‘bridges” or tonofibrils are easily distinguishable in phyllostomatids {Fig. 24). In the specimen of a long-nosed bat {Lepionycteris nivali.s) showm in Fig. 24, the tonofibrils connect the elongate ceils HIOLOGY OF THE PH YL.LOS'l OMATIDAE J63 Fkl l'S,-^Lvfr. Scanning electron micrograph of orthokeratotic layer (K) in Leptonyaeris nivalis. Nuclei of prickle cells also can be distinguished in this view (N). 1850X. Ri^hv. Oral epithelium in Snirnirn Ituiavki showing round, euchromatic nuclei of basal cells (Ni). elongate euchromatic nuclei of prickle cells (N 2 ), and a single heterochromatic nucleus (Na) of a cell within the otherwise orthokeratotic outer layer. Hematoxylin and eosin-Y. I628X. across a distinct intercellular space, forming a network of inlerfacial canals. Al¬ though these cells in phyllostoniatids have not yet been investigated by means of the transmission electron microscope, it is likely that the arrangement of tono- fibrils and desmosomes is similar to that in other mammals (Bloom and Fawcett, 1968). The orthokeratotic outer layer differs from the parakeratotic layer not only in presence or absence of nuclei, but also in staining characteristics. With the stan¬ dard hematoxylin and eosin-Y, for example, the orthokeratotic layer is pale yellow or essentially not stained, whereas the cytoplasm of cells comprising the parakeratotic layer stains the characteristic pink. The scanning electron micro¬ scope provides a striking view of structural features of an orthokeratotic outer layer (Fig. 23). The extremely flat, scalclike condition of the cells can be seen readily; in this instance, which is representative of the masticatory mucosa of the hard palate of a specimen of Lepumyctens nivaiis, at least seven distinct layers can be distinguished. Dark, ovoid areas below' the orthokeratotic surface layer, represent the nuclei of prickle cells, apparently altered by critical point drying or some other aspect of the SEM technique {Fig. 23). Fullmer and Lillie (1958) discovered and studied oxytalan fibers in the periodontium of several species of mammals. In the course of the present study it was found that txcasionally the outline of cells in parakeratotic layers of the oral epithelium of phyllostomatids 164 SPECIAl- PUB] iCATtONS MUSEUM TEXAS TECH UNIVERSITY Fig* 24, — Histological view of I he prickle cell layer in oral epithelium of Leptonyvteris nivalis’, these cells are characterized by ovoid cuchromatic nuclei (N) and interconnecting tonofibrils (arrows). Hematoxylin and eosin-Y. 2500X . would Slain deep purple following the peracetic acid-aldchyde fuchsin procedure for oxytalan fibers. The reason for this reaction is unknown, although Fullmer (personal comnuinication) also has noted such a reaction in other kinds of mam¬ mals. None of the other staining techniques employed in this study (see the sec¬ tion on materials and methods) produced similar results. The epithelial atlachement to teeth has received considerable attention from dental researchers (for example, see Sicher and Bhaskar, 1972; Loe, 1967; Glick- man, 1972, for citations). The area of attachment has the form of a cuff in which debris and bacteria can accunuilate and, thus, is a primary site for inflammation. In the phyllostomatids, the actual form of the epithelial attachment varies in¬ dividually as well as inierspecifically. Most commonly, the junction between enamel and cementum provides the site of attachment (Fig. 25). In some speci¬ mens the attachment is below this level, probably as a consequence of continuing eruption into the occlusal plane. In vampire bats {Desmodus), as show-n in Fig. 25, the epithelial attachment differs dramatically because the permanent teeth lack enamel (Phillips and Steinberg, 1976). In Desmodus, the attachment always is directly to the cementoid layer. Mast cells and numerous polymorphonuclear leukocytes, indicative of in¬ flammation, generally arc found within the area of the epithelial attachment to teeth (Fig. 25). It seems likely that low grade inflammation is a normal state of affairs in phyllostomalid bats. B[OI,OGY OF THE PHYLLOSTOMAT[DAE Fig. 25.— Top left: Epithelial attachment in Desmodns rotundus. Note the cementoid layer that extends up the coronal surface. Hematoxylin and eosin-Y. 148 X. Lower lefv. High magnification (light microscope) view of granular mast cells in the gingivum of DesmodtLs rottouius. Hematoxylin, eosin-Y, and PAS. 2072 X. The epithelial attach- ment in Ghssopha}>u sorivUuh which is more typical of phyllostomatids because of the lack of coronal cemenlum. Hematoxylin and eosin-Y, 370 X. Abbreviations for all illustra¬ tions are: C, cementum: D, dentin; EA. epithelial attachment; ES, enamel space; E. epithelial cells; EP, epithelium; M, mast cells; arrow, cementoblasts. Pathology Ora) and dental disease in wild mammals has not been studied to the extent warranted by the potential value of such data to our understanding of sysiemaiic and evolutionary biology. Among the several papers dealing with oral and dental diseases in w'ild species, the volume by Colyer (1936), although somewhat outdated, is by far the most complete. Publications by Hall (1940, 1945) also are valuable even though limited to relatively few species. Insofar as the Chiroptera are concerned, the only available information (aside from incidental comments) is that published by Phillips and Jones (1969, 1970), Phillips et ai. (1969), and Phillips (1971). The first of these papers deals with nonphyllostomatid bats, the second with Phyikmornits hcisfams and the last two w'ith glossophagincs only. Consequently, the following overview reflects this disparity and is based pri- 166 SPECJAl- PUHl.JCATIONS MUSEUM TEXAS TECH UNrVERSfTY mariJy on Phillips (1971), although a considerable amount of previously un¬ published histological data on periodontal disease has been included here. Dental Caries Dental caries involves many etiological factors, including bacteria, substrate characteristics (diet), and surface and structural aspects of teeth (Keyes and Jor¬ dan, 1963). Although this disease has been reported from representatives of many mammalian orders (Colyer, 1936), it appears to have a differential incidence in nature. Among the Chiroptera, dental caries possibly is uncommon, although gross identification of carious lesions in teeth of museum specimens does not necessarily provide an accurate picture of what happens in nature. In 1508 specimens from the families Emballonuridae, Noctilionidae, and Mormoopidae, Phillips and Jones (1969) found only one individual (a specimen of Mornioops tfie^^'alophylla) in which carious teeth could be recognized at the relatively low magnification (20 X ) used for the examination. The spear-nosed bat, Pfjyilosnnnns /uistatns, is clearly an exception because in this species dental caries is extremely common (Phillips and Jones, 1970). A 40 per cent incidence of caries was found in 52 specimens; additionally, an un¬ explained significant (P^O.99) difference in incidence w-as found between males (75 per cent) and females (19 per cent). Specimens of P. hasratns were compared to a series of 103 specimens of P. discolor, in w'hlch gross carious lesions w-ere lacking, in an effort to isolate some of the possible endogenous and exogenous factor.s involved. Certain specific differences between these species could be correlated in a general way with the surprisingly high incidence of caries in P. hastatiis. For example, in this species the large and robust teeth frequently ex¬ hibited stained fissures within which lesions usually w'ere located. Additionally, the structure of the oral mucous membrane and gingivum at the posterior end of the tooihrow was such that debris tended to accumulate there. The significance of a high incidence of dental caries in P. haskitifs is as yet unclear but two possible interpretations have been offered (Phillips and Jones, 1970): 1) one or more genetic factors that result in a high incidence of dental caries are associated with some characteristics, for example large size, that are of significant survival value; 2) some exogenous factors, such as an e volution- arily recent shift in food habits, are of considerable cariogenic importance, and selection has not yet produced a phenotype capable of ameliorating these environ¬ mentally produced conditions, or that the adaptive value of exploitation of a new' food source is of far greater benefit to the species than is prevention of dental caries. The study of positive allometry between body size and width of teeth dis¬ cussed in the section on evolutionary mechanisms (Pilbeam and Gould, 1974) can be applied to this particular problem. Selection for large size clearly has oc¬ curred in the evolution of P. hastaius. The large and robust teeth in this spe¬ cies probably are the result of positive allometry. An essentially “automatic” in¬ crease in tooth size In absence of modification of morphogenetic patterns, changes in mineralization, or even modifications of rate of development could w'ell ac¬ count for the absence of an effective system for modulating incidence of dental caries. BIOLOGY OF THE PHYLLOSTOMATIDAE 167 Carious lesions also have been reported in teeth of several species of glos- sophagine bats (Fig, 26) but are regarded as rare in incidence (Phillips, 1971). It is especially important to rementber, however, that the teeth in these bats are so small that recognition of dental caries is limited to relatively severe lesions. Sur¬ vey of specimens of glossophagines by means of the SEM would lead us to be extremely cautious about determination of rarity of this tbrm of pathology. For example, carious lesions were not found in examination of several hundred speci¬ mens of Lepfonyctens nivalis and L. sanhonti by means of a dissecting micro¬ scope (Phillips, 1971), but in three specimens of L. nivalis examined with the SEM, carious lesions were readily visible in molars of each individual. The lesions always were within basins and were characterized by loss of enamel and exposure of a surface of primary dentin (Fig. 27), Examination at 1500X re¬ vealed a general absence of dentinal tubules although in at least one lesion a group of tubules could in fact be seen. The overall appearance of the dentin ex¬ posed in these lesions was remarkably similar to that exposed by either of the components of bimodal attrition. A wide variety of materials, some of which could not be identified with certainly, were found wdihin carious lesions. Small spherical objects, measuring approximately one micrometer, were found in large numbers within the carious lesions. These had the size and appearance of coccus type micro organisms (Fig. 27). Two configurations are shown in Fig. 27; several fairly large clusters are clearly visible as well as several pairs of or¬ ganisms. The presence of coccus type bacteria at sites of carious activity is not surprising. Several kinds of acidophilic microorganisms, including lactobacilli and streptococci have been shown to have cariogenic potential (Orland ei al., 1955; Fitzgerald and Keyes, 1960; Keyes and Jordan, 1963) in primates and rodents. Nothing is knowm about the microtlora of the oral cavity of glos- sophagine bats and, therefore, identification of the microorganisms must aw^ait an opportunity for preparation of cultures. The paucity of exposed dentinal tubules in the carious lesions probably reflects the fact that calcium salts are deposited around exposed, degenerating odonto¬ blastic processes resulting in obliteration of the tubules (Sicher and Bhaskar, 1972). Obliteration of tubules is of clear-cut functional significance; the tubules not only can become packed with microorganisms (Johansen, 1963) but, more importantly, they communicate directly with the dental pulp. Periodotual Diseasc Periodontal disease involves inflanimation of the gingivurn and periodontal membrane and often leads to destruction of alveolar bone and dental tissue. This form of pathology probably is the major cause of loss of teeth in mammals (Coi- yer, 1936; Fullmer, 1966; Glickman, 1972). A w'ide variety of etiological factors have been associated with periodontal disease: these include microorganisms, calculus, oral hygiene, vitamin insufficiencies, irregular teeth, and salivary gland secretions (Fullmer, 1966; Klinkhamer, 1968), Protein deprivation and certain enzymes (kaliikrein, for example) also apparently play roles as etiological factors or mediators of periodontal disease but are as yet only poorly known (Sw'eeny, 1966; Narrod and Braunberg, 1966; Fullmer, 1966), SMEC[AL PUBLtCATJONS MUSEUM TEXAS [ECH UNIVERSITY Fkn 26.—Occlusal and labial views of dental caries in a glossophagine bat {Afiostm ^i'offroyi) Symbols are: a, carious lesion; b, exposed carious dentin; c, normal (noncarious) molar; d. wear facet; e, edentulous maxillary; f. exposed, partially resorbed root; g, h, areas of bone resorption. From Phillips (1971). Among the phyllostomatids, only the gJossophagines have been surx^eyed for periodontal disease (Phillips, 1971). Aside from a common localized rarefying osteitis, typical periodontal pathology is relatively rare in these bats, at least insofar as can be determined by examination of cleaned skulls in museum col¬ lections (Phillips, 1971). An unusual periodontal disease, thus far known only in giossophagine bats, has been reported and studied previously in some detail (Phillips e! at., 1969; Phillips, 1971; Radovsky ei al., 1971). In this disease, the major etiological factor is the presence of protonymphs of mites of the genus Radfordiella (Macronys- sidae), which become embedded in lesions in the oral mucosa and gingivum (Figs. 28, 29). The resultant pathological condition, which includes destruction of soft and hard oral tissues, frequently leads to exfoliation of teeth in life. The mites are highly host specific and consequently the presence or absence of pathology can actually be used as an unusual taxonomic character in separating Lep- lonycteris nivalis from L. sanburni. In Mofiophyiias, which is endemic to the Antilles, oral mites are found in M. rednuuii but have not been reported from M. pieihodon, in contrast to Leptonycteris nivatis, however, the incidence of in- mOLOGY OK THE PHYLLOSTOMATIDAE 169 Fig. 27. —Scanning electron micrograph of a carious lesion in a molar from a Mexican long*nosed bai, Ltpiatnycteris /uV(y//.v, Abbreviations are: RBC, red blood cells: DT. dentinal tubules; M I. paired microorganisms; M2, clusters of microorganisms. 1 140 X . testation in rednutni apparently is geographically variable (Phillips, 1971). Afioura is the third genus in which Radfonliella infests the oral cavity; once again the miles (both Racifardiella oricola and R. ^inourae) are found only in one species {A, ^ieoffroyf) but seem to have geographically variable incidence (Phil¬ lips, 1971). The reasons for species*specific and geographically variable incidences of infestation are as yet unclear. The long-tongued bats of the genus Lepfonyaeris have been examined most closely in this regard. As of this writing, no ecological factors or distributional characteristics provide good candidates for explana¬ tion of why only L nivalis is infested. The two species are partially sympalric and apparently have similar feeding habits (see chapter by Gardner, this volume). Many mammalogtsts have had difficulty distinguishing between the species, as is rejected by confusing and inadequate taxonomic arrangements suggested over the past 30 years (Martinez and Villa-R., 1940; Hoffmeister, 1957; Villa-R., 1967; Alvarez, 1966). In point of fact, however, the three currently recognized species not only are distinctly different in several phenotypic characters, but also are relatively easy to identify (Davis and Carter, 1962; Phillips, 1971). Superficial similarities between L. nivalis and L, sanhorni belie some apparently noteworthy differences in their oral anatomy and, indeed, their oral environ¬ ments. For example, light-level microscopic comparisons of their salivary glands has revealed structural differences, particularly in the parotid gland (see section on salivary glands). These differences are far from minor and carry through to the ultrastructural level (B. Wilder, personal communication). It is reasonable to assume, of course, that differences in subceilular structure and in structural char¬ acteristics of secretory granules suggests in turn that the oral cavities in these two species are bathed by a considerably different saliva. The role of saliva in miti- 170 S/>EC]AL PUH Lie AT IONS MUSEUM TEXAS TECH UNIVERSITY Fig. 28.— RadfordicHa oricola in the mouth of a Mexican long-nosed bat, Li’ptonyctet h niviifi'i. Symbols are: a, palatal ridge; b. upper premolar: c, mites. From Phillips (1971), gating disease and controlling both inflammation and microflora is well estab¬ lished and currently attracting considerable attention from dental researchers (For instance, see Klinkhamer, 1968). Most striking, perhaps, was the finding of Greenbaum and Phillips (1974) that the tongue of L. sanhomi, which species lacks oral mites, is characterized by highly keratinized, recurved hooks positioned so as to .scrape the lingual gingivum and adjacent oral mucosa (see section on tongues and asscxriated musculature). Anatomical and environmental differences such as these do not explain apparent geographic differences in incidence of in¬ festation within species, even though they certainly provide a basis for explaining species specificity of the oral miles. Tissue responses to infestations of oral miles are of special interest because of a general paucity of data on effect of parasites (Lavoipierre ei ai, 1967; Lavoi- pierre and Rajamanickam, 1968) and the potential, in this instance, for studying specific responses of periodontal and hard dental tissue. The results of initial BIOLOGY OF THE PHYLLOSTOMATIDAE 171 FiCi. 29 ,-—Scanning electron micrograph of Rudfordiella on'a^la in the mouth of a Mexican long-nosed bai, Li^pronycterts nividis. Abbreviations are: B, red blood cells; C, connective tissue: F, foreign (plant) materia); L, lymphocytes; G, gnathostome of mile- 533 X. studies of peritxiontal pathology associated with infestations of Radfordiella oricoki in Li'ptonycieris nivalis have been published (Phillips ei a!., 1969; Phil¬ lips, 1971). In the following paragraphs we have reviewed these findings and have included new, unpublished information based on both general histological analysis and studies with the scanning electron microscope. 172 SPEC[Al. PUBLICATtONS MUSEUM TEXAS TECri UNIVERSITY Macronyssid oral mites most frequently enter the oral mucosa adjacent to the first and second upper prcmolars. Occasionally, especially in cases of severe in¬ festation, the protonymphs also are found adjacent to other upper teeth or even along the midline of the palate. The resultant lesions (Figs, 28, 29) can contain up to at least 200 individual mites. Oral epithelium around the lesions is char¬ acterized by destruction of the orihokeratotic outer layer as well as the granular, prickle-cell, and basal layers. At the same lime, however, the size of the lesions apparently is in some way limited and does not continue to increase even in unusually .severe infestations. It is possible, of course, that size of lesions is limited by the mites themselves in response to some density-dependent factor(s). Average palatal and alveolar lesions are approximately 1.0 millimeters in length and width and 0.5 in depth. The edges of lesions grossly appear rounded; histologically it can be demonstrated that oral epithelium grows inward and, thus, provides a partial lining for the walls of the lesion. Oral submucosa or periodontal mem¬ brane (depending on the position of the lesion) or both are exposed in the bottom of the lesion. The mites do not burrow into the connective tissue; instead they become packed into the lesion with their gnathosomes and first pair of legs pene¬ trating the mass of connective tissue remanents and free cells at the bottom of the lesion (Figs, 29, 30). These parasites appear to be highly mobile and it is likely that they transfer from one lesion to another within the oral cavity. Indeed, wiien the mouth of a living specimen of Lepn^fiyaetis nivalis is forced open, it is pos¬ sible to observe the mites as they move about w ith considerable agility. Histologically, the lesions typically are packed with large numbers of lymphocytes and necrotic cells (Figs. 29, 30). Relatively few plasma cells tx;cur within or adjacent to these lesions. Staining with azurc-A and eosin-B also has revealed coccus-type bacteria wdthin adjacent oral epithelium and within the epithelial cuff of teeth adjacent to lesions. Additionally, microscopic plant materials {possibly spines from agave flow-ers) become embedded in the con¬ nective tissue at the base of most lesions (Figs. 28, 31). These materials have offered an excellent opportunity to study multinucleate foreign-body giant cells within the lesions. These large masses of cells have been studied with general histology and the SEM (Fig. 31). With azure-A and eosin-B, the cytoplasm around foreign plant material stains bright pink, whereas the basal cytoplasm is somewhat basophilic. Careful comparison of histological preparations with critical-point-dried SEM materials from only a few micrometers ('^ 10) aw'ay, has enabled identification of this cytoplasmic difference w ith the latter instrument (Fig.^ 31). By comparison of SEM photographs with adjacent histological ma¬ terials, it is possible to determine the three-dimensional appearance of these cells. It is interesting that at the apical .surface (adjacent to foreign material) the cell membrane conforms to the shape of the plant material and that the surface appears smooth and unbroken (at 5000 to 10,000 X ). It also is notew'orthy that foreign-body giant cells can be found in association with sites of alveolar and palatal bone resorption (Fig. 30) in a way similar to that reported by Irving and Handelman (1963). Resorption of palatal and alveolar bone might also result from localized circulatory disturbances. Increase in amount BJOLOGY OF THE PHYLLOSTOMATIDAE 173 Fuj. 30.—Lff/: Scanning elect ran micrograph of necrotic cell remnants, lymphocytes (L), and legs (LG) of a mite within an oral lesion. 1034 X. Upper right: histological view of a typical lesion. Abbreviations are; G, gnathostome; L, lymphocytes; M, body of mite; NT, necrotic tissue. Hematoxylin and eosin-Y, t67X. Loner right: A muliinucleated foreign- body giant cell (FBGC) adjacent to an area of palatal bone resorption. Abbreviations are: BV, blood vessel; MB, maxillary (palate) bone; N, nuclei; V, cytoplasmic vesicles character¬ istic of these cells. Hematoxylin and eosin-Y. 1900 X . 174 SPECiAl. EU BMC at IONS MUSEUM 7'EXAS TECH UNIVERSITY Fk:. 31.—XTomparalive histological (top) and scanning electron microscope views (bottom) of foreign plant material (F) and a muUinucleated foreign-body giant cell (GC). These two views represent sections through the same cell separated by approximately 10 micrometers. Abbreviations and symbols are: BV, blood vessel; FB> fibroblasts; L. lymphocytes; N, nucleus; arrow, apical cytoplasm of giant cell; arrow 1, conformation of apical cell surface to Surface of plant material. Hematoxylin and eosin-Y; lop. 1600 X ; bottom, 2000 X . of blood transported to the site of pcricxionlal infection alters the level and quality of transudate supplied to bone, a condition that can lead to resorption (Reich- born-Kjennerud, 1963). In histological preparations, it can be seen that venules at sites of infestations are considerably dilated. Response of the various types and stages of fibers (periodontal ligaments, reticular fibers, elastic and oxytalan fibers) also has been studied with both general histological techniques and the SEM. Except for remanents, or relatively BtOI.OGV OF I HE PHYLLOSTOMATIDAE 175 short lengths, fibers essentially are lacking from within the inflammatory sites. Collagenous fibers and reticular fibers (distinguishable with silver impregnation, after Lillie, 1965) generally are found at the periphery of lesions but are not as dense as elsew'here (in healthy areas). Elastic fibers have not been demonstrated in either healthy or infected areas of ora! submucosa in these bats. Oxytalan fi¬ bers, which have been studied extensively by Fullmer (1959, I960ri, 1960b, 1967) appear to be of special interest. First, it has not been possible to demon¬ strate these fibers in the periodontium of Lcptonycterh or other species of bats (see section on dental microanatomy). At the same time, however, in materials stained with aldehyde fuchsia following oxidation with peracetic acid, a dense, dark purple layer of oxytalan fibers sometimes can be found around the border of an inflammatory lesion. A possible role of these fibers in various types of fibrous pathology has been studied previously by others (see Fullmer, I960rt), Their relationship to inflammatory responses in the oral submucosa in bats with oral mites is as yet uncertain. A ttriliou at\d Eroskm Aside from dental caries and periodontal disease, there are at least three other causes of destruction of dental hard tissue (Sognnaes, 1963): abrasion; resorption from external causes; and erosion caused by both exogenous and endogenous chemical agents, mechanical factors, and idiopathic conditions. Abrasion is but one form of bimtxlal attrition (following Every and Kiihne, 1971; MacIntyre, 1966) and generally can not be regarded as pathological although specimens of glossophagines sometimes are seen in which the degree of abrasion suggests ab¬ normal occlusion. It has been pointed out that particular teeth and specific coronal surfaces are especially prone to abrasion in this subfamily (Phillips, 1971). For example, in Lepumyaeris, the metaconid of the first lower molar commonly is worn more greatly than are elements of other teeth w'ithin an In¬ dividual. Resorption of secondary teeth is not common in the phyllostomatids, wath ex¬ ception of teeth found adjacent to lesions caused by oral mites in Leptonycteris^ MonophylUis, and Anotim. The mechanisms of this resorption are unknown but can be regarded as an aspect of periodontal disease rather than some agent ex¬ ternal to the teeth and oral cavity. In contrast to pathological abrasion and resorption due to external causes, dental erosion (Figs, 32, 33) apparently is common in many species of phyl- lostomatids. To date, this pathology has been surveyed only in the Glos- sophaginae (Phillips, 1971), but a cursory examination of museum specimens, particularly of frugivorous genera such as Antheus, suggests that erosion is an important and common form of dental pathology among phyllostomatids. In the glossophagines, erosion is characterized by a generalized area of dis¬ solution (Figs. 32, 33) often beginning at sites of wear facets (Phillips, 1971). In these species, w-htch are characterized by small teeth, dental erosion frequently results in fracturing and loss of crowns. It is possible with the scanning electron microscope to study closely the characteristics of initial lesions representing 176 SPECIAL PL'liLlCA'MONS MUSEUM TEXAS TECH UNlVEkSllY 1 mm Fig. 32.—Dental erosion in the last lower premolar in a specimen of Anotira geoffroyL Abbreviations are: a. occlusal surface of remaining portion of tooth; b, exposed dentin; c, healed alveolar socket; d, normal dentary bone at dental cervix. From Phillips (1971). Fig, 33.—Scanning electron micrographs of dental erosion (arrow) on a lower molar of Olossop/iaga .utriiina. Abbreviations are: E, enamel surface: G, globular dental plaque. 350 X (left) and 700 X (right). dental erosion (Fig. 33). Such lesions differ from those of dental caries in that the former are not found in basins. In the example illustrated, which is a lower molar from an adult specimen of GI(KSS()p/ui},nj soricina, the outer enamel cuticle BIOLOGY OF THE PHYlXOSTOMATlDAE !77 is absent and the inner portion of the enamel layer has a rough, mgose appearance (Fig. 33). Tongues and Associated Musculature The gross anatomy of phyllostomatid tongues, except for the structure and distribution of papillae and for the tongues of glossophagines and desmodontines, is similar to that for other groups of microchiropierans. The longues of Arn'heus, CaroHki, and Pliyiiosfomus, as described or figured by several authors, are broad and usually rounded at the apex (Park and Hall, 1951; Lautenschlager, 1935). In many species, the distal portion of the tongue is progressively dorsoventrally compressed. Robin (1881) suggested that the tongue of Macroius usually is some¬ what more pointed and tapered than that of the others mentioned above and also noted that the tongue of Ariiihcus Cperspiciliains*') is substantially different in relative length and topography from that of megachiropterans despite the general similarity in food habits. Dorsal grooves generally seem to be lacking in tongues of fruit-eating phyl- lostomatids; however, a shallow depression has been described on the posterior surface in Carol I ia and Macroius {Park and Hall, 1951). Gross features of glossophagine tongues differ from those of other groups of phyllostomatids. The tongues of these nectar feeders are highly specialized for withdrawing liquid from elongate flower corollas. They are narrow', extremely elongate, highly extensible, and often have a pointed apex (Fig, 34). Winkelmann (1971) examined numerous adaptations for nectar feeding in glossophagines and identified two types of longues, each of which has sub¬ stantially different specializations for drinking nectar. The first group, which consists of Lonchophylla n^husia, L, moniax, Lkmycteris spurreiii, and Fkiialina ^enovensiunu possesses deep laterally placed longitudinal grooves, one on each side. These grooves probably enlarge to fill w-ith nectar during extension. The ventral surfaces of the grooves in Lonchophylla rohusta have numerous elongate papillae that are directed laterally. Other species in this group lack papillae in the grooves. The second group consists of Glossophaga soricitui^ Anoura geof- froyi, A. caudifer, MouophyliHS rednuinl, Lepionycteris nivalis, L, sanhornk Choeronycteris mexkana, and Musonycteris hatrisoni. The tongues of these lack lateral grooves. A dorsal trough widens posteriorly in the tongues of Li^pionycier'iS (Greenbaum and Phillips, 1974), Choeronycteris mexlcana, and Glossophaga soricina {Park and Halt, 1951). Most investigations of phyllostomatid tongues have focused upon the nu¬ merous, and sometimes highly specialized and elaborate papillae, which principally adorn the dorsal or, in some cases, the lateral surfaces (Fig. 34). Many species of the subfamily Glossophaginae are highly unusual with respect to papillae, and these specializations wdll be considered separately. Two large vallate papillae (Figs. 34, 35) are found on the posterodorsal surface of the tongue in most microchiropterans (Grasse, 1955), and thus their absence in Choeronyaeris and Desmodus, among phyllostomatids, is noteworthy. Papillae are otherwise highly variable in frequency, distribution, and shape among the I7« SPECIAL PUBI,1C:A [ K)NS MUSEUM TEXAS TFXJI UNIVERSITY Eui. 34.—Diagriifnmatic overview of tongues from Lt‘f>to/iyi reris .vttihotni ( A, C> and L. niviilis fB, D). Abbreviations are: mt. median trough; bp, hairlike papillae; hnp, horny papillae: bp, bifid papillae; hsp, hooklike singly pointed papillae; fsp, fleshy singly pointed papillae: pg. posterior groove; fnp, fungiform papillae; mvp, median vallate papillae: Ivp, lateral vallate papillae. From Greenbaum and Phillips (1974). few phyllostomatids that have been examined in detail. One species of Anihetis, as examined by Lautenschlager (1935) and Park and Hall (1951), has many large, broad, Hattened papillae within the posterior region that are unusual in being directed anteriorly. In Caroiiiit perspicilkaa, however, the large, posterior papillae are elongate and assume a basketlike appearance. Those of equivalent location in MacnHns califorfiicus and Dc.sniodas n/tundas are soft and flattened, but often terminate in a hairlike apex. In Piiyllosrfmins, Robin (1881) noted an abundance of conical papillae, which are largest in the midregion of the tongue. Fungiform papillae are found relatively infrequently on the tongues of most phyllostomatids. Park and Hall (1951) described the presence of a few' scattered ones in Carollhi (also described as rare by Robin), Desnunius, and Lepronycretis but reported their absence in Mmrotus, Choeronyaeris, Glossophaga, and ArilheiiS. Intraspecific variability in distribution of these papillae is possible judging from the fact that Fishman (1963) described the innervation of fungi- foriq kinds in his study of gustatory response in Ar it hens jamakensis. Green¬ baum and Phillips (1974) noted differences between two species of Leptonycteris with regard to fungifomu papillae, Leptonycteris nivalis has a few' large ones on each side of the posterior groove of the longue, w'hereas in L. sanhorni they are more abundant, but smaller. Variations of papillae that are most adaptive for particular foods arc found on the tongues of glossophagines. The distal portion of the Lonchopfiylla-type tongue of five of the glossophagines described by Winkelmann (1971) is covered w ith approximately two dozen papillae that decrease in size posteriorly. Nfost of the remaining surface in this species group is covered with small, nodulelike filt- BIOLOGY OF THE PHYLLOSTOMATiDAE 179 Fkl 35.—Scanning electron micrograph of a vallate papilla fVP) and singly pointed papilla (P) in a specimen of Leptonycteris su/ihorni 870X . form papinae. There are four or five bifid horny papillae (Fig. 36) vvtthin the midline about one-third of the way back from the lip of the tongue. Along the dorsolateral edge on each side of the anterior third of the Glosso- phaga-typQ tongue (after Winklemann), are rows of long hairlike vertical papillae. The number and sixe of these are highly variable among the species of Glos- sopfuiga, A tioura^ MotiophyUus, Leptonycteris, Choeronycteris, and Musonycteris, and those of G. sorcina have been described by several v/orkers as especially coarse. These elongate papillae probably load nectar by capillary action follow¬ ing extension of the tongue. Horny papillae, similar in shape and kx:ation to those in the LifnchophyilaAype tongue, are present in this second group of species. The remainder of the tongue is covered with smaller hairlike papillae. Of related interest are the studies of differences in tongue structure in relation to the presence of an unusual periodontal disease (see section on pathology) in only one of two extremely similar and sympatric species of Leptonycteris {PhW- lips, 1971; Greenbaum and Phillips, 1974). The macronyssid mite Radjordieiki orkoki produces lesions in the palate and alveolar bone only in L, mvalis. Dorsal and lateral filiform papillae on the posterior portion of the tongue in L. sanborni 180 SPt'CiAl. PUHI.JCAI IONS’ MUSEUM TEXAS TECH UNIVERSK Y Fig. 36.—Scanning electron micrograph of a pair of bifid papillae on ihe tongue of a specimen of Lfpitinycteris snnht>r/ii. Scalelike keratinized epithelial cells can be seen sloughing off from the surface (arrows). I020X. (Fig. 37) might prevent altachmcni of the mites in this species by way of a brush' ing action against the oral and gingival mucosa. The tongue musculature, both extrinsic and intrinsic, has been studied in few leaf-nosed bats. Because some glossophagincs probably feed almost exclusively with the tongue, they are of particular interest, and their tongue musculature has been the most thoroughly studied among phyllostomatids. The tongue muscula¬ ture of glossophagines and other phyllostomatids is qualitatively like that of other mammals. However, While (1954) described some noteworthy specializa¬ tions of origins and insertions of muscles in glossophagines that are highly con- iribulive to the great extensibility and mobility of their tongues. While (1954) observed that the genioglossus is highly developed into a leaflike form in Lep- louyaeris^ Athyura, Ltmchophylhi, and Choeronycteris and contributes to the great protrusibility of the tongue. The sternohyoideus has its origin on the ster¬ num, but inserts into the base of the longue to permit improved retractility. Also, the point of origin of this muscle shifts posteriorly from the manubrium of the sternum to the xiphoid process thereby increasing the length and force of tongue retractility, Thi.s, along whth particularly deep insertion of the sterno¬ hyoideus into the tongue, and insertion of the siylohyoideus on the lateral edges results in improved efficiency of manipulation. Intrinsic muscle fibers form a complex system of longitudinal, vertical, and transverse bundles that serves to reduce depth and diameter of the tongue, resulting in its elongation (Fig, 38). Fit;. 37,—Scanning etcctron micrograph of flai, fleshy papillae (top; arrow) in Lcpionycti’ris nivalis in comparison to the well-keratinized, hooklike papillae in L. xan- /jorn/ (bottom; arrows). Both views I 40 X. BIOLOGY OF THE PHYLLOSTOMATIDAE 182 SPECIAl- PUBIJCAL IONS MUSEUM TEXAS TECH UNIVERSITY Winkelmann (1971) developed a model for longue movement in gJos- sophagines. The great extensibility of the longue results from a combination of elongation through conlraction of intrinsic muscles and protraction of the base of the tongue by the action of the extrinsic muscles. Protraction is effected by relaxation of the steniohyoids (retractors), thus providing increased reach to the base of Howers. The mechanism of unloading of the tongue is unknown in any of the glossophagines but VVinkelmann (1971) assumed that shortening of the longue proceeds from proximal to distal to prevent unloading until the tongue is in the mouth. The tongues of vampire bats are structurally and functionally specialized to permit consumption of large quantities of blood. The tongue of the common vampire, Dcsmotliis nuundus, is rounded and has a deep dorsal fissure within the posterior half and a groove along the vent r> 1 ateral border on each side. The change in orientation of the blood grooves from front to back and the anatomy of associated papillae have been described by Glass (1970). During feeding, the tongue has been observed to move very rapidly in and out of the mouth; blood is visible only on the dorsal surface at the rear. Consequently, blood is thought to move along the ventrolateral grooves by peristalsis and emerge dorsolaterally on the tongue at the back of the mouth. A complex system of abundant motor endplates has been observed by Field and Holbrook (1969) in the tongues of Arnheiis limrafus and Des/nodits rontn- (lus, as well as in two species of vespertilionids. fhree to four individual fibers combine to form each endplate, each of which then splits into seven or eight terminal branches. No features of motor of sensory fibers or associated structures in the tongue were cited as unique to DesnKn/us ot Anihens. Suihers (1970) reviewed what is known about taste in bats, including data on experimental gustatory responses in Ai fihais ju/naicensis {Fishman, 1963) and Deswodu.s roiiitkliis. Application of dilute acid to the dorsal surface of the tongue yields greater responses in D. njiufidus than it does in several other bats for which data arc available, including, and especially, A. jafnaiceusis. In contrast, A. Janwicensis apparently is relatively sensitive to NaCl concentrations greater than that required for minimum responses and is unusual among the few bats that have been examined because its response to salt is greater than that for the other three principle taste categories. Salivary Gi.ands Salivary glands are an integral and important component of the oral biology in all mammals. Although the general impression often is that these glands are Ft(p. 38.—Diagrammaiic presemaiion of tongue masculature in long-nosed bais {Li'pioiiycieri:^). Toff, anterior musculature in the region of the horny papillae. Bodom: peripheral musculature in detail (box labeled X in top drawing). Abbreviations not ex¬ plained in Fig. 34 are: Im, longitudinal muscle; hm, horizontal muscle; vm, vertical muscle; Iv, lingual vein; la. lingual artery; il, lingual ligament; de, dorsal epithelium; Ip, lamina propria; cc, central muscle core. From Greenbaum and Phillips (1974). HIOl.OGY OF- THE PHYLLOSTOMATIDAH IK3 CC 184 SF^EClAl- PUBI.ICATIONS MUSEUM TEXAS TECH UNIVERSFTY only involved with the initial stages ot' digestive processes, in fact they have a w ide variety of functions that include provision of a protective coating for the teeth; production of antibacterial agents, toxic substances, and anticoagulants; maintenance of salt and water balance; and poxiuction of one or more hormones (Klinkhamer, 1968; Pearson, 1950; Rutberg, 1961; DiSanto, I960; Ito, 1960). The major salivary glands, particularly the submandibular, have been studied in considerable detail in man and in certain laboratory species (Leeson and Jacoby, 1959; Luzzato ei «/., 1968; Parks, 1962; Scott and Pease, 1959; Tamarin and Sreebny, 1965; Tandler, 1962, 1963, 1965), Interest in this particular gland partly is due to its tendency toward noteworthy specialization in many species. For example, it is the submandibular of the short-tailed shrew {Biarina brevicaiuia that produces a toxin {Pearson, 1950); the submandivular of the house mouse {Mtis }?iusai!ns) and Nonvay rat {Rauus norvef*icus) that displays sexual dimorphism (Junqueira et aL, 1951; Lacassagne, 1940; Caramia, 1966); the submandibular of the hamster ( McsiKTicetus cuitatux) that produces a sex¬ ually dimorphic mucin (Shackleford and Klapper, 1962); and the submandibular of the common vampire hat (DesnuHlus roimidifs) that produces an anticoagulant (DiSanto, I960). Investigational techniques employed in studies of salivary glands have pro¬ gressed from general gross anatomical and histological procedures to experi¬ mental cytological and physiological methods (Caldwell and Shackleford, 1967; Amsterdam et ai, 1969; Dressier, 1974; Scott and Pease, 1964; Castle et ai, 1972, 1975), The abundant literature on mammalian salivary glands shows w'ide variation in quantity and quality of information about all aspects of these glands in w'ild and domestic species. Indeed, most w'ild species are almost unknowm. The most notable exceptions are provided by a series of excellent papers by Shackle¬ ford and his co-workers on such species as the opossum (Wilborn and Shackle¬ ford, 1969), kangaroo rat and antelope ground squirrel (Shackleford and Schneyer, 1964), nine-banded armadillo (Shackleford, 1963), and squirrel monkey (CowJey and Shackleford, 1970u, 1 97Oh). The general paucity of studies on wild mammals is somewhat surprising, especially in view of the striking struc¬ tural and histochemical diversity apparent in salivary glands of mammals {see Shackleford and Wilborn, 1968, for a review). The variety of functions attributed to salivary glands notw'ithstanding, it is reasonable that hats {particularly the phyllostomatids) could be an extremely im¬ portant model for comparative studies of these structures. The impressive array of feeding habits that mark the evolutionary history of the Phyllostomatidae pro¬ vides an excellent opportunity for investigations of cellular evolution and, in the long run, could prove valuable to biosystematic interpretations. Systematic and evolutionary implications of differences and similarities in salivary glands gen¬ erally have not been presented because the species previously studied often w'ere only distantly related (Andrew, 1964) or, sometimes, because basic taxonomic misconceptions caused confusion and considerable analytic difficulty for the in¬ vestigators. For example, rabbits {Orycfolagiis) were regarded as rodents as recently as I960 by Ouintarelli and Chauncey (1960) who compared meta- BIOLOGY OF THE FHVE.LOSTOMATIDAE chromatic tinctorial characteristics of salivary glands of these animals to those of albinistic Norway rats {Rairus norvegicus) and house mice {Mus fnnsaiiits). Comparisons of even highly detailed features of salivary glands of a few spe¬ cies of rodents w'iih data from lagomorphs, man, dogs, and cats have been in¬ adequate for development of an understanding of the evolutionary process inso¬ far as these organs are concerned. The salivary glands of phyllostomatid bats, aside from two important excep¬ tions, are poorly known. Robin (1881) apparently was the first, and for many years the only w^orker, to describe salivary glands of various species of megachi- ropteran and microchiropteran bats. His descriptions included 41 recognized species. Glossitphaiiia .soricitia^ one of the phyllostomatid species studied by us, w'as included in his report. More recently, Dalquest and Werner (1951) reported on what they incorrectly termed “interscapular glandular adipose tissue” in Ariibeus Jafnaicensis, A series of other studies (W'erner er ai, 1950; VVemer and Dalquest, 1952; Dalquest et aL, 1952; Dalquest and Werner, 1951, 1954) also dealt w ith certain general histological aspects of salivary glands in a variety of microchi ropteran bats, Grasse (1955) briefly described the salivary glands of bats, with particular reference to production of an anticoagulant by the salivary glands of the common vampire bat, Dcsmodus rotumhis. The best, and most recent, studies of salivary glands of phyllostomatids are those by Wimsatt (1955, 1956) and DiSanto (1960). It w'as W'imsatt (1955) who pointed out that Dalquest and W'erner (1951) had mistakenly identified a portion of the parotid gland of Arriheus as browm fat; later, he (Wimsatt, 1956) provided an excellent, complete histological and histochemical analysis of the salivary glands of this frugivorous species. Likewise, DiSanto (1960) has undertaken a complete study of the anatomy, histology, and histochemistry of the salivary glands of DesfmtJus and provided a partial comparison with Aniheus. In Aniheusjamaicensis, the parotid secretory cells were judged by Wimsatt (1956) to be serous even though they were found to be morphologically more like mucous cells, at least w'hen viewed with the light microscope. These secretory cells are negative to mucopolysaccharide techniques such as PAS, Alcian blue, and mucicarmine, apparently do not pRxluce amylase, and are not serozymogenic (Wimsatt, 1956). Both the intercalated duct cells and striated cells of the striated duct are PAS positive, suggesting a secretory role in this species (W'imsall, 1956). In Desntodus^ the parotid secretory cells w'cre regarded as seromuccoid based on their histochemical responses, even though they appear to he serous cells from a morphological standpoint (DiSanto, 1960), In this species, the secretory cells showed prominent reactivity to five of six carbohydrate reactions and also revealed beta and gamma metachromasia to toludine blue (DiSanto, 1960). The secretory cells in parotids of vampire bats lack both acid and alkaline phosphatase activity, even though both enzymes ap¬ parently are present in the saliva (DiSanto, 1960). The fact that extracts of paro¬ tid glands from this species do not cause dissipation of blood clots suggests that they do not play a role in anticoagulation (DiSanto, 1960), Although the classical submandibular gland is characterized by mucous cells with a cap or demilune of serous cells (Sicher and Bhaskar, 1972), the opposite 186 SPECIAL PUHLICAMONS MUSEUM TEXAS lECH UNiVERSITY IS true in both A. Ju/naketisis und Deynuxius (Wimsatt, 1956; DiSanto, 1960). Additionally, it is the submandibular of Destntxlus that apparently pro¬ duces an anliclotling agent. This salivary component does not act as a true anti¬ coagulant because it does not prevent formation of fibrin; instead, it acts directly on formed fibrin by dissolving it (DiSanlo, 1960). The sublingual gland of Arfihcus j(unaicensis is niorphogically and, generally speaking, histochcmically similar to that of other mammals (Wimsatl, 1956). The sublingual gland of DcsnuHins, on the other hand, is unusual in that the secretory acini are comprosed of both mucous cells and meiachnimatic cells, which ex¬ hibit extensive mciachromasia through a broad pH range. The metachromatic cells might produce an unusual polysaccharide of low pH and could, according to histochemical data presented by DiSanto (I960), produce an anticoagulant such as heparin. At the same lime, however, extracts from the sublingual do not pre¬ vent coagulation of blotid. The lack of basic histological information about salivary glands in most phyl- lostomatids has led us to describe in following paragraphs the gross anatomy and general histology of the three major salivary glands of five additional phyl- loslomatid species. The following accounts, which are intended to serve as a basis for more detailed and sophisticated study, are preceded by brief, gencral- i/X'd overviews of both gross anatomy and cellular morophology. General Gross Anatomy Gross dissection and examination of the three major salivary glands (parotid, submandibular, sublingual) in Glosst^phaga soricina, Leptonyaeris nivalis, L. sanhorni, Anottra gi'offroyi, and Siarnira Indovici, revealed considerable dif¬ ferences in size and shape of the glands (Fig. 39). General findings about the gross anatomy of the major salivary glands are presented in the following para¬ graphs. The parotid gland varies in appearance, being either soft and loose or distinct and compact, and in size, ranging from extremely small to strikingly large (Fig. 39). It is found at the base of the auricle and is enclosed in a strong and tightly adherent capsule of connective tissue. Stenson’s duct can be traced from the anterior edge of the gland (having originated on the inferior surface) anteriorly, across the masseter muscle. A submandibular gland is found in all five species studied by us. Although it varies greatly in number of lobes, size, and appearance (Fig. 39), in all of the species it is encased in a capsule of dense connective tissue that appears to be continuous w'ith that of the parotid gland. The position of the submandibular varies somewhat, but generally the lobes are located in the mastoid region of the skull and are separated from the parotid by the external jugular. With exception of Anoara gcoffrtfyi, the submandibular possesses a single duct, to w'hich all of the lobes are joined. This duct typically passes anteriorly, under the digastric muscle, and along the lingual surface of the ascending mandibular ramus. In Aaonm the anterior lobe of the principle submaxillary has its own duct, which is described in detail in the account of this species. BIOLOCIY OF THE PHYLLOSTOMATIDAE 1H7 FjCi. 39.—Diagrammatic presentation of the gross anatomy of major salivary glands in A) Lt'pfofiycterix nivtiiis; B) Siurniru }miin’ici\ C) Anoura gcoffroyi; and D) Ariifu'ux jiimuicenxis. Abbreviations are; P. parotid; SM, submandibular; L, sublingual. The sublingual salivary gland is a delicate, sufl, loosely encapsulated mass of tissue located in a triangular-shaped depression bounded by the digastricus, sternohyoideus, and sternomastoideus and positioned directly ventral to the thyroid glands and larj'nx. The size and shape of the sublingual, as well as the size of the depression in which it rests and the depth at which it is located, varies greatly from species to species (Fig. 39). In all five species this gland has a single duct that originates on the dorsal surface of the gland and enters the oral cavity at the base of the tongue, from which point it travels anteriorly to the tip of the sub¬ lingual flap. General Microanatomy Diagrammatic representations of salivary glands of five species are presented in Figs. 40 and 41. The baste microanatomy is fairly consistent even though the cellular details and functions vary w idely from species to species. Terminal portions ,—The terminal portions of the salivary ducts are char¬ acterized by groups of secretory cells (acini) clustered around a narrow lumen SPECIAL PUHLJCA'nONS MUSEUM TEXAS TECH UNIVERSITY 1SS (Figs. 40, 41, 42). In phyllostomatids, the parotid secretory acini typically are comprised of a single type of secretory cell, which is either serous or seromucoid in morphology. The parotid acini usually are relatively small and somewhat round In shape and can be termed compound acinus. The typical phyllostomatid submandibular gland consists of elongate and often branched secretory acini comprised of at least two types of secretory cells (Fig. 41). In most species one type of secretory cell forms a cap, or demilune, at the distal end of the acinus. The submandibular secretory cells are a combination of serous, seromucoid, or mucousMype cells. The secretory acini of the sublingual gland are elongate (some- limes extremely so), branched, and comprised exclusively of mucous cells. Dramatic overviews of salivary gland microanatomy are provided by the scan¬ ning electron microscope (Figs. 42, 43). In Fig. 42 and 43, which show' the sub¬ lingual gland from a dwarf fruit-eating bat, Artiheiis phaeoiis, the branched, elongate secretory acini arc clearly visible. The exposed cytoplasm of the mucous cells is rough and trabecular in appearance, except at the basal margins where it is smooth (Figs. 42, 43). Certain other microanalomical features associated with the terminal portion of the duct system also are well illustrated by these SEM photographs. For example, in Fig. 42, one can easily distinguish a large capillary adjacent to an acinus; a major nerve bundle as well as smaller nerves that con¬ nect separate acini; and the basal lamina covering the secretary acini. In Fig. 43, the basal lamina can be compared to the connective tissue capsule, which is con¬ siderably thicker. The microanatomy of the acinar lumen also can be analyzed with the SEM. In Fig. 44, secretory granules about to be extruded into the lumen can be seen to cover the apical surfaces of the secretory cells. This view, which is from the .submandibular gland of Macnnus waierhousii, is characteristic of acinar lumina lined by secretory cells of the serous type. An interesting comparison can be made between SEM microanatomy and transmission electron microscopy. Although the TEM photograph shown here is from the submandibular gland of a speci¬ men of Phyll(mycteris, moderately electron dcn.se serous granules can be seen at the cell apex, adjacent to the acinar lumen. These granules are still covered by a membrane, which appears as a thin, dark line (Fig, 44). Ducts .-—rhe duct system leading away from the secretory acini can be divided into three segments; in order, they are intercalated ducts, striated ducts, and ex¬ cretory ducts. Certain microanalomical features allow for relatively easy recogni¬ tion of each type of duct. The intercalated ducts vary in length and degree of branching but typically have a narrow lumen and flattened appearance (Fig. 43). The transition between these duels and the so-called striated ducts is abrupt (Fig. 43); the latter are considerably thicker, although the lumen still is narrow' relative FiCi. 40,—Diagrammatic presentation of basic microanatomy of the parotid salivary gland in A) Glossophiiga soi k ina', B> Li’ptofiycicris sunhtn-tii: C) L. rtivaitA- D) Anitura j^eofffoyn and E) Siurnim imiovicL Fig. 41. -Diagramrriatic presentation of basic microanatomy of the submandibular salivary gland in the same species as shown in Fig. 40. BIOLOGY OF THE PHYLLOSTOMAT[DAE 1K9 190 SPECIAL J>UHIJCATI()NS MUSEUM lEXAS lECH UNIVERSITY Fig. 42.—Scanning electron micrograph of a secretory acinus (A), intercalated duct (ICD)* and striated (intralobular) duct (SD) in the sublingual salivary gland of Ariilu'ux phaeotix. Abbreviations are: C, capillary; L, acinar lumen; N, nerves. II70X. to the overall (outside) diamcicr. The luminal surfaces of these ducts vary con¬ siderably; they can be smtKHh, covered by microvilli, or even covered by micro¬ villi so long and densely packed that they resemble a brush border. An example of luminal microvillt is presented in Fig. 45, which is an SEM photograph from the submanibular of Macrotits. In this view, it is apparent that many of the microvilli are inter connected by narrow- bridges of cytoplasm. Such bridges have not been reported previously in salivary glands, and their significance here is unknowm. Excretory ducts, which carry saliva away from the lobes of the gland, are charac- BIOLOGY OF THE PHYLLOSTOMATiDAE 191 Fig. 43. —Scanning electron micrograph showing mucous secretory cells (MC) and extra- acinar space in the sublingual gland of Ardheus phtteofis. Abbreviations and symbols are: BL, basal lamina; C, connective tissue; EP, ergastoplasm; FB, fibroblast; arrows, fibrils (presumably collagenous). 1206X. terized by a wide lumen and narrow wall, the latter reflecting the low, almost cuboidal nature of the cells. Detailed Descriptions Details of gross anatomy and general histology of five phylloslomatid species are provided for the first time in the follow'ing paragraphs. 192 SPECIAL PUiil.lCATlONS MUSEUM TEXAS JECH UNIVERSITY Fig. 44.—Comparative transmission (top; and scanning (bottom: Mucrotus) electron micrographs of exocytosis of serous secretory product (SG) into the acinar lumen (L). Abbreviations and symbols are: GER, granular endoplasmic reticulum; MG, mucous secretory granule; arrows, direction of export into lumen. Both approximately 9730 X, soricitia Cross AnaUfmy Faroticl .—A moderate-sized gland extending from the lambdoidal region to the auricle (Fig. 39), A small lobe of the gland extends ventrally, over the niasseter and digasiricus muscles and lateral margin of the sublingual gland, Stenson’s duct BJOLOGV OF THE PHYLLOSTOMATIDAE 193 Fig, 45.—Scanning electron micrograph showing microvilli (MV) ai apical membranes (luminal surface; L, lumen) of cells in a striated duct in the submandibular salivary gland of Mticrotifs. Note the interconnections between microvilli (arrow's). Approximately 7600 X. originates on the inferior surface, near the anterior border, and follows the ex¬ ternal jugular along the base of the masseter to the anterior border, where it enters the connective tissue of the upper lip in the area of the canines. Stibmatuiihular .—A large gland consisting of four distinct lobes, each being ovoid and flat (Fig. 39). The two anterior lobes, which are slightly smaller than the remaining two, lie directly over the sublingual. The third and fourth lobes are located ventrally to the parotid and over the sternoniastoideus. The main ducts of the four lobes Join and pass over the digastricus before turning niediodorsad to meet the main duct of the sublingual near the inferior surface of the angle of the mandible. The two ducts enter the anterior portion of the oral cavity together, SiihlhiguaL —A relatively large, triangular gland consisting of three finely sub¬ divided lobes (Fig. 39). It is kKated at the angular process of the mandible and 194 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY FiCi, 46.—To/?: Secretory acini and striated ducts in the parotid of Glossophaga xorkina. Abbreviations are: A, acini; SD. striated duct. Masson's trichrome. 222 X. Bintom: Secre¬ tory acini and striated duct in the sublingual of Gloxxopha^ia surkina. Abbreviation not defined above is; ICD, intercalated duct. Hemato.\ylin and eosin-Y. 370 X. covered by the niylohyoideus muscle. The lateral edge of the gland extends from the digastricus to the mastoid region, passing under the stcrnomasioideus. Medial¬ ly, this gland extends to the ventral surface of the throat, under the lateral margin of rhe sternohyoideus. The main duct arises on the anterior margin of the sub¬ lingual and joins the main duct from the submandibular (see above). Hfsiology Parotid .—'(’he lobules of the parotid are densely packed with small, generally round, secretory acini (Figs. 40, 46). The secretory cells are seromucoid when stained with H&E. The small, round nucleus is positioned basally and hetcro- chromatic. The cytoplasm, which is most dense in the basal area around the nu- BIOLOGY OF THF PHYLLOSTOMAT[DAE ]95 cleus, is intensely basopiiilic {Fig. 46), The secretory cells appear to have large, irregularly shaped vacuoles between the nuclei and acinar tumina. These vacuoles appear to be empty {even at 1800X) in secretory cells stained with H&E, Mas¬ son’s trichrome, or PAS. The cytoplasm around the vacuoles, which forms a dis- tinct network throughout the apical portion of the cell, is intensely reactive to PAS, In some cells PAS-positive material can be seen adjacent to the vacuoles. The numerous lobules of the parotid gland have a moderately extensive duct system. The short, intercalated ducts are comprised of medium-sized cuboidal cells, each having a round, heterochromatic nucleus and faintly acidophilic cyto¬ plasm that nearly is obscured by the nucleus. PAS-positive granules are lacking in the intercalated ducts. The transition from intercalated to striated ducts is abrupt (Fig. 40). The cen¬ trally placed nuclei of striated cells are large and euchromatic {Fig. 46). When stained with Masson's irichrome, the striated portion of the cell is rust colored, whereas the apical cytoplasm around the nucleus is gray and that directly adjacent to the lumen of the duct is pale greenish gray. This region of the cytoplasm also is strongly PAS positive. The transition from the typical striated ducts to the inter¬ lobular ducts is gradual. It is characterized by a gradual increase in luminal di¬ ameter and a change in cell morphology. The nucleus becomes more centrally positioned and the cytoplasm adjacent to the lumen becomes only faintly acido¬ philic. This pale area is clearly visible in cells stained w ith Masson’s trichrome and is even more intensely reactive to the PAS technique than it is in the typical striated cells. This PAS-positive response suggests the presence of mucosubstance and pos¬ sibly indicates a secretory role for this portion of the duct system. Additionally, Mallory triple connective tissue slain reveals that the cells of this transitory region contain large, red granules. In comparison, only a few granules are found in the basal (striated) cytoplasm of the striated cells. Approximately 30 per cent of the cells comprising the striated duct system lack basal striations, are totally PAS negative, and have small, heterochromatic nuclei. These cells correspond to the “dark cells” that have been described by electron microscopy (see section on ultrastructure). The interlobular ducts are characterized by an increased luminal diameter and a decrease in cell volume. The cells of these ducts are nearly cuboidal and have a centrally placed, moderately heterochromatic nucleus. The cytoplasm is strongly acidophilic (with H&E), nonreactive to PAS, and uniformly brown w'ith Masson’s trichrome. When stained with Mallory triple, the cytoplasm is pale gray and con¬ tains numerous large, round, red granules. Submandibular .—The lobules of the submandibular gland are densely packed with compound tubular secretory acini of moderate size (Figs. 41, 47). Most of the acini are comprised of serous cells w'ith a mucous demilune, but some acini appear to be entirely mucous. These latter acini comprise approximately five to 10 per cent of the total acinar population. The relatively small, round nuclei of the serous cells are heterochromatic and basally restricted (Fig, 47), The small nuclei of the mucous cells are more euchromatic and basally restricted. 196 SPECIAL PUliLJCATlONS NfUSEt'M TEXAS TECH UNIVERSITY Fig. 47,— Top: Mixed serous (S) and mucous (M) secreiory acini and imercalaled duct UCD) in the submandibular of Gloi\wpluif*ti soridna. Hematoxylin and eosin-Y. 5K0X, BofUf/n: PAS-positive secretory granules (G) in serous cells of the submandibular of (ilossophafin .sorici/iii. 1972 X . tlKJLOGY OK I’HE PHYL(,OSTOMATIDAE 197 When stained with H&E, the cytoplasm of these latter cells is pale gray, almost achromatic, whereas that of the serous cells is dense and strongly basophilic. W'hen the PAS technique is employed, the serous cells are packed with large, strongly reactive, round granules (Fig. 47). The cytoplasm of the deniilunar cells, on the other hand, is characterized by irregularly shaped clear areas outlined by a fine, reticular network that is moderately PAS positive. This netw'ork stains dark purple when oxidized prior to staining wdth aldehydeTuchsin (this staining re¬ action is the only overlap with PAS in the submandibular). When Mallory^ triple is used, the serous cells arc pale pink and contain from one to four large, red gran¬ ules each. The mucous cells, on the other hand, are essentially nonreactive. With Masson’s irichrome, the cytoplasm of the serous cells is pale green and that of the demilunar cells is nonreaciive. AH four main lobes of this gland are histologically identical. The lobules of each lobe are characterized by an extensive system of ducts. The branched inter¬ calated ducts are comprised of medium-sized cuboidal cells with ovoid nuclei; the reduced cytoplasm is strongly acidophilic (Fig. 47), The system of intercalated ducts is extensive and the ducts are relatively long; in many small sections of lobules, only a few' striated but many intercalated ducts are found. Ihe transition from intercalated to striated ducts is abrupt (Fig. 40). The striated cells have large, round, apically displaced nuclei that are relatively eu- chromatic. The striated portion of the cytoplasm is intensely acidophilic, whereas that of the apical region usually is somewhat paler in cells stained with H&E. A narrow band of apical cytoplasm adjacent to the lumen of the duct is PAS posi¬ tive. Additionally, w'hen these cells are oxidized and stained with aldehyde- fuchsin, this area of the cytoplasm is pale purple. With Masson’s trichrome the cytoplasm is green in the striated region and pale green in the apical region adja¬ cent to the lumen. The striated cells are essentially nonreactive with Mallory triple connective tissue stains. The transition from striated to interlobular ducts is gradual; the nuclei become more centrally located and the basal striations are lost. The cells remain columnar but become slightly smaller; the luminal diameter in¬ creases slightly. As in the parotid, about 30 per cent of the cells comprising the striated ducts can be identified as dark cells with the light microscope. They have small, heterochromatic nuclei and are PAS negative. The cells of the interlobular ducts are nearly cuboidal; the nuclei are more heterochromatic than are the nuclei of the striated cells and are centrally located. The cytoplasm is strongly acidophilic, nonreactive to PAS, and pale green with Masson’s trichrome. The luminal diameter of the ducts is considerably greater than that of the striated ducts. SithlinguaL —The secretory acini of the sublingual are of the compound tubular type and are comprised of cells that have the classical mucous characteristics when stained with hematoxylin and cosin-Y. The small, round, heterochromatic nuclei are basally restricted (Fig. 46). In our formalin-fixed materials, these secretory cells were found to contain small, irregularly shaped, PAS positive gran¬ ules. W'ith Masson’s Irichrome, the secretory cells have dark brown cytoplasm, which is restricted to the area of the nucleus; the remainder of the cytoplasm of 19K SPECIAL PUEU.ICA f IONS MUSEUM TEXAS [ ECH UNIVERSITY these cells has small, dark granules and a faint reticular network. The sublingual gland is essentially nonreactive when oxidized w'ith peracetic acid and stained with aldchyde-fuchsin. All lobules of this gland are histologically identical. The duct system is only moderately developed. The branched intercalated ducts, which are not easily seen, are comprised of small, cuhoidal cells with heterochromatic, round, cen¬ trally placed nuclei (Fig. 46). The limited cytoplasm is only moderately acido¬ philic. Small PAS-posilive granules are found in the intercalated duct cells adjacent to the secretory acini. The transition between intercalated and striated ducts is abrupt. Within each lobule, the majority of the ducts can be classified as striated. The moderately heterochromatic nuclei are located adjacent to the lumen in the large, columnar, striated ceils; the basal striated portion of the cytoplasm is strongly acidophilic {Fig. 46). The striated cells are nonreactive to P.AS and are uniformly dark brown when stained with Masson’s trichrome. Approximately 30 per cent of the cells in these ducts are small, somewhat cuboid- al, lack basal striations, and have heterochromatic nuclei. The transition from the striated ducts to the interlobular (excretory) ducts is gradual and marked only by an increase in luminal diameter and slight decrease in cel! height. The nuclei of cells of the interlobular ducts are slightly smaller and more hetero¬ chromatic than those of the striated cells. The tinctorial properties, however, are essentially the same for these cells as for the striated cells, at least with the procedures employed by us. Additionally, the basement membrane of the inter¬ lobular ducts is consistently thicker and more fibrous than that of the striated ducts. LeptonycterLs sunhornt Gross Anafoffiy FarolieJ .—This is a mode rale-si zed gland that extends from the lambdoidal region of the skull around the auricle, and anteriorly until it overlies the posterior margin of the masseter (Fig. 39). Stenson's duct arises from the anterior border of the gland, passes across the masseter and anteriorly enters the connective tissue of the upper lip before entering the oral cavity near the upper canines. SuhnjwuJibii!(U \—This is a moderate-sized gland consisting of three to four compact lobes (Fig. 39). The posterior most lobe is the largest; it lies betw-een the sternomastoideus and the external jugular vein. The anterior lobes of this gland are flattened and ovoid and cover the sublingual gland. Wharton’s duct arises from the posterior most lobe and passes along the inferior surface of the aiilerior lobes receiving branches from each. From the anterior margin of the gland, the duct passes over the dorsal surface of the digaslricus and joins with the duct from the sublingual gland in the region of the angle of the mandible. These tw'o ducts run together along the tloor of the oral cavity and enter the mouth at a point directly posterior to the mandibular symphasis, under the sublingual flap. SnblhiguaL —In Lt’pfonycrerLs sanhorni, the sublingual is a moderate-sized gland that is triangular in outline (Fig. 39). It is located at the angular process of the mandible; the lateral edge lies against the digastricus, and the medial side BIOLOGY OF THE PHYLLOSTOMA‘1 IDAE 199 extends under the lateral margin of the sternohyoidcus. The anterior half of the gland is covered by the mylohyoideus. Posteriorly, the gland extends to the mastoid region of the skull and is covered there by the relatively wide sterno- mastoideus. The duct arises from the anterior margin of the gland and joins Wharton’s duct of the submandibular, as described above. Parolut —In this species, the parotid is a compound acinar gland; the secre¬ tory acini arc relatively small but numerous and, thus, densely packed (Fig. 40). The secretory cells best approximate the seromucoid type. They contain large, round, basally positioned nuclei that are moderately heterochromalic (Figs. 48, 49) . The nuclei are surrounded by dense, basophilic cytoplasm; the remainder of each cell is characterized by achromatic vacuoles of varying sizes. These are outlined by basophilic trabeculae that also are PAS positive (Fig. 48). The secretory cells are essentially nonreactive to Masson’s trichrome and Mallory triple connective tissue stains. Round, PAS-positive granules are found in many secretory acini at the junction w'ith the intercalated ducts. Although myoepithe¬ lial cells presumably are found in all salivary glands, they are especially obvious in the parotid of this species (Fig, 50). The lobules of the parotid are characterized by an extensive duct system com¬ prised of relatively complex cells. The intercalated ducts are long and highly branched. The distal cells (adjacent to the acini) are large, elongate or rectangu¬ lar, and have large, ovoid, moderately euchromatic nuclei. The cytoplasm ap¬ pears to be only slightly acidophilic with hematoxylin and eosin-Y; small, round PAS-positive granules often are found in the apical cytoplasm (Fig. 40). Ad¬ ditionally, some granules also are visible within the lumen. After oxidation with peracetic acid, these granules stain dark purple with aldehyde-fuchsin (Fig. 50) . The majority of cells in the intercalated ducts are flat, have heterochromatic nuclei, and are PAS negative. The transition from intercalated to intralobular ducts is abrupt (Fig. 40), The distal segment of the latter is comprised of large, columnar cells and has a narrow' lumen that generally contains PAS-positive material (Figs. 48, 49), The cytoplasm is only slightly acidophilic (with hematoxylin and eosin-Y); large, round, euchromatic nuclei are apically positioned and the basal one-half of the ceils have the characteristic striations. The striated cells are only weakly reactive to Masson’s trichrome; the cytoplasm has a pale rust-brown color. With Mal¬ lory triple the cytoplasm is pale pink, and with peracetic acid and aldehyde- fuchsin it is achromatic. The apical cytoplasm has an achromatic vacuole that is readily apparent when stained with hematoxylin and eosin-Y (Fig. 49). When examined with oil immersion optics, following staining w'ith hematoxylin, eosin-Y, and PAS, extremely fine trabecular PAS-posilive network wdthin the vacuole is observed. When stained wnth Masson’s trichrome, only the basal (striated) portion of the cytoplasm is reactive, being pale rust-brown. This same area of the cytoplasm is weakly acidophilic with hematoxylin and eosin-Y. Ceils comprising the proximal portion of the duct system lack the apical vesicle; 200 SOECJAt. PUBLJCAIIONS MUSEUM TEXAS I'ECH UNIVERSITY Fig. 48,—Scromucoid secretory acini and striated ducts in the parotid of Lepto/iycteris xttnhornL Abbreviations are: A, acinus: cap, capillary; dc, dark cell; SD, striated duct; T, trabeculae of cytoplasm. Hematoxylin and eosin-Y, Top, 610 X ; bottom, 2140X. BIOJ.OGY OF THE PHYIXOSTOMATIDAE 201 Fifi. 49.^— Top: Distal segment of striated duct in parotid of Lepionycteria Minborui Bottom: Proximal segment of the same duct as shown above. Abbreviations arc: BC, basal cell; DC, dark cell; E, ergastoplasm; L. lumen; N, nucleus; V, apical vesicle. Hematoxylin and eosin-Y. Both are 1720X . their nuclei are somewhat more apical in position (Fig. 49). Approximately 30 per cent of the cells comprising the striated ducts can be identified as dark cells (Figs. 48, 49). They are somewhat smaller than the striated cells, have heterochromatic nuclei, and are PAS negative. In the gradual transition to the more typical excretory ducts, the cells become less and less columnar (until they nearly are cuboidal) and consequently the luminal diameter increases considerably. The basal striations are lost, as is the apical achromatic vacuole, and the nuclei become more nearly ovoid. The stain¬ ing reactions of the cells generally are the same as those of the striated cells, at least with the procedures used in this study. Sitbmimdibutar .—The submandibular is a mixed compound acinar gland; the secretory acini are complex, numerous, and densely packed (Fig, 41). Two types of secretory cells are found; one, a serous type, is uncommon, representing 202 SPEC[Al, f’UB[,lC:AT10NS MUSEUM TEXAS TECH UNIVERSITY Fig. 50.— Top: Reactive secretory granules (G ICD) in the distal portion of the inter¬ calated duct in a parotid of Lt'pUinycicrh rnnhorni. Peracetic acid and aldehyde fochsin. 88HX, Lower left: Mixed secretory acini in the submandibular of L. sarthornL Abbreviations are: M, mucous ceils; S, serous cells; MC. myoepithelial cell nucleus. Hematoxylin and eosin-Y. n84X Loner right: A myoepithelial cell (MC) covering the surface of a secretory acinus in L. sanhoifii. Hematoxylin and eosin-V. 1184X, only about 10 per cent of the total number of secretory cells and is lacking from most secretory acini. Generally, these celts are clearly columnar and have targe, heterochromatic nuclei that arc basally restricted. These cells can best be de¬ scribed as serous when stained with hematoxylin and eosin-Y or azure-A and eosin-B (Fig. 50). The cytoplasm appears to be uniformly and densely baso- BIOLOGY OF THE PHYLLOSTOMATIDAE 203 phi lie with hematoxylin and eosin-Y, but w'ith azurc-A and eosin-B, the basal region is more distinctly basophilic than the apical cytoplasm. The latter con¬ tains numerous PAS-positive granules that are nonreactive to aldehyde-fuchsin after oxidation wdth peracetic acid. Additionally, the cytoplasm of these secretory cells stains rust-browm with Masson’s trichromc and is nonreactive with Mallory triple. The remainder (about 90 per cent) of the secretory cells have round, basal- ly restricted nuclei surrounded by a small amount of basophilic cytoplasm (hematoxyiiii and eosin-Y and azure-A and eosin-B). Most of the cytoplasm contains large vacuoles outlined by basophilic trabeculae and is thus similar in appearance to the almost achromatic mucous cells (Fig. 50). The trabeculae are PAS positive and also stain dark purple with aldehyde-fuchsin following oxidation with peracetic acid. The mucous cells are nonreactive to Masson’s trichrome and Mallory triple. Submandibular lobules are characterized by an extensive system of ducts. Generally, however, the cells of each portion of the duct system are relatively simple. The intercalated ducts are short, branched, and comprised of small, cuboidal cells with round, heterochromatic nuclei. The limited cytoplasm is w'eakly basophilic with hematoxylin and eosin-Y and lacks PAS-positive gran¬ ules. Peracetic acid aldehyde-fuchsin, Masson’s trichrome, and Mallory triple also are nonreactive. The intercalated ducts open abruptly into the striated ducts, w'hich are com¬ prised of large, columnar and moderately acidophilic (with hematoxylin and eosin-Y) cells. These cells have large, apical, euchromatic nuclei, and basal striations. The striated cells are nonreactive to PAS, with exception of a narrow band of apical cytoplasm. The cytoplasm of the striated cells is rust-browm with Masson’s trichrome and pale red with Mallory triple. After oxidation with per¬ acetic acid, small, round granules that stain dark purple w ith aldehyde-fuschsin are found in the apical portion of many striated cells. Approximately 20 per cent of the cells comprising this portion of the duct system can be classified as dark cells. They have small, heterochromatic nuclei and are positioned adjacent to the lumen. The transition between striated and excretory ducts is gradual, being characterized only by a decrease in cell height and increase in luminal diameter. The excretory duct cells are cuboidal and have more nearly ovoid nuclei than are present in the striated cells. The general staining reactions of the cells of the excretory ducts arc the same as those of the striated cells, at least w ith the techniques used by us. Stfblittgual .—The secretory acini of the sublingual are compound tubular. They are relatively large, branched, and loosely packed and are comprised com- pletely of mucous cells (Figs, 51, 52). The secretory cells have small, round or sometimes Battened nuclei that are heterochromatic and restricted to the basal piirtion of the cells (Fig. 51). The nucleus generally is surrounded by a small amount of basophilic cytoplasm (Fig. 52), w'hercas the remainder of the cell is pate, almost clear. Examination with oil immersion (1500X) reveals a dense but fine trabecular network (hematoxylin and eosin-Y) that probably represents the restricted cytoplasm between secretory vacuoles (Fig. 52), With 204 SPECIAL PUBLICATfONS MUSEUM TEXAS TECH UNIVERSITY Fig. 51.— 7\)p: General histology of mucous sublingual gland of Leptonyc/eris sanhifniL Boifom: Striated ducts and well-defined lumina (arrows) in secretory acini in the same gland as shown above. Abbreviations are: A. acini; cap. capillary; L, lumen of striated duct; dc, dark cell; pc. pale striated cell; SD, striated duct. Hematoxylin and eosin-Y. Top, 170X, bottom. 4)41 X . PAS, the basal cytoplasm of these cells is strongly reactive, whereas the re¬ mainder has a moderately reactive granular appearance and strongly reactive trabeculae. Aldehyde-fuchsin following oxidation wdih peracetic acid produces essentially the same staining reaction. The secretory cells are nonreactive to Masson's trichrome and Mallory triple connective tissue stains. The sublingual gland has a moderately extensive, but relatively simple, duct system. The inter¬ calated ducts are long, branched, and comprised of elongate or rectangular cells that have large, ovoid, nearly euchromatic nuclei (Fig. 52). The limited cytoplasm of these cells is only slightly acidophilic (hematoxylin and eosin-Y); the apical cytoplasm varies from slightly to strongly PAS reactive. The cells are nonreactive to aldehyde fuchsin following oxidation wdth peracetic acid, as w'ell as Masson’s trichrome and Mailory triple connective tissue stains. BIOLOGY OF THE PHYLLOSTOMATIDAE 205 Fki, 52.—Composite histological views of a sublingual gland in Leptortyctcrix sanhornL The large, euchromatic nuclei (Ni) of intercalated duct cells can be compared with the smaller, heterochromatic nuclei {N 2 ) of the mucous cells. The basal cytoplasm of the mucous cells is characterized by a distinct ergastoplasm (EP), whereas the remainder of the cytoplasm is dense (arrows) but greatly restricted by secretory granules and is, therefore, trabecular in appearance. Hematoxylin and eosin-Y, Both figures are 1184 X . The intercalated ducts open abruptly into striated ducts comprised mainly of large, relatively narrow, columnar cells with round, euchromatic, apically displaced nuclei (Fig. 51). The cytoplasm is only weakly acidophilic w'ith hema¬ toxylin and eosin-Y and non reactive w'ith the PAS technique. These cells stain pale green with Masson's trichrome and pale pink with Mallory triple. They are nonreactive to peracetic acid aldehyde-fuchsin. The transition to interlobular ducts is gradual and characterized by a decrease in cell height and increase in luminal diameter. Although the nuclei become more nearly ovoid, the basal striations are not lost. Indeed, the striations are found even in the cuboidal cells comprising the interlobular ducts. 206 SPECIAL PUBL[CATIONS MUSEUM TEXAS TECH UNIVERSETY Leptonycteris niviilis Gross A natomy Paroiid .—This is a nioderate-si^ed, elongate gland extending from the angle of the jaw to the posterior side of the ear. The parotid is separated from the main mass of the submandibular by the external jugular. Stenson’s duct arises from the inferior surface of the anterior end of the parotid and runs across the masseter into the connective tissue of the upper lip. It enters the oral cavity in the region of the upper canines. Suhfnmuiihular .—This is a large gland, consisting of a series of six or seven main lobes, each of which is extensively subdivided. All but one of these ovoid, tlat lobes are arranged, one behind the other, along the external jugular. The most posterior lobe is located dorsally to the external jugular, directly posterior to the parotid. The submandibular extends from the base of the neck anteriorly to the angle of the jaw; it covers portions of the sternomastoideus muscle and the sublingual gland. The jnain ducts of the various lobes join and leave the submandibular gland on the inferior surface of the most anterior lobe. Wharton's duct passes dorsal to the digastricus, in the region of the angle of the mandible, and is joined by the main duct from the sublingual. The two ducts together enter the anterior portion of the oral cavity. Sublingual .—This gland is large, triangular, and unilobular but finely sub¬ divided. It is located at the angle of the jaw, bordered laterally by the digastricus. Medially, the gland extends to the lateral margin of the sternohyoideus muscle. The posterolateral edge of the sublingual extends to the ma.stoid region of the skull, where it is covered by the sternomastoideus muscle. The main duct from this gland joins with that of the submandibular (see above). Histol(}gy Parotid .—The parotid is a compound acinar gland that is characterized by small, round secretory acini (Fig. 40). As in Lcptonycteris sanhorni, the secre¬ tory cells are of the seromucoid type. They are triangular in shape and have moderate-sized, somew'hat ovoid, hcterochromatic nuclei that are basally posi¬ tioned. The nuclei are surrounded by den.se, basophilic cytoplasm; the remainder of the cytoplasm is irabeculated, although basophilic granules can be distinguish¬ ed with oil immersion (1500 X , stained with hematoxylin and eosin). The trabec¬ ulae are intensely PAS positive but are essentially nonreactive to both Masson's trichrome and Mallory triple connective tissue stain. The lobules of this gland are characterized by an extremely extensive and complex system of ducts, I'he intercalated ducts, w'hich are unusally long and branched (Fig, 53), arc difficult to distinguish in sections stained only with hematoxylin and eosin. The intercalated duct cells are large and rectangular and have large, round, centrally placed nuclei that are euchromatic. The size of the nuclei in conjunction with the limited amount of cytoplasm and its almost achromatic appearance with H&E account for this difficulty. With PAS or alde- hyde-fuchsin following oxidation wath peracetic acid, the extensive,winding BIOLOGY OF THE PHYLLOSTOMATIDAE 207 Kit.. 51 .—^Two histological views of the highly developed, branched, secretory imer- calated duct system in the parotid of Upfonyclcrii; /livnHs. Large numbers of PAS-positive granules can be seen in the apical cytoplasm of these cells. Abbreviations and symbols are: A, acinus: ICD, intercalated duct; ICD-L, lumen of intercalated duct in cross-section: G(PAS), PAS-positive granules; SD, striated duct; arrow, junction between ICD and acinus. Periodic-acid SchiPTs. Right, 1044 X . left, 392 x, course of the intercalated ducts can be recognized because of the presence of positive staining granules in both the apical cytoplasm and lumen of the duct (Fig. 53). One to three cells are positioned between the secretory intercalated ducts cells and the abrupt beginning of the striated ducts. The former cells are PAS negative, have heierochromatic nuclei, and are more nearly cuboidal than rectangular. The striated cells arc large and columnar; they are slightly basophilic and only weakly acidophilic (hematoxylin and eosin-Y). The nuclei are centrally placed, large, round, and euchromatic. With PAS, the cytoplasm reacts faintly, giving the cell a pinkish cast. The only exception is a narrow band along the apical membrane, which is moderately reactive. The striated ducts are comprised of approximately 30 per cent dark cells, which can be recognized by their small heterochromatic nuclei and proximity to the lumen. The striated ducts give way gradually to interlobular ducts, which are characterized by reduced cell size, loss of basal striations, increased luminal diameter and decrease in number of dark cells. Submandibular .—The submandibular is a compound tubuloacinar gland; the secretory acini are relatively large and densely packed within the lobules of the gland (Fig. 41). The acini are comprised of two types of secretory' cells; the most abundant type has a dense, basophilic (hematoxylin and eosin-Y) cytoplasm, whereas the second type, which forms a demilune, has a pale, almost achromatic 208 SPECIA], F'Util-lCATK^NS MUSEUM t EXAS TECH UNIVERSITY Fig. 54. — Top: Histological overview of submandibular in Li^ptooycteris nivalis^ show ing mixed secretory acini. Abbreviations are: tCD, intercalated duct; M, mucous cells: SD, striated duct; SM, seromucoid cells: Hematoxylin and eosin-Y, 494 X. BoHom: Detail of seromucoid and mucous cells; note the trabeculae (T) and dense ergastoplasm (E) in the seromucoid cells. Hematoxylin and eosin-Y. I I 40 X. cytoplasm (Fig. 54). The first type of cell approximates the classical serous cell and the deniilunar kind is essentially a mucous type. The basophilic serous cells have a moderate-sized, round, basally located nucleus that is fairly hetero- chroniatic. The cytoplasm, when stained with hematoxylin and eosin-Y and observed with oil immersion optics (1500X), is uniformly stained but appears to contain many small vacuoles (or pale granules) of varying size. With PAS, these granules are essentially nonreactive, although the cytoplasm is faintly pink and coarsely granular in appearance (Fig. 54). The mucous cells have smal¬ ler, more hererochromatic nuclei that are basally positioned and surrounded by a small amount of basophilic cytoplasm. The cytoplasm of the mucous ceils contains a trabecular network that is intensely PAS positive. With Masson's trichrome the cytoplasm of the serous cells is pale gray and that of the mucous cells is essentially achromatic. BIOLOGY OF THE PHYl.LOSTOMATIDAE 209 The submandibular duct system is moderately extensive and comprised of relatively simple cells. The intercalated ducts are long and branched; the cells are cuboidal and have round, heterochromatic nuclei that are centrally placed {Fig. 54). The limited cytoplasm is only moderately acidophilic (hematoxylin and cosin-Y) and nonreactive to PAS. The lumina of these ducts also lack PAS- positivc material. Aldehyde-fuchsjn following oxidation with peracetic acid also is nonrcactive. The cells of the intercalated ducts are only slightly stained {pale gray) with Masson’s trichronie iind are non re active w-ith Mallor>' triple connective tissue stain. The intercalated ducts open abruptly into striated ducts (Fig. 54). The latter are comprised of large, columnar, striated cells that have large, round, central¬ ly-placed euchromatic nuclei. The cytoplasm is only moderately acidophilic, staining pale pink with hematoxylin and eosin-Y only after prolonged immersion in eosin-Y. A narrow band of apical cytoplasm is moderately reactive to PAS in the striated ceils. The same region stains purple with aldehyde-fuchsin after oxidation with peracetic acid. The striated cells are only slightly stained (pale gray) with Masson’s trichromc; the basal (striated) portion of these cells stains deep red wdth Mallory triple connective tissue stains. Approximately 20 per cent of the cells comprising the striated ducts can be classified as dark cells. They have heterochromatic nuclei and are PAS negative and are adjacent to the lumen. The transition from striated to interlobular ducts is gradual and characterized mainly by a decrease in cell height and an increase in luminal diameter. The nuclei become slightly more ovoid, and the basal striations are lost. The apical reactivity to PAS and peracetic acid aldehyde-fuchsin found in the striated ceils is tost in the transition. With Masson’s trichrome, the cells of the interlobular ducts are pale gray, and, unlike the striated cells, the basal portion of these ceils is non reactive w'ith Mallory triple connective tissue stains. Both kinds of cells are nonreactive to the peracetic acid aldehyde-fuchsin pro¬ cedure. Subdngtmi .—-The general histology of the sublingual in this species is essential¬ ly the same as that of the same gland in Lepionyctens sanhorni. The reader thus is referred to the description given in that account. Anoura geoffroyi Gross Anaiomy Parotid .—This is a moderate-sized gland located at the base of the auricle. It extends from the angle of the jaw, over a portion of the masseter muscle, to the auricle and, ventrally, to the mastoid region of the skull. This gland is separated from the submandibular by the external Jugular vein. Stenson’s duct arises from the anterior part of the parotid and runs anteriorly into the connective tissue of the upper Jaw' and enters the oral cavity al the level of the first premolar. Submandibular.—This is a moderate-sized gland consisting of two main lobes, each of which is extensively subdivided. The gland lies on the mastoid region of the skull and is positioned in the cervical fossa along with two lobes of the sublingual. The anterior lobe of the submandibular is elongate in shape; it ex- 210 SPECIAL PUBLICATIONS MUSEUM TEXAS IECH UNIVERSITY lends medially from near the cxicrnal jugular and mastoid region and covers a portion of the sternomasioideus muscle. The posterior lobe, to which the an¬ terior one is attached, extends from the lambdotdal region and covers the sterno- niastoideus and external Jugular. Wharton’s duct passes dorsally to the digas- tricus, posterior to the angle of the jaw, and Joins the main duct of the sublingual, Together, these ducts pass anteriorly into the muscle and connective tissue of the floor of the mouth and open into the region of the mandibular symphasis. StthlitttiimL —A relatively large gland that consists of three major lobes. One lobe occupies the typical site for this gland; it is positioned in a depression bor¬ dered by the sternohyoideus, sternomastoideus, and digastricus. This lobe is ovoid and Hat and is overlain by two more lobes. The anterior one of these latter lobes is the largest; it is somewhat triangular in shape and has a convex anterior mar¬ gin and a concave posterior margin. It is bordered by the parotid dorsally and the Sternohyoideus vcntrally. Posteriorly, within the concave margin, lies the last main lobe of the sublingual. It is elongate in shape and is bordered posteriorly by the submandibular gland, A short duct Joins the posterior and anterior lobes and then passes anteriorly, ventral to the digastricus, and joins the main duct from the lobe first described. The common excretory duct has been described above. Histology Parotid .—The secretory acini are densely packed and of the compound acinar type (Fig. 40), When stained with hematoxylin and eosin-Y, the cells have a classical serous appearance. They have small, round, heterochroniatic nuclei usually located basally and surrounded by a strongly basophilic cytoplasm (Fig, 55). These cells contain numerous small granules that can be seen easily when stained with H&E (formalin-flxcd). The secretory cells are PAS negative and stain pale gray-brown with Masson’s trichrome. The numerous lobules of the parotid are connected by an extensive, complex system of ducts. The short, nonbranched intercalated ducts are comprised primarily of cuboidat cells with helerochromatic, round or slightly ovoid nuclei. The relatively small amount of cytoplasm is moderately acidophilic. The inter¬ calated ducts are difficult to locale in any given section because of their short length, which does not exceed three or four cells at most. PAS-positive granules are lacking from these cells, although in many sections the most distal cells (at the Junction with the secretory acinus) do appear to contain some PAS- positive material and also have euchromatic nuclei. The transition betw^een the intercalated and striated ducts is abrupt. The striated cells of the distal pt>rtion are typical in that the large round nucleus is euchromatic and located near the apical membrane of these targe, columnar cells (Fig. 55). The cytoplasm is uniformly acidophilic and the lumen of this portion of the duct system is narrow. In sections stained with Masson’s trichrome, the cytoplasm is pale green, and in those stained with the PAS technique, the cells are nonreactive. Staining with aldehyde-fuchsin following oxidation with peracetic acid also is negative. Relatively small, dark cells having hetero- B10[.0GY OF THE PHYLl.OSrOMATfDAE 21 I Fig. .*55.— Top: Histological overview of the parotid in Anoitm m'offmyL Abbreviations are: A, acinus; BD, excretory duct; 1C, intercalated duct; SD. striated duct. Masson's trl- chronte. 494 X. Bouom: Portion of the striated duct apparently having a secretory function. Abbreviations are: am, apical membrane and adjacent cytoplasm; dc. dark cell; ep, ergasto- plasm; L lumen; v, apical "vesicle.” Hematoxylin and eosin-Y. 1140X. chromatic nuclei are found throughout the striated duct system and comprise as many as 30 per cent of the total number of cells. The proximal portion of the striated duct is characterized by striated cells that apparently are secretory. These large, columnar cells have large, round euchromattc nuclei that are centrally positioned (Fig. 55). The basal cytoplasm around the nucleus is strongly acido¬ philic and striated, whereas the apical cytoplasm is slightly basophilic (H&E), Although easily distinguished, the pale basophilic portion of the cytoplasm does not form a distinct vesicle (Fig. 55). In sections stained with Masson’s trichrome, the basal cytoplasm is green, whereas the apical cytoplasm is pale, nearly achro¬ matic. The entire cell is, however, nonreactive to PAS, The cells of the excretory ducts arc nearly cuboidal, have round, centrally placed, somewhat heierochroniattc nuclei, and lack basal striations. The cyto- 212 SPECJA[, PUH[J CAT IONS MUSEUM TEXAS TECH UNIVERSITY plasm is strongly acidophilic when stained with hematoxylin and eosin-Y and pale green when stained with Masson’s trichrornc. Additionally, these cells are non reactive to PAS. On the other hand, a considerable amount of PAS-positivc material typically is found in the lumen of this portion of the duct system. Sithnuuuiihitlar ,-—The submandibular is a compound tubuloacinar gland that is densely packed with small, round secretory acini (Fig-41). Tw'o types of secretory cells can be distinguished easily with standard stains. Both have the appearance of being serous cells when stained wdth hematoxylin and eosin-Y, The secretory cells (Fig. 56) that surround the acinar lumen are large, essentially rectangular or slightly triangular, and have nearly euchromatic nuclei. The cyto¬ plasm has a uniform, finely granular appearance (H&E). These granules are moderately PAS positive. The second type of secretory cell is relatively small and Bat but numerous enough to form a cap over the entire secretory acinus (Fig. 56). These cells arc basophilic and generally resemble the special serous ceils described elsewhere by Wilborn and Shackleford (1969). The nuclei are small, round, basally positioned, and hcterochromatic. When stained with the PAS technique, the cytoplasm is generally reactive and appears to contain large numbers of small secretory granules. The outer layer of special serous cells stains uniformly green with Masson’s trichrome, w'hereas the inner secretory cells are green, but distinctly paler. The duct system of the submandibular is of moderate length and of relatively simple morphology. The intercalated ducts are of moderate length, branched, and comprised of small cuboidal cells with round, centrally placed heterochro- matic nuclei. The cytoplasm is restricted but clearly acidophilic whth hematoxylin and eosin-Y. 4’hese cells are PAS negative and essentially nonreactive to Mas¬ son’s trichrornc. The transition between intercalated and striated ducts is characteristically abrupt. Judging from the relative paucity of striated ducts in any given section, this portion of the duct system apparently Is short. The major type of cell is large and columnar and has a large, round, centrally placed euchromatic nucleus (Fig, 56), Basal striations are easily discernible; the cytoplasm is typically acidophilic with hematoxylin and eosin-Y. The striated cells are nonreactive to PAS and stain pale green with Masson's trichrome, Approximately 10 per cent of the cells in the striated portion of the duct can be classified as dark cells, judging from their small and hcterochromatic nuclei and proximity to the duct lumen. The transition from striated to interlobular (excretory) ducts is gradual and characterized by an increase in luminal diameter, a decrease in cell height, and eventual loss of basal striations. Stihlifigtuii .—The numerous lobules of this gland have a moderate duct system comprised of relatively simple cells. The short, branched, intercalated ducts are easily seen. The cells comprising these ducts are nearly rectangular and narrow except near the Junction with the striated ducts w'here they appear to be some¬ what more cuboidal. The hcterochromatic nucleus is ovoid and occupies most of the cell. The small amount of cytoplasm is only moderately acidophilic (hem* atoxylin and eosin-Y) and nonreactive to PAS, Masson’s trichrome, and Mallory BIOLOGY OL THE LHYI.I.OSTOMATIDAE 213 Fig. 56. — Top: Histological overview of siibmandibular of A/ioura ffeojfroyi. Masson’s trichrome. 481 X . Bottont: Detail of a secretory acinus: note the extruded materials (arrows) at the apical membranes of the seromucoid cells. Abbreviations are; A, acinus: ED. excretory duct; L. lumen; SD, striated duct; SM, seromucoid cells; SS, special serous cells, Masson's trichrome. 1258X, triple connective tissue stains, PAS-positive granules were not found in the lumi- na of intercalated ducts. The secretor>' acini are of moderate density and compound tubular in nature. The secretory cells, when stained with hematoxylin and eosin-Y, have the clas¬ sical mucous appearance; the cytoplasm is relatively clear although a fine trabec¬ ular network can be distinguished- The nuclei are small, round, and hetero- chromatic. They are basally located and typically surrounded by a small amount of basophilic (with hematoxylin and eosin-Y) cytoplasm. The trabeculae of the cytoplasm of these cells are intensely PAS positive. Furthermore, they stain dark purple with aldehyde-fuchsin following oxidation with peracetic acid. The cells are essentially nonreactive with Masson’s trichrome and Mallory triple connective tissue stains. 2M SJ>EC]A[. PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY The transition between the intercalated and striated duels is abrupt. The striated cells of the latter are large and coluninar and have large slightly ovoid etichromatic nuclei that are apically displaced. The cytoplasm is strongly acido¬ philic with hematoxylin and eosin-Y, nonreactive to PAS, and pale green with Masson's trichrome. Additionally, these cells stain intensely red with Mallory triple connective tissue stains. The transition from the striated to interlobular ducts is gradual and characterized by a decrease in cell height concomitant with an increase in luminal diameter and slight decrease in size of the nucleus. The basal striations are lost. The staining reactions of the cells of the interlobular ducts, with the techniques used in this study, were the same as those of the striated cells. Sturnira JudovieJ Gross ^fiiito/tiy Parotid .—The parotid is extremely large (Fig. 39), its ventral margin extends to the ventral midlinc of the throat and overlies the posterior half of the sub¬ lingual gland. Dorsal ly, the parotid extends around the cervical region and covers the area where the digastricus and sternomastoideus cross, posterior to the masseter. A small portion of the gland extends anteriorly between the auricle and masseter. The remainder of the glandular mass lies over the mastoid, lamb- doidal, and occipital regions of the skull and extends into the cervical fossa where it is partially covered by the posterior half of the submandibular. Slenson’s duct arises from the anterior edge of the parotid and runs along the ventrolateral surface of the masseter in a groove between the masseter and digastricus, fol¬ lowing the contour of the masseter anteriorly and dorsalty. The duct continues anteriorly in the connective tissue and muscle of the upper lip and enters the oral cavity at the level of the posterior side of the canines. Sifhtnandibtdar .—-This is a large gland that fills the cervical fossa from the lateral midlinc to the ventral midline of the throat in the region of the sterno¬ mastoideus (Fig. 39), The gland is triangular in cross section; its posterior sur¬ face lies against pectoral muscles and tire inferior surface lies against the parotid gland. Wharton's duct arises from the center of the inferior surface, passes under the ventral margin of the parotid, over the dorsal surface of the digastricus in the region of the angle of the mandible. The duct runs along the floor of the oral cavity in the region of the ventrolateral margin of the tongue. It is joined by the main duct from the sublingual gland at the base of the tongue and together they enter the anterior part of the oral cavity, opening near the first tower pre- molar. Sitbiingual .^—'This is a moderate-sized gland that is triangular in shape and soft and loosely encapsulated in thin sheets of connective tissue (Fig. 39). This gland is unilobular but its numerous fine subdivisions can be seen clearly. The anterior half of the gland is covered by the posterior margin of the mylohyoideus; the medial edge lies against the sternohyoideus and the lateral edge is in contact with the medial surface of the digastricus. The duct arises on the inferior sur¬ face and joins the main excretory duct of the submandibular near the base of the tongue (sec above). BJOLOC.Y Oh THE PHYLLOS'J’OMATIDAE 215 Histology ParotuL —The parotid is a compound acinar gland that is densely packed with elongate secretory acini comprised of typical serous cells (Figs. 40, 57). These cells are basophilic with hematoxylin and eosin-Y; the cytoplasm appears to be finely granular when observed w'ith oil immersion (Fig. 57). The nuclei of the secretory cells are basally positioned, small and round, and heterochro- matic. Small, densely packed secretory granules can be distinguished in the apical cytoplasm when stained w'ith the PAS technique (Fig. 58). At the same time, however, these granules are not stained mth aldehyde-fuchsin follow'ing oxidation w'ith peracetic acid. With Mallory' triple connective tissue stain the cytoplasm contains relatively large red granules that apparently are not secretory product, judging from their size. The parotid is characterized by an extensive system of ducts. The intercalated ducts are of moderate length, branched, and comprised of two distinct segments. Adjacent to the secretory acini, the intercalated duct cells are large and rectan¬ gular; the nuclei are euchromatic and the cytoplasm contains relatively large, intensely PAS-positive granules (Fig. 58). The remainder of the intercalated duct is comprised of rectangular or cuboidal cells with heterochromatic nuclei, restricted cytoplasm, and no trace of PAS-positive materials. The transition from intercalated to striated ducts is abrupt (Fig. 57). The latter portion of the duct system is comprised of large columnar cells with cen¬ trally positioned euchromatic nuclei. The cytoplasm is acidophilic when stained W'ith hematoxylin and eosin-Y. Intensely stained, PAS-positive granules are found in the apical cytoplasm of these cells. These granules also stain with al- dehyde-fuchsin following oxidation with peracetic acid. Approximately 10 per cent of the cells in the striated ducts can be classified as dark cells based on their small and heterochromatic nuclei and absence of PAS-positive staining. The transition from striated to typical interlobular ducts is gradual and char¬ acterized by a decrease in cell height and eventual loss of basal striations. Submandibular .—The submandibular is a compound tubuloacinar gland with large and densely packed secretory acini (Fig. 41). The majority of each acinus is comprised of narrow columnar serous cells with basally positioned and somew'hat heterochromatic nuclei. The cytoplasm of these cells has a distinctly dense, basophilic basal region and a slightly paler but basophilic middle and apical portion (Fig. 59). The apical cytoplasm contains densely packed, small PAS-positive granules that also stain w'ith aldehyde-fuchsin following oxidation W'ith peracetic acid (Fig. 59). The demilunar secretory cells are of the mucous type; the nuclei are basally restricted and heterochromatic. The cytoplasm, with exception of a distinct basal ergastoplasm, is pale and somewhat trabecular when viewed with oil immersion (1500x). The vacuoles in the cytoplasm are PAS negative but the trabecular netw'ork, which presumably represents restricted cytoplasm between secretory granules, is PAS positive and reactive to aldehyde fuchsin peracetic acid (Fig. 59). The duct system of the submandibular is relatively simple. The intercalated ducts are short and branched but are difficult to locate in any given section. 216 Sf^ECIAL J^LJBLJCATIONS MUSBUM TEXAS lECH UNIVERSITY Bui, 57.-— Top: Parotid salivary gland of Ui{U)vici\ the arrow shows direction of flow of formative saliva from an acinus (A) to the striated duct (SD) via the intercalated duct (IC). The other abbreviation is: ED, excretory duct. Hematoxylin and eosin-Y. 900X. Botfom: Parotid acini showing a myoepithelial cell (M}, the thin connective tissue capsule (C). and distinct ergastoplasm l EP) of the secretory cells. Hematoxylin and eosin-Y. 1 125 X. The cells arc rectangular and have ovoid, heterochromaiic nuclei and a limited amount of slightly acidophilic cytoplasm. The intercalated ducts are generally PAS negative although some of the cells occasionally have a slightly pinkish cast. The transition to striated ducts is abnipt. The striated cells are columnar, have a centrally positioned, large, round, and euchromatic nucleus, and are PAS negative. These ducts arc comprised of approximately 20 per cent dark cells characterized by heterochromatic nuclei. The interlobular duct can be recognized by the decrease in cell height, toss of basal striations in the major cell type, and an increase in luminal diameter. Suhlingtuii .—The sublingual is a compound tubuloacinar gland comprised of mucous cells. I’he secretory cells have basally restricted, irregularly shaped, mOLOGY OF THE PHYLLOSTOMATIDAE 217 Fkj, 58.— Top: PAS reactivity in the secretory acini of a parotid gland from Snirnini tudovk i. Note the nonreactive ergastoplasm (EP) and striated duct (SD). Other abbreviations are: AL, acinar lumen; dc. dark cell: F, fibroblast; 1C, intercalated duct; pc, pale striated cell. Periodic-acid SchifFs and hematoxylin and eosin-Y. 481 X. Bottotn: PAS- positive granules (G) in intercalated duct cells (1C). Periodic-acid SchifTs. 814 x. heterochromatic nuclei. The adjacent basal cytoplasm is fairly dense and baso¬ philic. The bulk of the cytoplasm is pale when stained with hematoxylin and eosin-Y and is highly vesiculated when view-ed with oil immersion optics {1500 X). The trabeculae are intensely PAS positive and alsct reactive to alde- hyde-fuchsin following oxidation with peracetic acid. The duct system of the sublingual is relatively simple. The intercalated ducts are long and branched and comprised of flat or rectangular cells having little cytoplasm and ovotd or round heterochromatic nuclei. These cells show- only slight reactivity to the PAS procedure. The Junction betw'een intercalated and striated portion of the duct system is abrupt. The latter is comprised of large, columnar cells with apically positioned, large, round and euchromatic nuclei. The cytoplasm is acidophilic (H&E) and PAS negative. 21K SPECIAL PUBLICA'LIONS MUSEUM LEXAS TECH UNIVERSITY Fig. 59. — Top: PAS reactivity (G PAS+) in the serous cell (S) in the submandibular gland from Smrnint huiovici. Note the nonreactiviiy of the mucous cells (M) and the wide acinar lumen (L). Periodic-acid Schiffs and hematoxylin and eosin-Y. 988X. Bottom: Serous secretory cells, showing the granular apical cytoplasm (GCK euchromaiic nucleus (N), striated basal ergastoplasm (EP), Note the distinct intercellular canaliculus (ICC). Hematoxylin and eosin-Y. 1596 X. Comparisons Parotid and submandibular salivary glands are compared in the following paragraphs. We have not included the sublingual because of its conservative character. Our summary statements are based partly on our own observations and partly on Wimsatt (1955* 1956) and DiSanto {I960). Additionally* for ease of presentation we have compared species gland by gland rather than species by species. Parotid .—As pointed out in the general results of gross anatomical examina¬ tions, the parotid gland varies considerably in size* degree of compactness, and, to a certain extent, in position in the aural region. None of these features was found to be related to the general histological characteristics of the gland. In B[OLOGY OF TtlE HHYLLOSTOMATIDAE 219 comparison to the other species, this gland is relatively large in Artihcits, where it extends into the interscapular region, and relatively small in Desmodits. Comparisons of size or appearance, or both, do not reveal any relationship with taxonomic grouping, at least insofar as w'e could determine. The general histological features of the parotid vary as greatly as does the gross anatomy. The phyllostomatids studied thus far can be grouped relatively easily on basis of structural and tinctorial characteristics of the secretory acinar cells. In all seven species the parotid can be described best as compound acinar although the shape and size, as w-ell as number, of secretory acini vary consider¬ ably. For example, in Ariibeiis and Sturnira the acini are elongated and nearly tubular, w'hereas in Leptonycierls fiivalis, L. sanhonii^ and dossophaga they are relatively small and nearly bulblike. In the other species, the parotid secre¬ tory acini are irregularly shaped, of moderate size, and more typically compound acinar. As we have pointed out in the section on Materials and Methods, the terms serous, mucous, and seromucoid are used by us only guardedly and for purposes of description rather than on the basis of knowledge about chemistry of secre¬ tory product. In four species {Glossop/iiiga sp., lu^plonycteris nivalis, L, sanhorni, and Artibens sp.), the secretory cells have been described as seromucoid based on their relatively achromatic cytoplasm (when stained with either hematoxylin and eosin-Y or azure-A and eosin-B) and morphologically intermediate position. In the remaining species, the secretory cells have dense basophilic cytoplasm and are fairly representative of typical serous cells. On the basis of characteristics discernible as a result of the methods used by us, comparisons allow' for the following groupings. Three glossophagine species {Glossophaga soricina, Leptonyaeris nivalis^ and L. sanborni) are notably similar in both morphology (Fig. 40) and tinctorial characteristics. In each kind, the apical cytoplasm of the large, somewhat triangular secretory cells contain vac¬ uoles of variable sizes that with the light microscope appear to be outlined by a PAS-positive trabecular network. The notable differences within this group arc: 1) in Clossophaga, the PAS reaction is slightly more intense; and 2) the number (in terms of density) and size of the secretory alveoli are by far the great¬ est in Giossophaga and the least in L. nivalis. The parotid secretory acini of the fourth glossophagine species, Anouni geoffroyi, differ considerably from those of the others. The acini in this species are comprised of secretory cells that best can be described, as seen through the light micro.scope, as serous. A second grouping includes Sinrnira and Desmodus, in w'hich the parotid secretory cells are notably similar. In both species the acini are irregularly shaped and somew'hat tubular and the secretory cells have dense, generally basophilic cytoplasm that stains blue pink with hematoxylin and eosin-Y and azure-A and eosin-B. With PAS, the cytoplasm of these cells can be seen to contain small, evenly distributed granules. The remaining species, Artibi’us jcimaicensis^ is distinctly different from the others. The secretory cells morphologically are most similar to the seromucoid type, but the cytoplasm is relatively achromatic, lacks vesicles or trabeculae 220 SPECIAL PU Li Lie AT IONS MUSEUM TEXAS TECH UN i VERS [TV such as those found in ihe glossopliagines having seromucoid parotid secretory cells, and are PAS negative. Additionally, histochemical analysis suggests that they should be classified as serous ceils (Wimsatl, 1956). The intercalated segment of the duct system also varies greatly. In Ammnu it is relatively short, usually being only three or four cells in length, where¬ as in most species it is long (sometimes extremely so) and highly branched. In addition to differences in length and degree of branching, the intercalated ducts also differ notably in cellular composition as revealed with the light microscope. Each of the four, glossophagines studied are different, even at the congeneric level (Fig- 40). In Glossophugu the somewhat naitened cells apparently are nonsecretory or, at least, essentially PAS negative. In the closely related LL’ptonyaeris satiboruu the initial segment of intercalated duct is comprised of secretory cells (PAS positive), whereas the proximal-most segment resembles the intercalated duct of Glossophaga. In L, nlvaiis, on the other hand, the extremely long intercalated ducts are almost entirely secretory. Only the last one or two cells separating this segment from the striated duct are nonsecretory (Fig, 40). Anouta geoffroyi, the other glossophagine studied, is characterized by one or two secre¬ tory intercalated duct cells adjacent to the secretory acinus, but the remainder of the short duct is comprised of nonsecretory cells (Fig. 40). This pattern also is found in SttirninL In Anibtus and Desfuiulus, the intercalaied ducts contain PAS-positive material and the cells possibly are secretory, but their morphology at the light microscope level of magnification is slightly different from that of the secretory intercalated duct cells at least in the glossophagines. The striated portion of the duct system in all seven phyllostomatid species is characterized by a narrow lumen and large, columnar cells with basal striations (mitochondria and basal tnfoldings of the cell membrane) interspersed w'ith smaller cells distinguishable with the light microscope by their small and hetero- chromatic nuclei, A PAS-positive apical staining reaction of granules of various sizes suggests strongly that the striated cells in the parotids of Glossophaga, Lepumycteris uividls, Sturnira, and Desmodits are secretory and thus add directly to the formative saliva. The absence of PAS-positive granules (or materials detectable by the other stains and histochemical procedures used by us and by VVimsatt, 1955, 1956) in the striated cells in Anibens jamaicemis implies that these cells are nonsecretory in this species. Leptonyaeris sanbomi and Amnira geoffroyi differ from the other species in that the striated cells are PAS negative, but, nevertheless, the apical cytoplasm stains differently from that of the remain¬ der of the cell, suggesting a specialized function. The role of this somew hat vesic¬ ular-appearing region w'ill be undetermined until ultrastmctural or special histochemical studies are undertaken. SubmandibiiUtr .-—As is the case with the parotid, the submandibular varies considerably in gross anatomical characteristics. The variable features (size, position, appearance, and number of lobes) can not be used for meaningful tax¬ onomic groupings or for groupings rellccttve of apparent food habits or general histology. Two basically different types of secretory acini are found in the submandib¬ ular of the species studied. In all of the species except Anoura geoffroyi, the BJOLOGY OF THE PHYLLOSTOMAT[DAE 221 acini are both mixed and tubular in morphology. The glossophagtnes other than Afioura differ from each other but display a definite pattern. In Gi(>ss(^pha^>iiy the secretory acini include a large number of serous cells (Fig. 41) that contain large-sized, PAS-posltive granules. The acini are capped with demilunes of mucous cells. In basic light microscope morphology and proportion of .serous to mucous cells, the secretory acini of Leptonyvieris nivalis are similar to those in Giossopha^’n (Fig. 41). How-ever, some of the tinctorial reactions, especially that to PAS, are quite different; in L. nivalis, the serous cells are not densely packed with large-sized PAS-positive granules. Instead, the cytoplasm typically is packed with extremely small, evenly distributed PAS-posilive granules. Lc‘pto- nycteris sanharni differs greatly from the other two species in that there are far fewer serous cells and far more demilunar mucous cells per acinus (Fig. 41). At the same time, however, the PAS granules in the .serous cells are more like those found in the homologous serous cells of Glossophaga than they are like those of its congener, L. nivuiis. I'he other glossophagine, Anonni, is ex¬ tremely different. In this .species, the secretory acini are small and bulbous. Al¬ though mixed, both types of cells appear to be of the serous type. Most striking is the fact that the inner layer of secretory cells is covered completely by a layer of small, Hat cells (Fig. 41). This “demilune"’ is histologically and morphological¬ ly similar to the “special serous" cells found in the submandibular of the op- posum, Dideiphls (Wilborn and Shackleford, 1969). The degree of difference between the secretory acini of both the submandibular and parotid of Anouru and those of other glossophagines described here is striking, at least at the light microscope level. These notable differences, considered in view of dental, serological, karyological, and digestive tract differences (Phillips, 1971; Gerber and Leone, 1971; Baker, 1967, 1970; Forman, 1971, 1972), suggest that salivary gland histology might be useful in systematic analysis. AnibeiiS, Stnrnira, and Destnodns have tubular mixed secretory acini com¬ prised of serous cells containing PAS-positive granules and a demilune of mucous cells. Fhe serous granules differ from those in comparable cells in Glossophaga and Leptonycieris sanborni by being smaller and more densely packed. Overall, how'cver, the submandibular secretory acini of the phyllostomatids thus far studied, excepting Anonra, show' a considerable degree of conservatism in general histology at the light microscopic level. The intercalated ducts of the submandibular vary somewhat in length but are similar in all of these species. In Anibeus, the cells adjacent to the secretory acini apparently contain PAS-positive granules w'ithin their cytoplasm, but in all of the others the intercalated duct cells are histologically simple and PAS negative. The striated ducts also are relatively simple and similar in each of the species studied. Three species {Glossophaga soricina, Lvptonyaeris nival is and L. san¬ borni) can be grouped on the basis of PAS-positive staining reaction in the apical- most cytoplasm (distinct granules can not be discerned), whereas in all of the others the striated cells are PAS negative. 222 SPEC[AL PUBLICATIONS MUSEUM I EX AS TECH UNIVERSJ [Y Ultrastriicture General histological studies such as those described in the preceding paragraphs provide an overview of mic roan atomy but fall short of answering the most intriguing questions. Histochemical analysis, which can provide con¬ siderably different kinds of information about cellular activities and secretory products have to date been applied to only tw'o species, Arfiheus Jamaicensis and Desmdihis nHutuiits. Both of these techniques suggest, how'cvcr, that salivary glands of bats in general, and phyllostomatids in particular, wdll provide a fertile source of data on evolutionary biology and systematics. A third approach, com¬ parative ultrastructure, is clearly warranted; the histological differences observ¬ able with the light microscope illustrate an opportunity for analysis of evolution¬ ary changes at the ultrastructural level. Although the parotid and submandibular salivary glands of a reasonably wide range of mammalian species have been investigated, and to some extent compared, at the ultrastructural level, to date, none of the Chiroptera has been described. We recently have had an opportunity to study the ultrastructurc of the parotid secretory acini and duets and the submandibular secretory acini of the dw'arf fruit-eating bat, Arfiheits p}uu'(/!is. Our findings, which are presented in the following paragraphs, are intended to serve as an example of phyllostomatid salivary gland ultraslructure and as a basis for future comparative studies. Paro!'ui. —^Thc parotid salivary gland of Artilu'us phaearis is a compound acinar gland with tightly packed, slightly elongate secretory acini. From an ultrastructural standpoint, the parotid acinar cells are best de.scribed morph¬ ologically as seromucoid (Fig. 60), The basal plasmalcmma is delicately infolded and lacks concentrations of free ribosomes (RNP) and granular endoplasmic reticulum (GER). Consequently, there is a distinct margin that continues along the base of the cell and extends along nearly two-thirds of the lateral cell surfaces. The apical one-third of the lateral margin is either smooth or slightly infolded or has narrow intercellular canaliculi lined with microvilli. Each acinus in surround¬ ed by a typical basal lamina; myoepithelial cells typically overlay the secretory acini. The nuclei of the secretory cells arc basally displaced, considerably hetero- chromatic, and have a nitKlcrate-sized nucleolus. The outer margins of the nuclei frequently have a scalloped appearance, possibly as a response to pressure from adjacent membrane-bound secretory vesicles and organelles. Elongate mito¬ chondria are found throughout the cytoplasm but most often are positioned along the cell margins, just internal to the infolded plasma membrane. Stacks of GER are found in the basal portion of the secretory cell, usually cradling the nucleus. Short segments of GER also are distributed throughout the remaining cytoplasm. Densely packed free ribosomes also are numerous, particularly in the cytoplasm between secretory vesicles. Golgi complexes are located in the basal half of the cell frequently but not necessarily adjacent to the nucleus. Typically, they have only a few tlatiened lamellae, but numerous swollen cisternae contain¬ ing pale, ftocculent material and electron dense particles. Relatively large sac¬ cules containing these materials can be observed adjacent to cisternae and secretory vesicles. The presence of membrane-bound saccules of secretory nia- BIOLOGY OF THE PHYI.LOSTOMATIDAE 223 Fig. 60. — -Transmission electron micrograph (TEM) of parotid secretory cells in Ariihetis pfuieofis. Abbreviations and symbols are: arrow, basal lamina; BI, basal infoldings of the plasma membrane; GC, Golgi complex; GER, granular endoplasmic reticulum; M, mito¬ chondria; SG, secretory granules (note the electron dense material indicated by an arrow); S, saccules forming an immature secretory granule, 20,625 X . lerials within some of these vesicles, as well as at their boundaries, probably reflects the method by which the secretory vesicles are filled. When the con¬ tents of saccules are released, their membranes apparently become part of the limiting membrane of the secretory vesicle. The mature secretory vesicles usual¬ ly contain a relatively small accumulation of electron dense material as w-eil as barely visible flocculent material, at least under the conditions of fixation used by us. Generally, however, most of the area within a secretory vesicle is essential¬ ly electron transparent; it is possible, of course, that some components of the secretory product were solubilized in the extensive fixation employed. The se¬ cretory granules are negative to both toluidine blue {1 -micrometer epoxy sections) 224 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Fig. 61.—TEM of parotid intercalated duct showing cytoplasmic granules (arrows) in the pale duel ceils. Abbreviations not explained in Fig. 60 are: NU, nucleus; BCN, basal cell nucleus; TCN, triangular dark cell nucleus-. RBC, red blood cell. 4760 X . and PAS (7-micromeicr paraffin sections). The apical membrane of the sero- mucoid parotid secretory cell is relatively smooth. We found no indication of microvilli in available examples of the acinar lumen. The branched parotid intercalated ducts are comprised of at least two types of cells (Figs, 61,62). I he most prominent type is rectangular and has a relative¬ ly clear cytoplasm and a large, centrally located nucleus {Fig. 61). The short strands of GER are not regularly arranged, instead being found throughout the cytoplasm; the cisternae arc greatly swollen, giving the short strands a round, vesicular appearance (Fig. 63). At higher magnifications the cisternae can be seen to be filled with moderately electron dense, fiocculent material. Free RNP particles are found in considerable numbers most often in the juxtanuclear re- BlOl.OGY OF THE PHYLLOSTOMATIDAE 225 Fig, 62.—TEM views. Top: Branched parotid intercalated duct, showing the lumen (1.) and direction of flow of formative saliva. Abbreviation not explained in Fig, 60 is: AC, acinar cell. 2800X. BtHiom: Detail of apical membrane and cytoplasm of a striated dud dark cell. Note the flocculeni material (arrow) in the lumen (L) and the micropinocytotic vesicle(PV'). Abbreviation not explained in Fig, 60 is: C, ceniriole, 25,200X. gion. The marginal cytoplasm in these intercalated duct cells is characterized by large numbers of oriented filaments, particularly in areas adjacent to the zone of contact with other cells of the same type (Fig. 63). Additionally, the zones of contact between these cells are characterized by relatively flat plasma mem¬ branes with extensive desmosomes (Figs. 61-63). By way of contrast, the Junc¬ tions between these intercalated duct cells and the two other peripheral cell types, always lack desmosomes and tend to be moderately interdigitated (Figs. 61, 63). The pale cells have relatively small, elongate, branched mitochondria, w'hich 226 SPECIAI- PUBIJ CAT IONS MUSEUM TEXAS tECl I UNIVERSITY F:t appear either posilion- cd or oriented although they generally are lacking from cytoplasm containing concentrations of secretory product. In some instances mitochondria seem to be concentrated in the vicinity of Golgi complexes but this can not be determined with certainty. The Golgi complexes, because of their extreme size, are prominent cyto¬ plasmic feature (Fig. 66). Golgi components and several initial steps in synthesis of secretory granules are show n in Fig. 68. The border of the complex consists of flat lamellar membranes. The apparent steps of granule formation within the Golgi complex, as illustrated in Fig. 68, are as follows: 1) There is develop¬ ment of a tubular, membrane-bound unit that becomes swollen with accumula¬ tion of small, eiectron-dense particles, pale tlocculent material, and distinct (membrane-bound) clear vesicles. 2) The small vesicles increase in number and the tubule loses its swollen appearance. 3) Spherical, membrane-bound sac¬ cules containing several small vesicles and some tTicculent material become isolated in the cytoplasm. 4) The small vesicles disintegrate, producing a larger granule w ith a distinct, thick outer boundary that is either the result of the origin¬ al limiting membrane being joined by the membranes that bound the vesicles or is due to an accumulation of electron-dense materials at the outer margin of the interior of the granule. The granules at this stage contain irregularly shaped clumps of moderately electron-dense material and a paler, uniform background material. These granules frequently coalesce with one another. The secretory granules resulting from these steps at the Golgi complex oc¬ casionally are found throughout the cytoplasm but most often arc restricted to the apical onc-half of the cell. In general they can be classified into three distinct morphological types: unorganized strands, concentric rings, and tubular networks. The first type (Figs. 69, 70) probably represents an intermediate (condensing) stage betw'een the Golgi complex and the mature product. The other two possibly are stable, mature forms. Serial sectioning demonstrates that the two presumed mature forms of granules are structurally distinct, rather than different in appear¬ ance due to sectioning angles (Fig. 70). The simpler form (concentric rings), actually is comprised of concentric spherical shells of electron-dense material. The other type is comprised of convoluted tubules. Occasionally, secretory gran¬ ules of mixed morphological type or granules containing areas of electron-dense material also arc found. That all of these forms represent mature granules and possibly retlect slight chemical variability, is suggested by the fact that they alt arc extruded from the cell regardless of their morphology. The secretory' IHOLOGY OF THE PHYLLOSTOMATIDAE 233 Fig. 68.— I EM detail of seromucoid cel! Golgi complex, showing numerically the prob¬ able packaging sequence {numbers correspond to those in text) and condensing vacuoles (CV). 12,716 X. granules are negative to both toiiiidine blue (1-micrometer epoxy sections) and PAS (7-micrometer formal in-fixed paraffin sections). The method of apical extrusion does not appear to be unusual. The membrane of the secretory granule becomes fused with the apical plasmalemma; microvilli are lost or at least lacking from the site. The contents of the granule are extruded into the lumen and apparently are broken down immediately as the only material found within the lumen appears as a pale, fiocculcnt background. Discussion ami comparisons .—The ultrastructure of the parotid and sub¬ mandibular salivary glands of Ardhetts phacoiis can not yet be compared directly to that for any other chiropieran. Judging from obvious interspecific differences St^EClAl. PUm.lCATIONS MUSEUM TEXAS TECH UNIVERSITY 2? 4 Ft(k 69.=—A TEM comparison of mature (SG) and immature seromucoid secretory granules (ISG). Note the tube! ike morphology of some granules (arrow). 24,480 X . in general histology of salivary glands, such comparisons, when possible, should prove to be unusually iiiieresiing and valuable to both oral biology and evolution¬ ary studies. The parotid secretory cells of ArithvKS pfuu't^th qualify morphologically (but not tinctorially) as seronuicoid based on the definition given by Shackle lord and Wilborn (1968). Ehe GER is flat and the Golgi complexes prominent. Serous secretory cells, by way of contrast, frequently exhibit swollen GER with floc- culent material visible within cislernae (Parks, 1961; Kayanja and Scholz, 1974; Tandler and MacCalliim, 1972) and, thus, resemble pancreatic acinar cells (Jamieson and Palade, 1967^;, 1967 /j, 1971). The secretory granules are con¬ siderably more electron lucent in A. p/u/co//.v than are those found in parotid cells of other mammals for w'hich data are available. For example, in alb ini Stic house HiOLOGY OF THE PHYt.LOSTOMATlDAE 23 s Fig. 70.—TEM serial sections through the concentric shell (a-k) and "tubular" fi-s) se¬ cretory granules in the submandibular seromucoid cells. 35,000X. mice {Mm mmcuim) ihe secretory product is denser although often bizonal (Parks, 1961) and in ungulates it typically is uniformly electron dense (Kayanja and Scholz, 1974). Aside from the relatively small, irregular masses of electron- dense material, the secretory granules in A. phacoiis morphologically resemble 236 Sl^EUAl. PUBLICATIONS MUSEUM TEXAS TECH UN[VERS]TY mucous granules. It is surprising, therefore, that the parotid secretory cells in this phyllostomatid are reactive to neither PAS nor toluidine blue. The extensive infolding of the basal plasma membrane and basal two-thirds of the lateral plasma membrane suggests a specialization for intake of raw materials. The arrangement of mitochondria adjacent to the apices of membrane folds probably is more reflective of the role of mitochondria in supplying ATP for active transport (DeRobertis tT aL, 1970) than it is a reflection of lack of space elsewhere in the cytoplasm. The relationship between adjacent parotid secretory cells in A. phacniis differs somewhat fnmi the usual pattern. In pliaeoiis, the intercellular canaliculi are narrow in diameter and of uncertain length, and in no case do adjacent cells have a loose inierdigitaLion. In the squirrel jnonkey {Saimiri mureus) and in several species of ungulates and rodents, adjacent secre¬ tory cells generally are loosely interdigitated and have large intercellular can- aliculli (Cowley and Shackleford, 1970;7; Kayanja and Scholz, 1974; Shackleford and Schneyer, 1964). The duct system of the parotid of A. phaeolis is not unusual at the ultra- structural level although it differs in detail from that of other species for which data have been published. The intercalated portion apparently is secretory, judg¬ ing from the presence of PAS positive granules of variable electron density with¬ in the cytoplasm of the major cell. The general cytoplasmic features of these cells, particularly the presence of large numbers of fibrils, are consistent in a variety of species in both the parotid and submandibular salivary glands (for example, see Shackletbrd and Wilborn, 1968, I970U, 1970/); VVilborn and Shackleford, 1969). The ultrastructural characteristics of the intercalated duct cells in A. pluieoiis do not suggest adult functions aside from a possible secre¬ tory role, even though the possibility exists that the intercalated ducts are in¬ volved in certain physiological functions (Rutberg, 1961). The striated portion of the ductal system is nonsecretory in A. phaeotis, The absence of sjiiall, electron-dense granules in the apical cytoplasm and the smooth apical membrane in the pale "striated” cells make them different from homol¬ ogous cells in rodents and certain primates (Parks, 1961; Cowley and Shackle¬ ford, I970^d* The complex, loose infoldings of the basal plasnialemma and ob¬ vious association of oriented mitochondria found in A. phaeods is typical of striated cells. The striking ultrastructural similarity between these cells and those of the renal distal tubule has led to frequent physiological and ultrastructural comparisons (Rhodin, I958u, 1958/); Rutberg, 1961; Tandler, 1963), Although these striated cells vary considerably from species to species, active resorption likely is one consistent role (for example, see Rutberg, 1961). The presence of dark cells (characterized by dense cytoplasm) among typical striated cells is common in mammals. Histologists generally either have over¬ looked, or at least have not reported, this type of ceil even though it can be rec¬ ognized in typical histological preparations because of its small size and small hetcrochromatic nucleus located adjacent to the ductal lumen. The function(s) and origin of these cells arc unknown although in A. phaeotLs the available micro¬ graphs strongly suggest pinocytotic activity along the luminal surface. We BJOLOGY OF THE PHYLLOSTOMATIDAE 237 found no evidence indicating that the dark cells are in any way necrotic although others (for example, Kayanja and Scholz, 1974) have reported possible mito¬ chondria) destruction in similar cells. The ultrastructure of the submandibular of A. phaeoiis is unique among studied species of mammals. The arrangement of serous cells capped with an extensive seromucoid demilune differs from the structural features of primates and rodents (Cowley and Shackleford, 1970b; Shackleford and Schneyer, 1964). In man, for example, the submandibular is basically a serous gland although there also are isolated mixed alveoli in w'hich mucous cells are capped by a demilune of serous cells (Sicher and Bhaskar, 1972). The arrangement in A. phaeoris, which is typical in the submandibular of chiropterans, structurally resembles that found in the European hedgehog, Eriimceiis eim^paeus (Tandler and MacCallum, 1972). Useful ultrastructiiral comparisons presently possible can be made between Anibeita phaeotis' and Erituiceus europaeus. In A. phae(?ns, the submandibular serous secretory cells can be described as typical, whereas the seromucoid demi- lunar cells are highly unusual. In the European hedgehog, on the other hand, the demikmar mucous cells are typical and the serous secretory cells are highly un¬ usual (Tandler and MacCallum, 1972). In the latter species the immature secre¬ tory product of the serous cells morphologically resembles, to a remarkable extent, the mature secretory product of the seromiicous cells in A. phaeotis. In both instances the granules have the appearance of concentric rings of alternating pale and dense material. As Tandler and MacCallum (1972) pointed out, similar complex secretory products have been found in a wade variety of cells in both vertebrate and invertebrate species. From an evolutionary point of view, it can be argued that apparently the sero¬ mucoid cells in the submandibular of A, phaeotis have evolved from a more primitive demikmar mucous cell. Consequently, the seromucoid secretory product in Anihetf,s phaeotis is not produced by a cell that is homologous to the serous cell of the submandibular of Erinaceus europaeas, even though the secretory products are morphologically similar. Aside from secretory granules, the two types of cells are different in most ukrasiructural aspects including those characteristics that retlect the process of synthesis of secretory product. In A. phaeotis, the extensive GER is flat and slacked, whereas in the European hedgehog it is short and greatly sw'olteii and the cisternae contain flocculent materials during the active phase of synthesis (Tandler and MacCallum, 1972). The Golgi complexes in A, phaeotis consist of flat lamellae, swollen cistemae containing small vesicles, and adjacent condensing vacuoles, whereas that of the hedgehog secretory cell primarily con¬ sists of lamellae that give rise to small saccules that in turn become condensing vacuoles (Tandler and MacCallum, 1972). Furthermore, the secretory granules in .4. phaeotis are negative to both PAS and loluidine blue, w'hereas those in the European hedgehog are PAS positive. Overall, it can be said that the striking uUrastruclural similarity between the tw'o types of secretory product is not the result of a similiar process in synthesis and possibly not a reflection of similarity in basic chemical composition. 2U SPECIAL FHJBLICATICNS MUSEUM TEXAS TECH UNIVERSITY AcKNOW I EDC. M ENTS Most financial support for the investigations reported here was from an NIH (National Institutes of Dental Research) Grant, DE 03455-02, to Phillips. Ad¬ ditional support was from a National Science Foundation Instituiuional research grant to Hofstra University (funds awarded to Phillips in 1972 and 1973), a grant from the Hofstra University HC’LAS Executive Committee, 1974 (to Phillips and Grimes), and from the Department of Biology, Hofstra University. We are especially grateful to Dr. Irving Gal insky for his support. Additionally, some specimens used in this study were obtained in Jamaica by a joint Hofstra University and Texas Tech University field expedition which was supported in part by NSF grant (GB-41105) to Robert J. Baker. We also are pleased to acknow'ledge the assistance of Dr. Bernardo Villa-R. and Ticul Alvarez-S., Departmento de C'onservacibn, Secretaria de Agricultura y Ganaderia, Mexico, and John R. Richardson, National Institutes of Health, Atlanta, in arranging for necessary collecting and importation permits. Several present and former students at Hofstra University made noteworthy contributions to various re¬ search projects summarized here. These persons, and the phases in which they were involved, are: Arnold Conrad, dental anatomy; Brett Oxberry, Barry Steinberg, Michael Titler, dental microanatomy; Paul Billeter, Sharon Butler, Laura Corio, Brenda Wilder, salivary glands; Ira Greenbaum, longues; Reena J. Harm, foreign body giant cells in oral lesions. Especially important technical assistance with such time-consuming tasks as photography, staining, and main¬ tenance of scanning and transmission electron microscopes, was provided by Joseph Adler, William Oxberry, and Perry Simons. Valuable field assistance was given by Dr. Edward Snoek and Messers. Brett Oxberry^, Paul Billeter, Paul Danker, and Michael Con fort i in association with the Department of Bi¬ ology, Hofstra University. Others who contributed materially to field work in¬ clude Dr. Stuart A. Marks, Department of Behavioral Sciences, St. Andrews College; Biol. Arturo Jimenez-G., Uni vers idad Auto noma de Nuevo Leon, Mexico; Biol. Cornelio Sanchez Hernandez, Universidad Nacional Autonoma de Mexico; Dr. Robert J. Baker, Dr. Hugh H. Genow'ays, John W. Bickham, John C. Patton, and Dr. Diiford C. Carter (and field parly), Texas Tech Uni¬ versity; and Dr. Rollin H. Baker (and field party), Michigan State University. Fluid-stored museum specimens representing many phyllostomatid genera kindly were loaned to Hofstra University by Drs. Clyde Jones and Don Whlson, Bird and Mammal Laboratories, U.S. Department of Interior (National Museum of Natural History, Smithsonian Institution). We are especially grateful to our typist, Mrs. Cecelia Sokolow'ski, Hofstra University, who w'orked so patiently with the several drafts of our w-rilings. Lastly, w^e also acknowledge both the understanding and notable assistance of our respective w'ivcs, Linda, Suzanne, and Elaine. Ll TERATURE CiTED Al l EN, G. M. I9J9. Bats. Harvard Univ. Press, Cambridge. x + 36k pp. Ai-VAREi', T. 1966. Redescubrimienio dc alguno.s tipos de murcielagos Mexicanos que sesuponian perdidos. Rev. Soc. Mexicans Hisi, Nai., 27:191-197. BJOt.OGV OF [HE PJIYLLOSTOMATIDAE 239 Amsierdam. a., 1. am> M. StusANUt. 1969. Dynamic changes in the ulira- siructure of the acinar cell of the rat parotid gland during the secretory cycle. 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Milk dentition in the broad-footed moles, Stiipunus tittitnittnix. J. Mamm., 53:354-355. -. 19726. Process of mammalian tooth development as illustrated by dental ontog¬ eny in the mole. Satpatuts laiimunus. Arch. Oral Biol., 17:61-76. ZJMMERStAN. S. 1968. Physiochemical properties of enamel and dentine. Pp, 70-91, in Dental biochemistry (E. P, Lazzari, ed.k Lea and Febiger. Philadelphia. ix-b 322 pp. ECHOLOCATION AND COMMUNICATION Edwin Gould Interpreting the sounds of members of the Phyllostomatidae offers a special challenge to the descriptive and experimental zoologist. No family of bats in the New World has radiated with such diversity in kind {137 recent species, Jones and Carter, 1976:7), food habits (fruit, nectar, pollen, blood, insects, and verte¬ brate prey), and habitat (deserts, grasslands, forests, and woodland clearings from the low'lands to the highlands) as have the Phyllostomatidae. Is the richness of their ecological adaptations matched by equally diverse options of communica¬ tion and echolocation systems? Extensive descriptive studies essential to investi¬ gating diversity should reveal clues to the variety of phyllostomalid sound systems. As many as 11 species may simultaneously occupy the same roosting site (availability of roosting sites is thought to be one of the factors that limit the pres¬ ence of bat species in a region, Tamsitt, 1967). Herein may lie the clues to se¬ lection for the distinctive communication signals that typify phyllostomatids. Diversity in terms of vocalizations will be described. Microchjropteran bats, including all of the phyllostomatids studied, orient acoustically by responding to the echoes of their own ultrasonics. Bats typically emit pulses at increasing repetition rate as they approach an object, take off, or land. Search phase, approach phase, and terminal phase have been designated as the three phases of altered pulse emission during goal-ortented flight. The sounds are high frequency, frequency modulated, and of short duration. From the echoes of their emitted pulses a bat can determine direction, distance, and velocity and some aspects of size, shape, and nature of objects (Novick, 1971), Despite the diversity of phyllostomatid food habits, the families Phyllostomatidae and Vesper- tilionidae (a family with most species having similar food habits) display similar pulse-emission patterns. Phyllostomatid bats have been used for a number of experiments in the study of echolocation. This work w'ill be reviewed briefly. The intensity of sonar calls of phyllostomatids has received considerable attention and also will be discussed. Another feature of interest in the phyllostomatids is the nose leaf and its possible function. The major aim of this review is to discuss ultrasonic vocalizations emitted by phyllostomatid bats. Most of the published literature relates to calls that function as echolocation signals, I will present some new' descriptive and functional information on the calls of young and adult bats; some of these calls in young bats are precursors of echokx;ation sounds in adults. Other calls in young and adult bats probably function as communication signals. The function of some other vocalizations is not clear; perhaps depending on circumstance, these calls serve as communication as well as echolocation signals. At the very' least, these new data will indicate a greater diversity of ultrasonic vocalizations than has hither to been described. 247 248 is PEC I Al. PUBl.lCATJONS MUSEUM TEXAS TECH UNIVERSETY Why consider vocalizations used for communication in the same discussion as vocalizations used for echolocation? Konstantinov (1973) worked w'ith Myoiis oxygmithtts and with Rhinolophits fetrufm’quimmr, he demonstrated that onto¬ genetic continuity between sounds of the same class emitted by young and old bats are communication and echolocation signals respectively. Woolf (1974) confirmed Konstantinov’s work by demonstrating a continuum of communication vocalizations emitted by infant Eptcsicas with sonar vocalizations emitted by the adult; W'oolf has referred to this continuum as a sonar family. Gould (1971) reasoned that in the course of evolution, sonar calls were nuxlificalions of already existing communication signals. The social behavior and associated vocal communication of most bats are still so poorly studied that we can only speculate on the extent to which some ultrasonic signals are used for echolocation or communication. In a preliminary examination of communication, Gould e! al. (1973) described species-specific ultrasonic comnmnication calls in five species of phyllostomatids. I will attempt to examine here some of the variability not reported on by Gould et al. Terminology The following terminology, the first six terms of which are from Strulisaker (1967), will be used in this paper. Synonyms used in other publications are in¬ cluded. Unii .—-'rhe unit is the basic element of a sound uninterrupted by periods of silence or abrupt changes in frequency. The unit is represented as a continuous tracing along the temporal (horizontal) axis of the sonogram. “Note" is a syno¬ nym. Phrase .—The phrase is a group of units separated from other similar groups by a time interv'al greater than any time interval separating the units within a phrase. [ioiif, —bout is a grouping of one or more phrases separated from other simi¬ lar groupings by a time interval greater than that separating any of the phrases within a bout. No/iiotuil unit. —A non tonal unit is composed of sound that is more or less continuously developed over a wide range of frequencies; synonyms are “noise” (Andrew, 1964) and “harsh noises” (Rowell and Hinde, 1962). Tonal unit. —A tonal unit is composed of sound characterized by one or more relatively narrow frequency bands and has been referred to as “clear calls” by Row^ell and Hinde (1962) and “sound" by Andrew' (1964), Units with a multi¬ harmonic structure are included in this category. Conipoutul unit .-—A compound unit is composed of both nomonal and tonal sounds that appear as a sequentially continuous tracing on a sonogram. Mixed unit .—Units composed of both tonal and nontonal sounds that occur superimposed (simultaneously) on one another are called mixed units. The tonal and nontonal aspects are more or less separated by differences in frequency. Is(7la!ion call |/-ca]l (Gould, 1971)].^—The /-call is a tonal unit with nearly constant frequency with a duration of about 20 to 60 milliseconds (msec.). BJOLOGY OF THE PHYLLOSTOMAT[DAE 249 Synonyms are Stimmfuhlungslaute (Kulzer, 1962), attractive pulse calls (Kon¬ stantinov, 1973), and Veriassenheitslaut (Schmidt, 1972). FM pulse. —Woolf (1974) described FM pulses as tonal calls that are ‘Tiiono- tonically decreasing, frequency modulated vocalizations which sw'eep through roughly one octave.” The sweep is not linear and has a duration of about 1 to 7 milliseconds. It appears that some infant bats may emit FM pulses that function mainly for communication. Thus, terms that refer exclusively to physical char¬ acteristics of the calls will prevent any premature and prejudicial designation as to function. Fhe FM pulse frequently has been referred to as an echolocation signal or sonar call by Griffin (1958) and throughout the extensive literature on bat echolocation; Konstantinov (1973) used the term “location signals.” Oc¬ casionally, monotonic frequency modulated vocalizations increase in frequency; these are designated as an ascending FM pulses. FM glide (FMG),—The FM glide is a single unit, frequency modulated pulse that sweeps about one octave and for whiich frequency is held more or less con¬ stant at the beginning (FMGB) or at the end (FMGE). Mixed pulse (Suthers, 1965) is a synonym, but this term is avoided because of its inconsistency with Struhsaker’s reference to an unrelated term, “mixed unit” (see above). Double m>ie (Gould ei ctL, 1973; Woolf, 1974).—DN is identified by the close temporal association betw een tw o notes, a long and a short call regardless of their order. Repetition rate of DNS (plural of DN), even in bouts, is usually less than that of FM pulses. The DN varies widely, particularly in regard to the number of notes; a long-short call may be follow'ed by two FM pulses, and, sometimes, one of the units of the DN is repeated once or twhee (thus the designation “and higher multiples”). Rarely do the notes number more than four in a phrase. Verlas- senheitslauten (Schmidt, 1972) is a synonym, \^^arhle .—-A short, single tonal unit in which frequency rises and falls two or more times. Infants separated from their mothers for 15 minutes or longer were placed 20 or more centimeters away from their mothers. From the time of placement to the time that the infant attached to the mothers nipple is a reunion. Conmci- —The moment any portion of the mother or infant’s body touched the other is the moment of contact. Methods Bats were collected from the following localities: Lepionycierts sanborui, Sonora, Mexico; Macrolus californicus, southern Arizona; Carotlia perspicillata, Trinidad; Phyllosiomiis hasiatuSy Trinidad; Desmodus romnduSy Costa Rica; and ArnheuSy Mexico. Sample size and ages of bats used for recordings are sum¬ marized in Table 1. LeptonycierlSy MacroruSy DesmoduSy and Artibeus w'ere kept in darkened cages that measured 23 by 23 by 37 centimeters. Tsvo Ariibeus in healthy condition (judged by their vigor and the condition of their pelage), but abandoned by their mothers, were received from Roy Horst, but they failed to survive more than two weeks. Macrolus were maintained on glop, mealworms, and crickets. Lepio- 250 SPFX’lAl- HUBlJCAT[ONS MUSEUM TEXAS TECH UNIVERSITY TAHi.b L— Siitnple size atul of Imus usetl for riionlitif^^s. Aye in finnthcr of ilays ix followcii by stonpU' size in pureniheses i/yreuttr than one. R iilemifiex recordings t)/ renniotix', Atf Oil nils. Dc^nuuftt': Li'/mmyiu-rh PhyUiofu/tiii', MtariHiis Curtflhti .Artihi'u:* 1 (2) 1 (2) R 1 (2) R 1 (2) R I (2) 13(2) 20-30(2)* 4 2 2 2 R 2 16 R 5 R 4 R R 7 7 17 6 5 R 4 11 R 4 19 7 7 7 R 9 5 20 R 9 S R IS Ad (2) 7 R 21 16 R 9 45 R 9 24 17 15 R Ad (2) 10 25 20 41 6.‘5 ISO Ad (2) Ad (2) Ad (4) 40 * Age in days is approjftmate. nycieriSy Camilla, and Phyllosiomu.s maintained on diets identical, or similar, to those described by Rasweiler (1973). Desmoiius were fed fresh and frozen beef blood. Tw'o hand-raised infants (from 2 and 7 days old) were fed Esbilac (Taylor ef ai, 1974), The Phyih.^R}miiS infants used for sound recordings lived more than 10 months and reached adult size. Despite long periods in a 77 by 56 by 52-centimeter cage, they could fly w'ell and high (7 meters) in a 18.3-meter geodesic dome. Lc'pto- nyaeris and Macroiu.'i were released in the wild after the observations w'ere complete. The Lf’pionyaeris flew vigorously, and the Macrotus (infants 15 to 21 days of age and their mothers) were released in apparently healthy condition at the place of their capture, a concrete chamber beneath a bridge. Most of the recordings of Caroiiia were obtained from D. Kleiman’s colony at the National Zoological Park in Washington, D.C. T wo reunions of 7 and 16- day-oid Caroiiia were observed and recorded in the environmental chamber. The entire colony was removed from the chamber. An infant was removed from its mother’s nipple. In an isolated room, sounds of mother and infant w'ere recorded separately under various circumstances. Then the infant was placed, facing a microphone, on a slender, horizontal branch near the center of the dimly lighted chamber. The microphone was placed 7.5 cm from the infant and behind the branch. With an assistant 1 watched the reunion through the chamber window. The mother was released in the chamber. Her attempts to approach the infant from the rear were thwarted by baffles placed behind the microphone. Thus, whenever the mother attempted to reunite w'ith her infant, she Hew tow'ard the microphone. The moment the mother landed next to the infant w-as obvious be¬ cause her claws scratching the branch were clearly audible on playback. Scratch¬ ing w'as also recorded as the mother took off w ith the infant attached to her nipple. BIOLOGY OF THE PHYLLOSTOMATIDAE 25 J Observations of reunions of other species were conducted w ith the use of cages equal in size to those in which the bats w-ere maintained (see above). An infant bat was removed from its ntother, often by detaching it from a nipple. The mother w'as placed in another cage; then the sounds of the isolated infant w'ere recorded. The door of the mother’s cage was opened; the infant w'as placed as far from the mother as possible Lmd the dtxir was shut. A microphone was fastened inside the cage and oriented so as to optimize detection of calls from both mother and infant. My commentary on mother-infant behavior was tape-recorded on one channel of a Precision Instrument tape recorder at 76.2 centimeters per second; the bats’ sounds W'ere recorded on the other channel (see Gould, 1971, for details). Timing of the reunion began w hen the infant w'as introduced into the cage and it continued until body contact between mother and infant occurred. I used a dim red light to observe reunions. Sounds emitted by precocial new-born bats often closely resembled calls emitted by adults. To facilitate my ability to discriminate the calls of mother and infant during reunions, I first recorded the sounds of isolated mothers and infants. Later, w'hen I analyzed recordings, I listened to the slowed recordings of isolated in¬ dividuals and then listened to the recordings of reunions. Experimental Studies of Echolocation Investigators have demonstrated the skillful ability of bats to avoid obstacles in a rather standardized experimental design utilizing evenly spaced vertical wires. The bats’ skills are about equal regardless of taxonomic position (Grinnell, 1970). Obstacle avoidance ability is impaired when ears are plugged or the bat’s voice is altered by cutting the motor branch of the superior laryngeal nerves, The ability of leaf-nosed bats to avoid a plane of vertical w'ires by means of echolocation has been demonstrated several times. Grunimon and Novick (1963) demonstrated that Macroms could avoid wires down to 0.27 millimeter in diameter and could even avoid wires 0.19 millimeter in diameter at a rate better than that expected by chance alone. Their unprecedented large sample indicated considerable variability of individual performance. Macroms, Carol!la, and Arftbeiis are equally adept at avoidance (see also Griffin and Novick, 1955), Glossophaga performed better than did Macrotus, CarolUa, and Artiheus by scoring 89 per cent misses when avoiding w'ires 0.175 millimeter in diameter. Howelf (1974) has evaluated dental and skeletal morphology and food habits in combination with obstacle avoidance behavior in selected bats {Glossophaga S(}ricina, Anotira gcoffroyi, Lcpionycfcris sanhorni^ Choeronycteris mexicana) that feed on nectar, pollen, and insects. The ability of Glossophaga soricina and Anoitra geoffroyi (wild-caught specimens of which How-ell reported to contain 46 and 90 per cent insects, respectively) to avoid w-ires 0.28 millimeter in diam¬ eter was superior to that of Lepamycteris safihijrni and Cfunronyaeris mcxica/ia (w-hich Howell found rarely to eat insects). Vampyrum spec!nan is capable of using echolocation to make rather difficult discriminations between targets of different shape or size (Bradbury, 1970). This species could detect and select one of two lucitc targets at distances be- 252 SPECIA]. PUBL[CAT(ONS MUSEUM TEXAS TECH UNIVERSITY tween 50 and 150 centimeters. All pulses emitted on discrimination flights were of the so-called FM variety. There was a steady decrease in pulse amplitude and an increase in the rate of frequency modulation as targets w-ere approached. Bradbury (1970) concluded that one individual had used frequency dependence of the echoes to effect the discriminations whereas another had used the overall amplitude cues* This suggests that more than one option of information proces¬ sing is open to bats. Perhaps in the course of ontogeny an individual bat would learn to favor one particular system over another. The variable repertoire of echolocati%^e pulses implies that these bats may use different types of pulses to extract different kinds of information about targets (Bradbury, 1970). Different species of bats frequent different habitats, feed on different foods and echolocate by means of different types of orientation signals. The orientation sounds differ in intensity, duration, frequency, pattern of frequency change, and repetition rate. Sounds that differ in such fundamental w ays presumably code information about target location and properties in fundamentally different forms. One might therefore expect to find significant differences in the mechanisms of neuroprocessing used to extract this information {Grinnell, 1970). Various neuromu-scular .systems that demonstrate a bat's ability to respond to echoes of its own emiued ultrasonic pulses have been described (Simmons, 1973; Grinnell, 1973; Griffin, 1973; Suga and Schlegel, 1973). Great specificity of the nervous system seems to be a prime characteristic of the bat’s auditory nervous system. Echoes of a particular intensity are probably processed optimally (Novick, 1973), A bat’s nervous system is sharply tuned to specific frequencies, intensities, and temporal relationships of stimuli. In studies of the comparative auditory physiology of neotropical bats, Grinnell (1970) found that Phyiltfsto/fius kin6 Cu/Yj/Z/f/(as well as three nonphyllostomatid bats) have auditory systems that are most sensitive approximately in the range of the bat’s emitted frequencies. He found a correspondence between audiograms and emitted frequencies, implying a specialization of the cochlea and neural ap¬ paratus for analysis of the same narrow band of frequencies used in emitted pulses. Carollia showed auditory responses to .sounds of 140 to 150 kilohertz (kHz), a level higher than that found with any consistency in other species studied. One of the five Carollia differed from others in show'ing approximately equal sen¬ sitivity to all frequencies between 30 and 120 kHz. Vernon and Peterson (1966) found that Dcstnodus has maximum cochlear sensitivity at frequencies (50-70 kHz) higher than those in their echo locating pulses (42-24 kHz) as reported by Novick (1963). Removal of the ear pinna and tragus had little effect on cochlear sensitivity (Vernon and Peterson, 1966). Howell (1974) speculated on the differences in cochlear potentials of Choero- nycte/is and Camllia compared to other phyllostomatids; she suggested that her observations support the notion of polyphyletic origins for nectar feeders. Grinnell (1970) described an '‘off’ respon.se in the Mormoopidae that might help in detection and distance determination by bats employing long or constant wave length signals. The “off’ response was prominent in Pieronatus (= Chilo- nycferis), as well as in the emballonurid Sewcopferyx, present but of higher thresh- BIOLOGY OH THE PHYl.LOSlOMATIDAE 2^3 old in Pserorwtii^^ {sensii strkiit)^ and essentially absent in Phyilostifnius and Caroiiia, as it is the case in Myatts and Plccotus. The effects on sound emission of unilateral and bilateral sections of the motor branch of the superior laryngeal nerves of several phyllostomalid bats {CaroUia, Phyliosiomits^ and GU}^sophaga) and some vespertilionids were similar (Novick and Griffin, 1961). Unilateral section resulted in lowering of sound pulse fre¬ quency; bilateral section resulted in additional lowering of frequency as well as disappearance of FM. Novick and Griffin (1961) concluded that in the Vesperti- lionidae and Phyllosiomatidae, and probably in the Nociilionidae, Enibal- lonuridae, and Molossidae, orientation pulses are produced by tensing special iaryngeal membranes w-ith the cricothyroid muscles under vagal motor inner¬ vation via the muscular branch of the superior laryngeal nerve. Suthers and Fattu (1973) concluded that the cricothyroid muscle contracts just prior to each ultra¬ sonic vocalization and relaxes during phonation. Cricothyroid muscle relaxation may gradually decrease the tension on the membranes and create the down¬ ward frequency sw'eep characteristic of most pulses. Sound with Two Components Vampyrtim spectruni emitted pulses with two components during discrimina¬ tion experiments (Bradbury, 1970). One component was alw'ays higher in fre¬ quency than the other; both components swept simultaneously to lower values. In genera] the upper sw'eep began around 110 to 115 kHz and terminated around 80 to 85; the lower sweep started at about 95 to 100 kHz and dropped to about 65. Sonograms of FM pulses emitted by Vampynmi (Bradbury, 1970), adult Phyllostomus {¥\g, IN), one-day-old Desmodus IG, H) and MflCTO/nA’(Fig. IP) show' similar characteristics: they reveal two different but simultaneously emitted sounds. One interpretation is that the components are related by a com¬ mon submultiple. If the two components are not harmonically related (and this appears to be the case), they may derive from two different vibrating systems. Perhaps asymmetrical or unequal tension on the two vocal membranes of the larynx accounts for the emission of the tw'o distinct but simultaneously emitted sounds. Perhaps this call occurs in Desmodus only during a brief developmental stage in sonar ontogeny, a stage in w-hich the immature animal has yet to achieve complete bilateral coordination of the larynx. The phenomenon of two acoustical sources existing in the avian syrinx w'as described by Greenew'alt (1968). This possibility in bat vocalizations requires more investigation. The Nose Leaf Caroiiki, and presumably other species of Phyllostomatidae, differ from the typical pattern found in the Vespertilionidae by being able to emit sound through either the mouth or nostrils. In addition, phyllostomatids typically scan their surroundings by extremely active movements of their external ears. Experiments with the nose leaf of Hipposideros and Rhinoiophtis have revealed that altering nose shape changes directional qualities as well as amplitude char¬ acteristics of sonar calls (Novick, 1958). How-ever, Grummon and Novick (1963) 254 SPECIAL PUliLiCATlONS MUSEUM TEXAS TECH UNIVERSITY -A , iV , . D 98- : i 64 32- 0 a _ 98 I 64- % 0 3 O’ £ 4 Ik 96- 6? 2; LiKluAnMMkAikkJki k ^ 1 >1 t ^ ' ' H I 4 > I 111 . ; ' I’ * iiii ' V i jv*!' .y , j y f, .,^%. ^ ^ 0 p 96 64 32 0 K L K N t ) 1 ' ' h 1 *1 ,Vi t I I V 1 , ' I ; ' \i V V ^ ^ ^ \ \ ^ ^ ^ \ 0 ■ ' (J> mmic. Time Fid. I.—Ultrasonic calls of five genera of phyllostomaiid bats. Hashy dark areas al the bottom of most sonograms and to about 32 kHz and 9b kHz in some are due to background noise. A. Five FM pulses of a lb-day-old Otroflia during reunion. Note the prominence of what is probably ihe fundamental starting at about 64 kHz and dropping to about 32 kHz. B and C. Three FM pulses of mother and seven-day-old CttroUkv, the FM pulse on the left in each case is the last unit of a DN; the next two to the right are the mother's FM pulses. What appear to be second and third harmonics are more prominent than pulses shown in A. D. Four FM pulses emitted by a four-duy-old Lepro/iyrter/v during a reunion; its mother was quiet. Mother and infant were at opposite ends of the cage and the microphone w as pointed directly at the infant. E. Three FMGE emitted by adult Lep/onyt ter A during reunion w ith its four-day-old infant. F. G, H, and I. FM pulses of a one-day-old Desmoditx exploring a bo.x. Note two components that may not be harmonically related in G, H, and 1. J and K. FM pulses from a 41-day-old Dextncnlux that had been hand raised from the time it was seven days old. Note what appear to he harmonically unrelated conponents in K. Recorded while the bat was held in hand. L. F.M pulses emitted by llO-day-okI Dismodus during four meters of flight; it maintained altitude and climbed slightly. .M. FM pulses of 65-day- old Di’xmoiiu.'i walking on substrate. N. FM pulses of adult Phylluxnfuiiis flying in 18.3- meter geodesic dome cage. Intervals of eight pulses averaged 107 msec. (100-110 msec.); the example is indistinguishable from many others recorded under same circumstances. P and Q. Nineteen consecutive sounds emitted by a one-day-old Macrofnx. The last two (Q) are a DN. Notice the second component that appears as a vertical line just below the beginning of fundamental (about 16 kHz). RIOLOGV OF THE PHYLLOSTOMATIDAE 155 failed to link the function of the nose leaf in with echolocation. Likewise, ccholocation in Carol!la is not effected by amputation of the nose leaf (Griffin and Novick, 1955), Thus, the function of the nose leaf in the Phyllostomatidae remains obscure. Whispering Bats Leaf-nosed bats have a curious behavior that makes their sounds difficult to study; they characteristically emit low' intensity FM pulses. Griffin (1958) first characterized FM pulses of certain phyllostomalid bats as “whispered” pulses. Among the species of whispering bats, he recognized Glossophaga soricoja^ Aniheus Jama ken Uroderma hihha/um, Desmodus roiunduSy and Carol lia perspk'illata. Griffin estimated the intensity of these w-hispered pulses at about 3 to 5 dynes/cm- or about 100 to 1000 fold less intense than Myotis FM pulses. When the mouths of three Carollia were tightly sealed, the pulses were either quite normal or slightly reduced in intensity (Griffin, 1958:248); when their nostrils w'ere covered, tw'o Carollia emitted pulses. Sealing the nostrils or the mouth of Carollia had little effect on vocal emission. ''Carollia (and presumably others of the Phyllostomatidae) thus differ from the typical [species of ] Ves- pertilionidae in being able to emit sound through either the mouth or the nostrils as well as in the frequent scanning movements of their external ears” (Griffin, 1958). Their very short pulse durations set the phyllostomatids apart from the horseshoe bats, which emit long duration sounds through the nose. Probably the methods of phyllostomatid echolocation are more like those of the Vespertilioni- dae. A number of investigators, including Griffin, have noted that not all phyllosto' matid pulses are “w'hispered.” Desmodus rotundns on some occasions emits fairly intense 2 or 3-msec, pulses having a constant frequency between 20 and 30 kHz (Griffin, 1958). Bradbury (1970) in Trinidad recorded pulses from yampyrnm spectrum that differed from pulses recorded during discrimination tests in the laboratory in the United States. The unique pulses were longer and louder than the discrimination pulses and, based on a fundamental of 20 kHz, included a complete harmonic series up to the seventh harmonic. In one such record, the bat shifted in one jump from the long harmonic pulses to the short discrimination ones. The shift in one jump and the interval of about 20 msec, between units (Bradbury, 1970, fig. 6) are similar to DNS described below. Sealing the nostrils of Macrotus limits vocal output to abnormally long duration pulses, possibly cries of emotion (Grummon and Novick, 1963). Sealing the mouth did not .seem to affect the duration of recorded pulses; however, pulses tended to be at higher amplitude than normal. In both cases, the mean frequency was unchanged from that of the normal bat. But bats with nostrils blocked were unable to orient in an obstacle course. In Macrotus, nasal emission appears neces¬ sary for acoustic orientation. Grummon and Novick’s (1963) observations of Macrotus suggest that high intensity calls are emitted via the nostrils. These ex¬ amples of high intensity sound emission provide little information about function. 256 SPRCLAL PUlil.lCATlONS MUSEUM ] EXAS TECH UNIVERSITY However, Howell (1^74) has presented evidence that some phyllostomatkls con¬ trol amplitude during obstacle avoidance. During analysis of ultrasonic vocalization of six phyllostomatid species 1 have noted frequent cases of bats emitting calls of much higher intensity than whis¬ pered EM pulses. Some of these high intensity calls were FM pulses. Others ob¬ viously were not and w ill be described below. High intensity calls were recorded from Lcpiouycteris, Carollku Phyiiosfimuis, Desmodus, and Macrotits (Fig. 2). In some cases, these calls were detected only in the infants; in others, the calls w'ere obtained from adults as w-ell (see details that follow'). Decibel differences between the two calls were measured in three to five bouts; Curoliia, 15-22 decibels (db); Artibetts^ 8-12; Lepumycteris, 8-12; Phyilostomiis, 12-14. Annotated List of Ultrasonic Vocalizations Emitted HY Ado).t and Infant Phyllostomatid Bats Although Howell (1974) and Novick (1963) have speculated on the potential interspecific variability in FM pulses of the Phyllostomaiidae, only sparse data support their suggestions. For example, Novick (1963) slated that Desmodits pulses (FM) are, indeed, not describably different from most of the Phyllosto- malidae such as Artibeits or MacroUts'' Novick (1963) urged caution in the in¬ terpretation of earlier data from Griffin and Novick (1955) on PhyUosiomits^ Umchorhhui, and MacrophyUufu, Griffin’s and Novick’s (1955) evaluation should probably be considerably revised in view of technically more successful recordings show'ing a frequency modulated, harmonic pattern found generally among the Phyllostomatidae (Novick, 1963). Howell (1974) noted that all genera studied could emit calls at an intensity of approximately 2.5 dynes/cm-. As the wire diameters decreased in an obstacle avoidance experiment, GUnsophaga, Anowa, and Chocronycterh increased their pulse amplitude to 5, 4, and 4 dynes/cm-, respectively. Specimens of Lepto- nycteris did not increase the amplitude of their cries beyond 2.5 dynes/cm-. Howell did observe, however, that lj.'ptonycteris emitted pulses of longer dura¬ tion than did the three other phyllosiomalids studied. This verified Novick’s (1963) observation; Novick also noted that the frequency pattern was similar to that of Glossophaga and Lonchophytta. In some of the sonographs, the ability of phyllostomatids to emit long or short FM pulses is apparent (Figs. 4F, 4G). Diversity of recording situation has repeatedly been mentioned as a key factor in determining the characteristics of FM pulses. The review of species bekw (Fig. 3) as w'cll as Fig. 1 seem to indicate that all phyllostomatids emit brief, low' intensity, multiharmonic FM pulses. There is little indication of a correlation between the ability to emit higher frequencies and size of the bat. The highest frequencies detected in leaf-nosed bats are from Giossophaga\ but Vampyrutu specintm, the largest member of the family, can emit sounds above 100 kHz, which is somewhat higher than sounds emitted by other species (Fig. 3). Similarly, in birds there is no consistent relationship between frequency in the song and size of the bird. Large birds such as the osprey and Laysan albatross sing at high fre¬ quency whereas the morning dove and several small owls have deep voices (Greene- walt, 1968). BlOl.OGY OF THE PHYLLOSTOMATIDAE 2^7 *96 ^64 01 ^32 Ol £ a „S( mA iiliMiri ' } 211 to “l— 60 Time ^MSECI LOO Fig. 2.—A-F. OscilJographic records of DNS and FM pulses emitted in the same bout of vocalizations. Long duration, high intensity units are DNS: shori duration, low intensity units are FM pulses. Upper lines of dots represent a lime base of 60 dots per second. Record speed was 30 ips (inches per second); reproduce speed, I % ips. A. about 14+ days of age. B and C. Lepto/iycferis, 1 days old; pulses in C occurred about 400 msec, after those in B. D. Phyiktsiomus, 1 days old. E and F, CuroUUi, 1 days old. G. Sono- graph of the five units shown in F; these include a DN of three units, one long and two short follow'ed by two FM pulses that are indistinguishable from FM pulses of a flying adult. Note the intensity difference between longer unit.s (DNS) and shorter units (FM pulses). Listening to slowed recordings of CitroUuu I heard several hundreds of bouts containing DNS and FM pulses. Species differences in intensity, duration, and frequency probably have to be worked out in the context of obstacle avoidance experiments. Bradbury’s (1970) description of the sounds made by Vampymm spectrum is the only de¬ tailed study of the variation in FM pulses emitted during different phases of echolocation. The lack of data in this area is because studies of that sort are tedious. The following descriptions of vocalizations include data obtained from re¬ cordings of six species of bats during several experimental situations in which bats were: reunited (mother and infant), hand-held, isolated and exploring a strange box, walking on the floor, and flying in a geodesic dome. Nearly all of the information on FM pulses is taken from Novick (1963) and Pye (1967); my recordings of FM pulses generally corroborated their findings. Macroius californicus FM pulses (Fig. IP).—^Duration, 1.7 to 3.9 msec.; frequency fundamental beginning at 35 to 40 kHz and ending at 26 to 30 kHz; second harmonic beginning at 78 kHz and ending at 34 kHz. When the fundamental frequency dropped below 34 kHz (34-29), a third harmonic appeared and swept from 102 to 81 kHz. Calls resemble those of Artibeus and (Novick, 1963). 25S SPECiAL PUBLICATEONS MUSEUM TEXAS TECH UNIVERSITY INCREASING SIZE — ♦ Fiti, 3.—Frequencies of FM pulses of 16 difl'erenl genera of phyllostomatid bais. Data were obtained from Novick (1963), Pye (1967), and this study. The figure is somewhat over¬ simplified because there is no indication as to which harmonic or harmonics contain the greatest energy. Vertical lines indicate the range of frequencies' occurring in FM pulses. Broken lines indicate the harmonic structure; the lower line may be the fundamental, and the lines above represent the second and third harmonics. For example, in E the fundamental is about 40 to 30 kHz, the second harmonic about 80 to 40 kHz. and the third harmonic about 112 to 80 kHz. The numbers adjacent to the letters are the approximate weights in grams of the adult bat.s, and the species represented are; A, Clossophaga longirostiis, Ghs- sophanii soricitui, and Anoura }*i’tiffroyt B, Ariibeus iinereua', C, Vampyrops heUeri\ D, Miuroiits codfornk'HS', E, CaroUkt perspiciUafir, F, Centurio .vene.r; G, Siurnlra Uihun and S. lihitie", H, Lcptofiyiteris nivatis' 1, Phyliodtrimi xtetuips', J, Destnodus rotitiidiis^ K, Chirtidermu vsUoxunr, L, Artihvus januikensix {Pyc, 1967, listed the second harmonic as 65 to 42 kHz and the third harmonic as 92 to 55 kHz; Novick, 1963 reported the primary component to be 49 to 56 kHz falling to 32 and the second harmonic at 104 to 64 kHz); M, Ariiheus litHratu-% N, Phyliossotnus huxuitus and Phylloxiomus discolor, O, Vompyntm spectnon. The data have been arranged into three size groups in order to point out the lack of correlation between weight of the adult and frequency of emitted pulses. DN (Figs. IQ, 4T).—^Duration of phrase, 61.5 msec. (43-91.5, 10); fre- cjuency beginning at 13 to 32 kHz, with a middle of 7 to 11 kHz, and an ending, 11 to 32 kHz. DNS were emitted by two isolated infants (one and 11 days old) and adults; three adults emitted DNS during reunion with their offspring. Care* BIOLOGY OF THE PHYLLOSTOMATIDAE 359 A, m’ B4 -'s \ \i* D| . \ 96~ “iN '«<- 32-S to M* it*.- h \. I A ''’■ \ ■■S 1 ^' < H I 1 j 1 9« 64 1 52 * 'w 'S L M N Ml ' nS^>^ A ' * ■ Vl»' S, V- 4 ' * iJo ' li) T \ V V V \i k TIm Fig. 4 .—DNS of five genera of bats. A and B. DNS of (he same specimen of Ariibeus, an infant of about 14 + days of age. C, D, E, F, G. Sonograms of Oirallia DNS- C, D, F, and G are from the same 7-day*otd infant described under "profiles" in text; E, 16 day-old infant described under “profiles" in text. Note similarity between D and E, Call in D was emitted while the mother was roosting quietly. Call in E was emitted near the moment of bodily contact of mother and infant. C, D, F, and G show calls that were emitted w hile mother was roosting quietly. H, I, J, K, L, M. N. Sonograms of Dt’s/nodus DNS arranged to show order of increasing complexity: H, seven-day-old; 1 and J, 5-day-old infant; K, one-day-old; L, seven-day-old; M, 20-day-old; N, 20-day-old. P. Q, R. Sonograms of an isolated specimen of Phynoxtomus: P and Q, one-day-old; R, aroused 45-day-old, isolated bat. The latter sound¬ ed tike a DN in a mixed unit; compare with Fig. 7A (65-day-old Dexmodus). S. Sonogram of an aroused four-day-old DextHodiis that had been hand-raised from day one. Recorded on a Uher tape recorder at 7 Pi ips and slowed to I ips. T. Sonograms of adult Mucrotus responding to its calling one-day-old infant. At the time of this recording, the infant was removed from the range of the microphone. Record and reproduce W'as the same as in S. 260 SPEClAl, HUHt.lCATJONS MUSEUM TEXAS TECH UNIVERSITV K 96 ’ 64 e Dt E| F 16 - G tt ; 4 - \ ' I'P ' II in |o M' -II c K Time M w. Fig. 5,- — Selecied sonograms showing short duration units that result from omission.s in long units; Woolf (1974) has referred to this continuum as a "sonar family." A, B, C, D. Calls of the same isolated one-day-old Oitollia. Compare the second unit (from the left) in A with the second unit in B w ith the first and second units in C and with the first and second units of D. This scries shows breaking up of a longer unit into componems. The ter¬ minal portion of the second unit in A resembles the fundamental of the FM pulse, C is a typical DN with three units; a long, a short, and after a longer interval a second short note. Eand F. DNS emitted by an isolated 13-day-old CurttfUa. Compare sonograms in A and E. These are recordings from different bats. F appears to be a variant of the Iasi three units of E. Dark hashing between first and second units of E is caused by background noise: background noise produced similar effects on other sonograms. G, i'-call emitted by one-day-old ftycicris, H. DN emitted by same bat as in G; note the FM features of the second unit. L DN emitted by cight-day-old Leptonyittri.',. J, K, L. M. Calls of an adult Macrotus just after its vocalizing from the recording area. Compare the frequency contours of the units in K, L. and M w ith those of the single unit in J. ful Study of this call may reveal that its units derive from a division of a single unit (Fig. 5J-L), Sound frequency of DNS dropped at about the same rate as the sweep of FM pulses; it then rose to slightly more or slightly less than the frequency at which it started. Bouts usually contained 2 to 6 units. Infant and adult Despiodus emit a similar call (Fig. 4S). Warhic (Fig. 5J).—Duration 62.9 msec. (57.5-70,0); frequency 16, 6, 32, and 24 kHz (measured at four extreme points). Although this call is long and modula¬ ted, it does not fail in the higher frequency range typical of the warble of Des/uodus and Lc^ptonyetpris, This call seems to have two continuous but distinctive com¬ ponents; it begins with an intermediate frequency, drops to about 6 kHz, and rises to about 32. It w'as recorded from adults during reunion with infants in the same circumstances described above for DNS. Sometimes only the low fre¬ quency portion, other times only the high frequency portion was emitted. Other variants are a two and three-part call as shown in Fig. 5 K-M; these seem to re¬ sult from partial deletions of the complete call. BIOLOGY OF THE PHYLLOSTOMATIDAE 261 C ami a a persp i cill a ta FM pulses (Figs. lA-C).—Duration, 0.5-1.0 msec. (Pye, t967; my record¬ ings); frequency beginning at 80 kHz (Pye, 1973); 112 kHz (this study) and ending at 55 kHz (Pye, 1973); 80 kHz (this study). Recordings of FM pulses were obtained from two mother Carollia as they flew toward their infants (and the microphone). FM pulses were also emitted by a seven and a 16-day-old in¬ fant of CamlUa when their mothers were flying in the room or near them (see details below). Some FM pulses of young bats (a 17-day-old for example) had their primary energy concentrated between about 24 and 40 kHz. Other young bats emitted calls that closely resembled those of adults (Fig. 1). The funda¬ mental sweeps from about 48 to 24 kHz, the second harmonic from about 80-48 kHz, and the third harmonic from about 112 to 80 kHz. Usually the greatest energy is in the second and third harmonics, DN (Fig. 4C-G, 5 A-D).—Duration of phrase, 65.5 msec (13,5-90, A/= 14); frequency beginning at 16 to 24 kHz and ending at 40 to 66 kHz. DNS were de¬ tected from isolated Carollku one to 24 days old (Fig. 4C-G). Calls from a one-day and two 13-day-old bats w'ere very similar; most had three units. The three units have a quality like that of a bird song when slowed 16 times; their sonograms are as complex as the sonograms of a Connecticut w'arbler's song (.see sonograph in Greenewalt, 1968, fig. 34). The variability of the call may be seen in Fig, 5A-D. The sequence of four calls in Fig. 5 indicates the way in which a single unit breaks up into units that resemble DNS and FM pulses, FMGB itmi FMGE (Fig. 4C, F).—On three occasions, a 16-day-old Camdiu emitted DNS composed of FMGE when its mother flew close to it. On one of these occasions, this ! 6-day-old CaroHia was emitting FMGE; the approaching mother emitted one FMGE a fraction of a millisecond after the FMGE of the infant. The adult’s call sounded quite different from the infant’s. Sometimes DNS were composed of FMGB or FMGE. Frequently, FMGE w^as emitted just before or Just after DNS; FMGE then graded into FM pulses. It was also emitted in pairs by a seven-day-old bat that was recorded during a reunion. FMGE is usually of much higher intensity than FM pulses. Buzz.—Carollia rarely emitted this call. However, good recordings were ob¬ tained from one 19-day-old bat that became very aroused during handling. The buzz occurred as a mixed and as a compound unit in association with FM pulses and FMGE, Lepiofiycferts sauborni FM pulses (Fig. ID).^—Duration, 2.0 to 7.9 msec,; frequency beginning at about 58 and 100 kHz (third and fourth harmonics) and ending, at 25 and 50 kHz (third and fourth harmonics); fundamental, 12 kHz. Sound emission appears to be nasal (Novick, 1963). The dominant frequency sweep begins at 58 kHz and drops to 25 kHz. Series of harmonics with one or tw'o harmonics predominate, FM pulses have a longer duration than in Glossopfiaga and Louchophylla. Both Novick (1963) and Howell (1974) were impressed with the variable duration of 262 SPECiAl. PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY pulses emitted by Lepicmycteris nivalis and L sanhorni (2 to 8 msec, Howell, 1974). DM—Duration of phrase, 64 msec. (53-79, M=10; see Fig. 9); frequency beginning at 8 to 30 kHz and ending at 24 to 66 kHz{M=9), isolation calls break up into DNS as in Eptesicns (\\ooU\ 1974). Thus, frequency contours of DNS are similar to those of /-calls. Note that the lime from the beginning of the first to the end of the second pulse in the DN shown in Fig. 5G and 5H is roughly the duration of an /-call. This is typical of DNS emitted by Lt’ptonyi tens, On occasion the /-call breaks up into three notes and contains a short, slightly more modulated first note. Isolation call (Fig. 5GJ.—Duration, 64.6 msec, (55.5-77.5, M—10) emitted by one and four-day-old infants; frequency, 14 to 25 kHz. Compared to FM pulses, /-calls have relatively constant frequency. Isolation calls were recorded from infants one to four days old that w-ere hand-held and from infants that had recently been placed in a cage w ith the mother after 30 to 60 minutes of isolation. FMGHancI FMGE .—This call w-as described briefly by Novick (1963) as part of the adult repertoire; the constant frequency portion lasted I to 2 msec. An FMGE is shown in Fig, 1E. Warble (Fig. 6B, D, F),—^Duration, 70,6 msec. (52-1 12.5, M = 5); frequency, 70 to 48, 17 to 40, 48 to 50, 19 to 32, 48 to 64, and 28 to 40 kHz. Frequency was measured at six extreme points on five sonograms of adult calls. Four sonograms of calls from an 8-day-old bat measured 35 to 40, 16, and 32 to 64 kHz at three extreme points. This call was detected during close contact of mother and infant (one four days old and another eight) after a separation of 20 to 60 minutes. In some instances, a rather stereotyped call sounded like an unsuccessful attempt by infants (one, eight, and 15 days old) to emit a warble (Fig. 6D, F); two units are separated by a change in frequency rather than by a silent period as typified by a DN. Destnodiis roinmlns FM pulses (Fig. IF-M),—Duration, 1.1-2.3 msec.; frequency beginning at 38 to 42 kHz and ending at 24 to 29 kHz. Vampires appear to emit their orienta¬ tion sounds nasally. Second harmonic at high amplitude: 76 to 83 kHz (Novick, 1963; Pye, 1967). “ The beginning may typically have two component frequencies, 40 and 80 kHz, the middle three, 30, 60, and 90 kc and the end three, 27, 54, and 81 kc” (Novick, 1963). FM pulses of Desniodus are not describably different from those of other phyllostomatids, such as Macrotus or Artiheus (Novick, 1963). DN (Fig, 4H-N, S).—^Duration of phrase, 83 msec. (45.5-146.5, M—10); frequency fundamental usually ranged from 16 to 32 kHz, beginning at 10 to 16 kHz and ending at 32 to 34 kHz (M= 10). Harmonics are apparent in most of the sonograms. Desnwdus DNS seem to possess more variation wdthin individuals than I could detect in DNS of other species. DNS were emitted when Desniodus infants w-ere hand-held, during reunion w'ith the mother, and w'hen isolated. The quality of many DNS was to my ear indistinguishable from that of Artiheus when BIOLOGY OF THE PHYLLOSTOMATIDAE 263 ISI Flg, 6. —Warbles of Di^sfnodus ant! L^ptouycreriy. A. Warble emitted by mother of five-day^old Desmodus at the mcmeni of bodily contact during a reunion, B. Warble emitted by mother of five-day-old Lepionycienx at the moment of bodily contact during a reunion. C, Isolated 5-day-old Desmodus exploring fiat substrate; compare with E. D. Leptonycteris, 15 days old approaching its mother during a reunion. This call and the one in F, both from different animals, resembled the warble emitted by their mothers (for example B). It seemed as though the infants were attempting to imitate their mothers, E* Isolated 41-day-old Dvsrfwdus exploring flat substrate; this bat had been hand raised from day seven, F. Leptonycieris, an eight-day-old infant, near contact with its mother. G. Isolated five-day-old Desmodus whWe being held in hand. 264 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY B 96 - !64 32 ! 0 ^ 211 Mi niSEc c Di 12 8 4 ; qL r fSi It' ,». ,t 'i I Time ( 'ik Pin. 7.—Mixed and compound units; A. Mixed unit emitted by a hand*held, 65- day-old DcsfUiHliis. A buzz is superimposed on an FM pulse and an ascending FM pulse. B and C. Calls of one-day-old Di^s/ntHlus. Mixed units of buzz superimposed on slightly more structured pulses. D. Compound unit of DNS and buzz emitted by a 41-day-old Di’s>nos of vtH uHzatious attitu'd hy iiuiivuiuals of five f^enera of phyllosto/nufkl hots iinrin}> iliff'ercf}! circnnt.shifives. Nionhers represent ajie of imlividiiti! hats in thiys\ .v, infant sepnnitedfrom mother or mother silent ilitrin^ n lennioir, a, nOfthef s approach ilnrini’ rennion: t', mother emitiinp FM pnise.s diirifip rennii}n; t , mother makinp contact ditrif!}> rettnkfn. Curattiu h'pf‘»iyru‘rts PhyUoMinitu> Mae raff iS f-call 1 s. 1 s. 4s DN Is, 7saec, 16saec. 1 s, 7ces, 8s Ics, 7 as .''sa, las, la Is Is 11s 13e, 17s 17s, 9a FM pul.se 7es. 16ae, 9s 4s I 4s Is Is Is Ms FMGB 13s, 7c Novick (1963) FMGE 16a iri .ft Warble 4dt, 8dtt 41s*. 5c Buzz t9d 5 c, 180**, 210**, Is. 180s, 41s An iisKTisk itlfnriOes ii haiid^raiscd infiini; double asterisks, data I'roni Schmidi (1972), ase in days is approstmaie; dagger, vocalizations probably from moihcr only; and double dagger, vocalizations probEibly from mother and infant (.see Pig. bD). .Novick (1963) recorded FMGU from adull Lepnntyaeris. (Gould, 1970, 1971). This specificity in Caroiiia is consistent with the behavior described below. The vocal exchange belw-een mothers and infants in the following tw'o de¬ scriptions typifies reunions that I observed of other species in somewhat diftereni circumstances. Seven “day-old Carol I ia Minus 0 .—infant was placed on horizontal branch and remained there until reunion with mother, 0-23 sec .-—Tape recorder on. .Mother relea.sed in chamber. Mother flies the width of the room several limes. Infant emits FM pulses (low intensity unless otherwise noted) and numerous DNS. 24-30 sec .—^ Mot her hangs up on ceiling, infant slops emitting FM pulses; infant continues emitting DNS. 51 sec.-2niin., 9 sec .—^Mother continues flights across room and intermittently hangs up. Infant emits DNS and FM pulses when mother flies. 2 tnin., iO sec.-S min., 40 sec .—Mother flies back and forth across the width of the room and hangs tip three times. Infant emits DNS w hen mother hangs up. While mother flies but does not approach infant, infant emits DNS and FM pulses. In one sample of 33.7 sec. of infant vocalizations in which FM pulses graded into and out of DNS. infant emitted 20 DNS and 302 FM pulses. During mother's three hang ups lasting a total of 50 seconds infant emits 11 DNS and no FM pulses (Fig, 10). 3 min.,41 \ec-3 min.,43 sec .—Mot her Dies toward infant and approaches wi thin 12 centimeters of infant. Infant increases rate of DN and rate of FM pulses (Fig. 10). 3 min., 44 sec.-lS min. Mother approaches infant 12 times; infant increases rate of DNS and FM pulses during each approach. Mother hangs up 16 times. Mother flies back and forth 7 times. }8 tnith, 5 .sec .—Mother approaches infant. Infant emits FM pulses, 0 to 19.5 + pulses per second (in eleven 0.95-sec. consecutive samples) and DN 0.4-2,3 calls per second (in the same 11 samples). During the same II samples, FMGB and FMGE sounds are emitted, probably by the infant (Fig. 4C F). Mother hangs up next to infant and guides infant to HIOLOGY OF THE PHYLLOSTOMATIDAE 269 R ATE OF V OCALIZATION BY A DAY OLD CAROLLlA HOTnfA fLY^NC hl9TM(ft fltim M hfClHCA MD (W*lET □ I Z Fu.. 10.—Rale of vocalizations by a seven-day-old Caroflui during a reunion. The three sets of data were obtained as the infant remained motionless on a branch and the mother flew toward the infant or roosted quietly at a distance from the infant. Four arrows indicate the limes that the mother flew toward the infant. The mother’s FM pulses were not detected on the tape recording when the mother was flying about the room or when the mother was quiet. When the mother approached the infant, her FM pulses were detectable. The rate of her calls was not determined. Rates of DN and FM pulse emission while mother was flying and approaching (left and middle set of data) were obtained by slow ing the recordings 128 times; the number of infant FM pulses was counted during the intervals between DNS. From these two tallies, the number of FM pulses per DN interval and the DN interval were used to compute (he two rates. Intervals were mea.sured from the beginning of one DN to the beginning of the next. Note that the vertical scale on the right refers to DNS; the vertical scale on the left refers to FM pulses. nipple. Infant attaches to nipple. Just before mother flies from branch with infant attached, mother emits six FM pulses. The tape recorder was on for 6 minutes, 15 seconds during which time infant emitted 162 DNS. The ratio of DNS to FM pulses varied from 1:3 to 1:4. Note that the tape recorder was only on about 35 per cent of the time. I plotted rate of FM pulse emission against time between DN phrases. The rate of FM pulse emission decreased as the time between DNS increased. The relation¬ ship is most apparent from the data obtained during the mother’s approach to the infant. The plot of data obtained when the mother was flying but not approach¬ ing the infant is scattered. Sixteen-day-old Carol! ia Minus Or —Infant was placed on horizontal branch and remained there until reunion. 0-i min., 35 sec .—^Tape recorder on. Mother released in chamber. Mother flies back and forth across room and then hangs up. Mother hangs up seven limes. Infant emits a few FM pulses while mother flies. Infant is silent when mother hangs up. / min., 40 sec.~l min., 45 jcc.—Mother approaches infant within 12 centimeters for the first time. Infant emits burst of FM pulses with increased repetition rate during mother's approach. 1 min., SO sec.-25 min .-—Mother hangs up 22 times. Mother approaches infant within 12 to 20 centimeters 16 times. Infant emits burst of FM pulses each time mother approaches. During eight approaches, infant also emitted a total of 16 DNS that included FMGB (14) and FMGE (2). In three approaches, infant FM pulses could be distinguished clearly from those of the mother. During those three approaches, infant FM pulse rate emission increased in a similar pattern; in 940 msec, following the infant's first call, it emitted 8, 9.1 and 8,5 270 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Table 3. — -DN L’/nissio/! decreaxes ux the hai matnrt'\. R idi‘nfifi(’x dma idyicdiu'd front rvci>niifj}?x of leunionx-, the remainder of the data ii'irv obtained frooi recordififis of ixitluied iitfnntx. Species* Age Number of DNS Time sampled in sec. Raie/sec. Leptonyeierix !R 89 246 0.37 15R 24 4,50 0,04 PhyUoxtotnux 2 38 120 0.3 7 0 60 0 Desmodns i 107 12,4 8.6 5 60 16.8 0.3 *Each set of data derives from a dtlTcreot hat. pulses/second in the three approaches respectively; in the next 940 msec., 8, 16, and 7,5 pulses/second, and in the last 940 msec., 1.6, 0.6 and 2.1 pulses/second. All three outbursts (composed of a total of 26, 23 and 17 FM pulses) were preceded and followed by long periods of silence from the infant. The mother's calls also were evident on the recording. 25 ntim, W .rer.— Mother lands near infant. Six DNS with FMGB units are emitted, probably by the infant. Mother guides infant to her nipple. Just before taking off, mother emitted FM pulses of higher intensity than any of those recorded from the infant. The tape recorder was on for 8 minutes, or 31,8 per cent of the time; it was turned off primarily when the mother was hanging. The infant emitted 34 DNS. The DNS were so infrequently emitted that 1 usually could not determine inter¬ val between calls. Four DN intervals measured 1.3 to 1.6 sec., and four measured 4 to 11.25 sec. Table 2 shows the incidence of various calls during reunions and other experi¬ mental situations. In general, older bats emit few'er DNS (Table 3). For the 16- day-old Ccirollia and 15-day-old Lt'ptonyaens, DN emissions were associated with approach and contact; younger bats emitted DNS more often and under more varied circumstances than did older ones. The rather abrupt decrease in DN emissions from CaroUia at about 18 days of age is coincident with the time at which young begin to Oy (Fig. 11). At 18 to 20 days of age, these bats should be able to take a more active part in their attempts to reunite with their mothers. Mac rot us I have observed reunions of mother and infant in eight genera of bats. None of the mothers displayed as much determination to make contact with the infant as did those of Macrotus vaUfornicus. In 7 trials with three different mothers and their one-day, two-day and 11 -day-old infants, the behavior was the same. As the calling infant w'as brought within hearing range (at least 3 meters) of the mother, the mother lunged from the back of her cage onto the door where she clung and ‘TVantically” emitted DNS. When the infant was beyond hearing range of the mother, the mother was quiet. DNS are probably the mother’s responses to the infant’s calls. Tw'o adult Macrotus responded to the calls of their infants by emitting DNS; the infants were held 0.75 meters from the mother and the microphone BJOLOGY OF THE PHYLLOSTOMATIDAE 271 g CO g S UJ tj_ o LJ a: RATE OF ON EMISSION 8Y CAROLUA U.y A 0.8 A A fly, turn, goin altitude 07 A 0.6 < L 05 AR 0,4 0,3 0,2 A 0.1 - ^ A -,-^ -*---♦ -r-1-1-T-"-1-f 0 5 10 18 20 30 40 AGE IN DAYS Fig. Mr—Rate of DN emission of IS individual OirotlUi at ages from two lo 40 days. Recording samples ranged from 51 to 480 seconds (mean, 99; mode, 116). An R next lo the symbol means that data were obtained during a reunion of the mother and infant. All other data points were obtained from isolated bats that experienced similar handling and explora¬ tion in a box. According lo Bleier et al. (this volume), Carollia can maintain and gain altitude and turn and avoid obstacles at 18 days of age. Emission rate is expressed as pulses per second. was placed a few centimeters from the mother. A sample of 74 phrases (usually two units per phrase) in six bouts had rates varying from 2.9 to 15.0 phrases per second (mean, 8.8). As soon as the infant was placed in the cage, the mother guided it to her teat even while the infant was still in my hand. The mother’s and infant’s DNS were so similar and so often emitted by both mother and infant that data analysis during close contact was next to impossible. Greater separation in a large enclosure is essential for further study of this species. A reunion in a large enclosure (366 by 732 by 183 centimeters; Bee Tent, Chicopee Mfg. Co., Cornelia, Ga, 30513) revealed unexpected maternal behavior from an II-day postpartum mother. Macrotus califoniicus generally gives birth to a single infant. Two infants, one and 11 days old, were placed about 15 centimeters apart on a fence post. The mother of the older infant w'as released in the enclosure and photographed with a 16~millimeter motion picture camera that was synchronized w'ith repetitive flashing strobe lights (about 2.1 meters from the post). The film showed clearly, and substantiated our direct observation, that the mother approached the younger infant first and guided it to her nipple, she then crawled to her own infant and guided it to her other nipple and flew off with both infants securely attached Before the mother landed, the younger infant called incessantly while the older infant called occasionally. 272 SPECIAL PUBLJ CAT IONS MUSEUM TEXAS f'ECH UNIVERSITY Lc^pt(Htycteris As two infant (one and eight days of age) Lepfouyaeris sanborni approached their mothers, they increased emission rates of DNS just prior to contact. In both reunions, the mother remained motionless and often quiet in the corner of the cage most distant from the door (during the first week or so after birth, Ltpto- nycteris infants can walk; they then lose this ability and never regain it, Dietz, 1973). During the 13.8 seconds before contact, the 8-day-old infant emitted six DNS having the following intervals: 560, 180, 4009, 4009, 130, and 130 msec, respectively (range of repetition rate, 0.3 to 7.7 per second). The last three DNS w'ere emitted just before, or when, the infant made contact with its mother. Both infants flapped their wings as they hung from the roof of the cage opposite their mothers. Each bout of wing flapping was accompanied by DN emissions. The force of the flapping raised the infants to a nearly horizontal position. Tw'o to three-week-old young also Happed their wings; however, wing flapping in this case was accompanied by FM pulse emissions. Warbles w'erc detected on recordings of reunions of four and eight-day-old infants w'ith their mothers. 1 could not determine whether the mother or infant emitted the calf At least three naturalistic observations of Leptonycteris sanborui suggest the importance of acoustic communication. 1. In Colossal cave, near Tucson, Arizona, large concentrations of these bats gathered in small groups of 50 to 75, circled and then departed in a “string’" composed of many small groups (four to six individuals each, Hayw'ard and Cockrum, 1971). Other species leave their cave roost in a .single ma.ssed evening flight. In the dark of a cave, group flights may be organized by acoustic signals such as w'ing Happing, FM pulses or DNS, or the like. 2. Hayward and Cockrum (1971) suggested that twittering (probably the audible component of DNS) seems to serve as a bond in keeping members of a LAfptonyvteris colony together; females hovered over, and landed on, sacks containing loudly twittering ^adults. 3. During reunion, the mother would wrap her forearms about a Juvenile in front of her, draw it to her breast, then release her hold on the ceiling and fly away. This apparent search for, and identification of, a particular young bat was noted many times. Other young might attempt to attach but they were given no notice other than a fending off motion of the forearm. Of course, olfaction may play an important roll in the reunion. Destnodus Reunions of mother and infant Desmodia mtimdus w'ere similar to those of Lt'pumycteris scmbonii in that the mothers remained motionless in the rear of the cage and only occasionally vocalized. Infants called often, frequently emitted DNS, and took the initiative by approaching their mothers. At or near contact, infants increased their DN repetition rate. Schmidt (1972) described reunions typified by more active mothers. His descriptions, including the vocal exchange between mother and infant, resemble my observations of Macrotus caiifornicus. BiOLOGY OF THE HHYLLOSTOMATIDAE 273 This resemblance in our data could be derived from population variation or to undetermined disturbances in my colony; the infants in my study did reach adult size. Schmidt {1972) noted the relationship between DN emission and the relative distances of a young of Desmodits rp?i), Pp. 281-324, in Social communication among primates (S. Allmann, ed.), Univ. Chicago Press. Chicago, xiv + 392 pp. SucA, N.. AND P, ScHLEGEL. 1973. Coding and processing in the auditory systems of EM-signal-producing bats, J. Acoust. Soc. Amer., 54:174-190, SuTHERS. R. A. 1965. Acoustic orientation by fish-catching bats. J. Exp, Zool., 158: 319-348. —, 1970. Vision, olfaction, taste. Pp. 265-309, iti Biology of bats (W, Wimsait, ed.), vol.2, Academic Press, New York, xv + 477 pp. SuTHERS, R. A., AND J. M. Fattu. 1973. Mechanisms of sound production by echolocat¬ ing bats. Amer. Zool,, 13: L215-1226. Tamsitt. j. R. 1967. Niche and species diversity in neotropical bats. Nature, 213: 784-786. Taylor, H., E. Gogld, A. Frank, and N, Woolf, 1974. Successful hand-raising of one week old bats. Eptesicus and Antroious, by stomach catheter, J. Mamm., 55: 228-231. Vernon, J., and E. Peterson. 1966, Hearing in the vampire bat, Desmodtrs rotitiiditx as shown by cochlear potentials. J. Aud. Res., 6:181-187. Woolf, N. 1974. The ontogeny of bat sonar sounds: with special emphasis on sensory deprivation. Unpublished ScD thesis, Johns Hopkins Univ., Baltimore, 140 pp. THERMOREGULATION John J, McManus There is a subsJantial literature on mammalian thermoregulation, and several comprehensive reviews are available for different orders of mammals (Whittow^ 1971). Excellent surveys of existing literature on temperature regulation in bats were prepared by Stones and Wiebers (1965), McNab (1969), and Lyman (1970) and these should be consulted for general coverage of the Chiroptera, Also, be¬ cause many species of bats enter torpor, the order is frequently discussed in con¬ text with the evolution and adaptive significance of heterothermy (Hudson, 1973; VVhittow, 1973). Most earlier papers on bat metabolism and thermoregula¬ tion dealt with temperate zone microchiropterans (Hock, 1951; Reeder and Cowles, 1951) or tropical megachiropterans (Burbank and Young, 1934; Bartholo¬ mew ei aL, 1964). A rough pattern emerged from these studies, which contrasted the relatively precise homeothermy achieved by the large, frugivorous mega¬ chiropterans with the pronounced thermolabiiity and capacity for torpor shown by the smaller, insectivorous species that inhabit areas with more rigorous ther¬ mal environments. Within the past decade, many additional species of tropical microchiropterans have been examined. Most of these have proved to be inter¬ mediate in their temperature regulation and, collectively, they illustrate the continuum of thermoregulatory strategies that exists within the order Chiroptera. These recent works include data on temperature relation in more than two dozen members of the Phyllostomatidae, and the scope of this chapter will be restricted principally to a summary of this information. Some attempt will be made to place w'hat is known of thermoregulation in phyllosiomatids in evolutionary perspective and to suggest possible directions for future studies on New' World leaf-nosed bats. Status OF Thermoregulation by Phyllostomatids Problems of Measurement Even the most casual survey of the literature indicates that within the Chiroptera there is an extraordinarily wide variation in the quality of temperature control. This variability makes generalizations regarding thermoregulatory strategy difficult and forces a careful analysis of the sources that contribute to w'ide dif¬ ferences in results, even at the species level, Lyman (1970) has enumerated some of the more important factors that should be considered in assessing temperature regulation in bats: 1) phylogenetic antiquity of the order; 2) high species diver¬ sity; 3) broad geographic distribution; 4) wide differences in habitat preference, feeding habits, and daily and seasonal levels of activity; 5) preponderance of smaller size classes; 6) high surface area to volume ratios; 7) capacity for energet¬ ically costly sustained flight; and 8) peculiarities of behavior and habitat require¬ ments in the w'ild that often make it difficult to simulate ecologically realistic 281 282 SPEC I A], PUBLICATIONS MUSEUM TEXAS TECH UN I VERSO V conditions for testing the tliernioregiilalory capacity of captive animals. Thus, it is not surprising that various investigators have employed quite different methods and criteria for assessing temperature control in bats. All of the above constraints apply to the literature of the Phyllostomatidae and these should be noted when the following comparisons are made. A tabular summary of recent work on phyb lostomatid thermoregulation is given ( I’able 1). EJfeci ofBoily Wcigiu on Basal Heal Froclaciitni In general, maintenance of a differential between body temperature and re* duced ambient temperatures requires the generation of metabolic heat at levels just equal to the rate at which heat is lost from the body. Because heat production occurs within the volume of the body and heat loss is primarily a surface pheno¬ menon, the ratio betw'een surface area and body volume is important in establish¬ ing the level at which heal production and heat loss are balanced. Smaller animals, in w'hich the surface area is high in relation to btxiy mass, w ill have pro¬ portionally higher rates of heat loss and heat production than would be found in larger species. The predictive relationship between weight-specific metabolic rate (heal production per unit w'eight per unit time) and btxjy weight takes the general form M — AW*, where M is weight specific heat production, W is body W'eight, and k and /) are constants. When heat production for many species of mammals at thermoneutrality is calculated as milliliters of oxygen per gram per hour (ml. O 2 g/hr), the constants k and h lake the values 3.8 and -0,27, such that M = 3.8W - 02 ^ (Brody, 1945; Kleiber, 1961), McNab (1969) compared the basal metabolic rates of 23 species of New- World leaf-nosed bats to the rates predicted for mammals within the weight range of nine to 97 grams (Fig. I). As in other bats, the basal metabolic rates of phyl- lostomatids are inversely related to body size, and, for most species, the measured rates of heat production tend to be higher than those predicted by btxiy w'eight. This is expected in view' of the high surface area to volume relations caused by the presence of w'ings, large ears, and associated membranes. The level at w-hich body temperature is maintained is dependent on the ratio of heat production to heat loss and appears to be weight dependent (McNab, 1969, 1970). Smaller species of bats tend to have lower body temperatures than do larger species and this generalization seems to hold true for those phyllostomatids for which body temperatures have been recorded (Fig. 2). It is obvious, however, that smaller species of leaf-nosed bats show' considerable variation in resting body temperatures and in many instances it is not clear w'hether this is the result of species differences or a reflection of such confounding variables as differences in physiological state, nutritional status, activity level, or methods of body tempera¬ ture measurement (Lyman, 1970; Studier and Wilson, 1970). Nevertheless, be¬ cause heat loss depends directly on the differential betw'een internal and external temperature, even slight reductions in the set-point temperature by smaller phyl¬ lostomatids are advantageous in that they decrease the levels at w'hich heat must be produced for maintenance of homeothermy. BIOLOGY OF THE PHYLLOSTOMATIDAE 2K3 Effect of Diet on Basal Heal Production Phyilostomaiids can be placed in categories based on their feeding habits— fruit and nectar feeders, larger carnivorous feeders, insect feeders, and blood feeders (McNab, 1969, 1970). Excepting the sanguivores and perhaps some of the most specialized nectar feeders, it is likely that insects are taken to varying degrees by all leaf-nosed bats, but the preceeding categories will be used as approximate indices of the bulk of the diet. Fnigivorous .—The leaf-nosed bats as a group are principally fruit eaters and this dietary commodity apparently has exerted a strong influence on temperature regulation in the family. McNab (1969) argued persuasively the notion that seasonal availability of fruit and nectar in Neotropical areas has allowed frugivo- rous phyllostomatids the luxury of elevated metabolic rates (Fig. 1) and has made nonessential the diurnal and seasonal torpor characteristic of temperate-zone insect feeders, the food supply of which is subject to severe daily and seasonal fluctuation. Arata and Jones (1967) posed a similar hypothesis and extended the reasoning to include tropical insect feeders. Despite a high degree of variation, resting body temperatures of frugivorous leaf-nosed bats (Fig. 2) tend to be held at relatively high levels, approximately 5 to I0®C higher than those of insectivorous species of comparable size (McNab, 1969; fig. 31; Lyman, 1970: table I). It seems reasonable that this could be achieved only if energy sources were readily avail¬ able to support the high levels of heat generation needed for such homeothermy. Carnivorous. —The meat-eating phyllostomatids ( Tonaiia bidens, PhyHosiomus discolor^ P. hasiatus^ P. elongams, Chrotopterus auritus) appear to approximate or slightly exceed the basal metabolic rates predicted by w'eight. They also are among the largest of the phyllostomatids. Practically nothing is known about the diet of carnivorous bats in the wild, but I presume that vertebrates comprise its bulk. Considering the relative stability of vertebrate populations in tropical areas, compared to the more pronounced fluctuations in abundance and availability of populations in temperate zones, one can speculate that larger carnivorous phyllostomatids have adjusted their energy expeditures in response to food re¬ serves that remain relatively fixed in supply throughout the year. Availablity of food and large body size contribute to their fairly precise control of body tem¬ perature. However, the carnivorous species of leaf-nosed bats apparently depend on food supplies that, while temporarily available, may be spatially distributed in a way that cannot support second-level consumers. McNab (1971) observed that carnivorous species are the first to disappear from bat faunas on tropical is¬ lands with depauperate vertebrate communities. insectivorous, —Although Arata and Jones (1967) postulated that tropical insect feeders may resemble fruit and nectar feeders in thermal ecology, McNab’s (1969) study suggested that they are quite different. Tropical insect feeders more closely resemble temperate zone taxa in their proclivity to relax thermoregulatory control when at rest and in their tendency to have low'er basal metabolic rates. This apparently results from the fact that insectivorous species tend generally to be smaller and gain considerable metabolic savings by reducing body temperature- 2«4 SPECIAL F’UBLlCAi'IONS MUSEUM TEXAS TECH UNIVERSITY Fk;. L—Basal merabolic rates of phyllostomatid bats. The regression line indicates the level of metabolism expected for a given weight based on the general relationship for many species of mammals: BMR = 3.SW “0T7 (Kleiber, 1961). Data are based on McNab's (1969) study. ambient lemperature differentials. In addition, for many tropical areas, the pre¬ sumption of a constant level of availability of insect foods is unfounded; tropical insect populations may respond seasonally to rainfall in a fashion similar to the response of temperate zone insects to sea.sonal temperature changes (Janzen and Schoener, 1968). Thus, tropical insect feeders do not utilize a dietary item in constant supply and cannot afford the luxury of the elevated metabolic rates seen in frugivorous species. Of the phyllostomatids studied, metabolic data are available for only one insectivorous species, Tonaiia sytvicoia (Fig. 1). It appears to conform to the pattern seen in other tropical insect feeders. Sanguivomas .—Because of their unique feeding habits, vampire bats have attracted the attention of several investigators, and details of the thermoregulation and bioenergetics of one species, Desmodus rotundas^ are as well knowm as for any bat. Early observations (Wimsatt, 1962) indicated that despite its moderate size {30 grams), Desmodus rotundas was surprisingly ineffective at controlling body temperature. At rest, body temperatures of vampire bats fell close to tmibient temperatures (even at environmental temperatures as high as 33^*0, Lyman and Wimsatt, 1966), and responses of individuals to reduced ambient temperatures varied markedly. McNab (1969, 1973) provided data for Diaemus youngii and Diphyita ecaudata, as w-ell as for Desmodus rotundas, and both basal metabolic rales and body temperatures tended to be low' in these species (Figs. 1,2). The effect of diet on the thermal economy of desmodontines w'as considered extensively by McNab (1969, 1973) and the following is a summary of his findings. The use of mammalian or avian bltxyd by a vampire bat requires transporting all or part of the blood meal in flight. The size of the meal that can be ingested, therefore, is limited by the ability of the bat to carry it back to the roost (Crespo et aLy 1970). Furthermore, w'hole blood has a relatively low caloric density BIOLOGY OF THE PHYLLOSTOMATIDAE 285 • truir 40 - O Jniactj X = 35.10 X = 36.62 o 38 ! 8 # blood • • ® • • m • 36 • @ • o . u o - © ' 34 32 > o i 30 -L.i_1_J. J-1 J _1_t_1_1_L_ 1 _1_ i _i ■ .1-1 J 20 40 60 80 100 Body Woighi - grams Fig. 2, —Resting body temperatures of phyllostomalid bats. Temperatures measured at ambient temperatures greater than 25°C, Based on data in Table 1. and, although some concentration of the meal may occur prior to flight (by urination while feeding—McFarland and Wimsatt, 1969), vampires appear to be limited by the amount of energy they can acquire and process per foraging flight. Such limitations on energy intake would presumably be most severe for females near the end of pregnancy w'hen load-lifting capacity is lowered and ab¬ solute energy needs arc greatest. The conclusion drawn, then, ts that the type of food employed by vampire bats forces them to conserve energy at times other than flight by relaxing control of body temperature while at rest and by sustaining relatively lowered rates of basal metabolism. Resistance lo Hyperthermia With the exception of a few' species, high temperature tolerance has not been studied systematically in leaf-nosed bats. The temperate zone species Macrotus californicus responded to increasing ambient temperatures (T^) by initiating a series of slow wing-flapping movements when body temperature (Tb) reached 32.6(Reeder and Cowles, 1951). At Tb 34.7®C, wing movements increased in frequency and at Tb 38.7®C constant fanning occurred; after 26 minutes, T^, was held at 39.0'"C in a T^ of 40.6®C. Wimsatt (1962) reported Desmodus ro¬ tund us to have surprisingly little tolerance of even moderately high air tempera¬ tures and suggested that the critical tolerance level w'as within the T^ range of 27 to 30°C. In controlled experiments, Lyman and Wimsatt (1966) found this same species unable to tolerate Ta’s of 33 to 34°C for more than two hours. No wing fanning or salivation responses were noted. Unpublished observations (McManus and Nellis) on Art then s jama ice nsis indicated that individuals of this species can tolerate T^ 40°C for up to five Table 1 . — Thermon’f'nUiiiOfi liam for various phyllantomatiJ hats. Mb is hasai metuholk raw e.xpressi’il us mi. Ozl^tlhr. C is thermai c<}rjtiuctance give ft as fill. O^lgfhrf^ C. Mil temperufares are in degrees cent igraiSe. 286 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY (K ^ CJ f-l > ^1 it an - z a ^ a a £ = S 3 2 £ ■t: -£ i ■“ -5 >> 1 — —> I 1111 m 1 m gj ^ 51 ill I u - (5 a 5 “ ?5 -- fs fl Wl«iJ3U31>3y § £ H o. "5 -Q a T a T £ > £ Q SH - — O ■- £ (5 li; c « I I 5 I I I r-l r4 — — 2 «- ^ ? -s: i. II 5 5 5 T I o -t ri ^ — SSE Tf wa ^3 5; :: a I 1=5 .£ a £ = III ^ O rt _; ^n o — I -1 I »*ii m m 2 E ^ 2 ^ 4 xiuHfa frtiJi/y/i'r 1 K 5 (many) , 16.5 3.71 0,36 36.5 34.5 34-3 5 40-2 34 — — fruit. nectar CaroJIijnac Caroitia perspicillafa 14.9(itiimy) 36.4 2.89 0,35 22-37 17-3S 12-37 39-40 37-39 — — fruit liictaiing 9 mamtained Tb 6J0 at Ta's 7.1-26.3" Rhinopfjytlu pumiiist 9.5(9) 34,7 1,96 0.37 34-36 19.5-34.5 ,34 3«.8 37.5 — — fruit — 10 BIOLOGY OF THE PHYIXOSTOMATIDAE w-i j-n d ao -c J? n s 1 E-g -C o la so o S 'Y' > rt 'I' ri ^ H o » « H U d p 7- C “ d g A ,E ^ JG £ ad o 5 if I”^ 3 a *"H 13 .Ed « i_i 2 r3 ^ ^ “ 2 ''' JS oi > hi O 11 II y E rt -Ji M t d d A J5 H E o a w d ^ g «-s? I f E I I I I [Ml I I I S 2 .2 .2 I 2 I I * I I I M I I T 5 i ! I o o o o r>^ l*-| <1 't O' a rt f-i — (s r4 la-j i-n Tf O r- r^i fJ —* ^ ^ vT (N S = S III X 'C I I A I I 0 A H A eu o « E T (-J U f'. Q f*i Js S' ? E ^ — Es sJ. ^ i w V ^25 I f I I r' rj « rj d d (M " S' do 3 .5 q q E I ^ "5 c s (fl s ‘“ 'S n '*' 9- 2 # f* at S is E « e S « rt ei ^ qj ^ « ? s; in - u I 5 “v.. u > O 'O u — c 5 5:: Hi ^ f P4 li- ■^1 s ’ll 3 ih S: S ^ 5 X) U ^ f- « 287 (1967); 7, Carpenter and Graham (1%?); 8, Morrison and McNab (1967); 9. LsVal (1%9); 10, McNab (1969); 1), Ras^eikr (1970); 12. Smdjer and Wilson (1970); 13, McManus and Nellis (1972); 14, McNab (1973). 288 SPECIAL PUBLICATIONS MUSEUM TEXAS LECH UNIVERSITY hours, but die within two hours at 45 Lethal body temperature is near 43^C Carpenter and Graham (1967) found that A. hirsuius begins panting vigorously at T^, 38”C and suggested that these bats probably cannot survive higher ambient temperatures. The same authors reported that Leptonycieris sanhorni maintained a body temperature between 39.2 and 41.5“C after a four- hour exposure to 41.5”. Although Ij.’pkmyctens was quite efficient at resisting hyperthermia, no conspicuous thermoregulatory behaviors such as wing fanning or salivation occurred. T'he lethal body temperatures listed in Table 1 usually w'ere recorded after accidental deaths during oxygen uptake tests. Collectively, they resemble those of other mammals, although lethal T^ values for Carollia perspicillata and Rhinophyfla pnmilio arc suspect because death probably re¬ sulted from causes other than hyperthermia. Levels of ambient temperature tolerated by phylbstomatids tend to be high, but not exceptional. This probably reflects the moderating nature of most roost sites and the nocturnal habits of the animals. RcsisUince (o Hypolhermkt Asa rule, leaf-nosed bats .show little tendency to experience large reductions in b(xly temperature when exposed to low ambient temperatures for short periods (Table 1), As exposure time increases, however, body temperatures vary erratical¬ ly, particularly at ambient temperatures outside the normal range of temperature encountered in the vvild. Additionally, the length of time betw-een capture and testing appears to confuse the issue (McNab, 1969; Studier and Wilson, 1970) and nutritional status undoubtedly influences the magnitude of Tb-Ta differentials sustained. In several species, exposure to low ambient temperatures causes a drop in body temperature, but to some relatively constant level above the ambient temperature. Studier and W'ilson (1970) computed regression formulas to de¬ scribe the relation betw'een T^ and T^ for two species at various ambient temper¬ atures. Betw'ecn T^ 33,2 and 8.0”C, Arfiheus januiicensis gave the response Th = 8-8 + 0.9333 Ta whereas betw'cen 33.1 and 7,8”C, Vampyrodes caraccioloi show'cd Th= 12.0 + 0-813 Ta. McManus and Nellis (1972) found A.Jamaicensis able to maintain Tt, above 35” after six hours at 10”C and one individual w'as able to keep Tb above 18” for at least 14 hours at 10”. Brachyphylla cavernarum held Tb above 25” after 24 hours at 10”. There are no knowm species of phyllostomatids that naturally enter deep torpor-—even those species, such as Matroiuscaiifo miens or Lepfonycierls sanhorni^ which are native to, or regularly enter, temperate areas. Carpenter and Graham (1967) observed that Li’pfonyaeris sanhorni and Artihens hirsiifns w-ere able to regulate bexiy temperature within levels of precision equivalent to those of other small mammals of comparable size, and McN ab’s (1969) study extended this pattern to many other species. An interesting observation made by McNab (1969) on PhyUosiomus discolor, Toncuia hidens^ and Ciirofoptcms anriuts was that these species may show' a temporary relaxation of temperature control at moderately reduced ambient temperatures (approximately similar to those of their roost sites), yet are capable BIOLOGY OF THE PHYLI.OSTOMATIDAE 289 of more effective thermoregulation at lower temperatures. The implication is that temporary hypothermia may be “intentionally*’ tolerated and serves to de¬ crease the differential between ambient and core temperature. Such a strategy would reduce energy requirements during periods of inactivity and may explain in part the rather pronounced diurnal variation in resting body temperatures noted by Morrison and McNab (1967). Nevertheless, the overriding tendency among the majority of leaf-nosed bats examined is to maintain btxly temperature at a roughly constant level when exposed to moderately low- ambient temperatures. When forced to withstand temperatures well below' those normally encountered, responses vary markedly, with larger species tending to conserve body heat more effectively than smaller kinds (Table 1). Compared to temperate zone micro- chiropterans that regularly enter deep torpor, the lower lethal minima for phyl- lostomatids appear to be higher by 10®C or more. S(>ci a I Tfierm oreguJm ion With one exception, the energetic significance of clustering as a means of behavioral thermoregulation has been neglected in New World leaf-nosed bats. In torpid microchiropterans from temperate areas, clustering is regarded as a means of reducing variations in body temperature by decreasing the exposed surface area of any individual in a cluster. In contrast to the result of such behavior in strict homeotherms (Tertil, 1972, and references therein), clustering in hiberna¬ ting bats is not intended to conserve heat, but rather to keep body temperature low' and to avoid temporary increases in ambient temperature (Twente, 1955). Such behavior promotes the most prudent use of stored fats during hibernation by keeping metabolic rate low (Hock, 1951; McManus and Esher, 1971). Apparently a similar strategy is employed by at least one species of phyllostomatid, Phyllostonuis discolor. McNab (1969) found that clustering at 20°C resulted in a drop in both mean body temperature and mean resting metabolic rate of bats in a cluster as compared to that of single individuals. Whether such behavior is widespread among leaf-nosed bats is unknown. Additionally, the object of social thermoregulation in species with more precise thermoregulatory control than P. discolor (see preceeding section) may in fact be heat conservation rather than maintenance of reduced body temperatures. Finally, for species of bats that regularly form clusters while roosting, the testing of an individual in a metabolism experiment is an abnormal situation and may contribute greatly to variation in physiological performance. 1 suspect that this area would yield interesting re¬ sults with further study. Torpor Deep torpor and seasonal hibernation are found only in the families Vespertil- ionidae, Rhinolophidae, and to a lesser extent in the Molossidae, Viewing this as one extreme condition and precise endothermy as the other, phyllostomatids appear to occupy a broad portion of the spectrum. Hudson (1973) regarded the ability of Neotropical bats to tolerate low body temperatures as “an ancestral phenotype (as well as genotype) which could easily be modified to give the ‘deep’ 290 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY or True’ hibernator. Under tropical conditions, such a thermoregulatory per¬ formance represents a solution for which there is no problem,” A similar thesis was advanced by Studier and Wilson (1970) who suggested the possibility that the thermolability of tropical bats may be nonadaptive. Although this may be true for frugivorous and perhaps carnivorous bats, the facultative capacity to experience and tolerate slight reductions in body temperature during periods of inactivity is clearly advantageous for small insect feeders and sanguivores. Conclusions The family Phyllostomatidae presents a varied mosaic of thermoregulatory strategies, but the problems of measurement, coupled with natural variation in levels of temperature control among species, make broad generalizations difficult. Perhaps the easiest w'ay to summarize this review is to indicate those parameters that appear to have infiuenced the development of thermoregulation in leaf-nosed bats. Body size appears to affect resting rales of metabolism and resistance to hypo¬ thermia in much the same w-ay as it does in other mammals. Qualitatively, smaller species have higher basal metabolic rates and poorer temperature control at reduced temperatures than do larger species. Quantitatively, the peculiar surface area to volume ratios unique to bats cause the levels of metabolism to be higher in general than those of other mammals of comparable weight. This disparity is most obvious in small and intermediate-sized phyllostomatids. Diet and trophic position seem to be of particular import in determining thermo¬ regulatory performance. First-level consumers feeding on spatially and temporally available foods such as fruits and nectar achieve higher basal rates of metabolism than do most second-level consumers. Carnivorous species have fairly precise thermoregulation, but this is associated with their generally larger body size. Insect feeders and blood feeders, for which fot>d supplies are temporally or lo- gistically limited, or both, seem to have developed either a more relaxed pattern of temperature control or have reduced the set-point at which metabolism proceeds. One glaring insufficiency in our knowledge of thermoregulation in leaf-nosed bats (and most other bats) is the paucity of data on food intake under natural conditions or in the laboratory. Associated with this is the scarcity of field temper¬ atures for bat roost sites upon which any type of bioenergetic analysis must de¬ pend. Stx'ial aggregation is suspiciously well developed in several species of leaf-nosed bats and, along with such obvious factors as availability of roost sites and synchro¬ nization of reproductive activities, the thermoregulatory significance of such behavior merits investigation. The climatic history of New World leaf-nosed bats during their evolution seems to have allowed considerable latitude in the degree of temperature control. Thermal stresses comparable to those encountered in temperate regions have not been present in sufficient strength to force the develop¬ ment of precise homeothermy, nor have they caused the adoption of “deep torpor” capabilities typical of bats at higher latitudes. One broad generalization that can be made with respect to the family Phyllostomatidae is that they are extremely diversified in details of their temperature control. BIOLOGY OF THE PHYLLOSTOMATIDAE 291 Acknowledgments I am indebted to Diane Ruffino for typing several drafts of this manuscript. Many of the ideas incorporated into this review are based on the studies of Brian McNab, and I urge a perusal of his papers for fuller treatments of several aspects of bat energetics included herein. Literature Cited Arata, a. A„ and C. Jones. 1967. Homeoihermy in CttrollUt (Phyllostomatidae: Chi- roptera) and the adaptation of poikilolhermy in insectivoroiLS northern bats. Lo^.ania (Acta Zoologka Columbiana), 14:1 -10. Barthoiomew, G. A-, P. Lbitner, and J. E, Nelson. 1964. Body lemperalure, oxygen consumption, and heart rate in three species of Australian flying foxes. Physiol. ZooL, 37:179-19H. Bradshaw, G. van R. 1961. A life history study of the California leaf-nosed bat, Mcurotns californicHs. Unpublished Ph.D, dissertation, Univ. Arizona, Tucson, 89 pp. Bkodv, S. 1945. Bioenergetics and growth. Reinhold Publ. Corp., New York. xii + 1023 pp. Burhank, K. C.. and J. Z, Y'oljnc. 1934. Temperature changes and winter sleep of bats. J. Physiol., 82:459-467. Carpenter, R. E., h\nd J. B. GRAttAM. 1967. Physiological responses to temperature in the long-nosed bat. Lvptofiycferis stifihorni. Comp. Biochem. Physiol., 22:709- 722. Crespo, R, F.. R. J. Burns, and S. B. Linhart. 1970. Loadlifting capacity of the vampire bat. J. Mamm., 51:627-629. Hook, R, J. 1951. The metabolic rales and body temperatures of bats. Biol. Bull.. 101:289-299. Hudson, J. W. 1973. Torpidity in mammals. Pp. 97-165, in Comparative physiology of thermoregulation (G. C. Whittow, ed.), Academic Press, New York, 3;xii+ 1-278. Jan ZEN. D. H., and T. W. Schoener. 1968. Differences in insect abundance and diversity between w etter and drier sites during a tropical dry season. Ecology, 49:96-110. Kleiber, M. 1961. The fire of life. John Wiley, New'York, 454 pp. LaVal, R. K. 1969. An example of unusual hehavior and temperature depression in ///nrfitjo (PhyllostomidaeL Bat Res. News, 10:42. Leitner, P., and S. P. Ray. 1964. Body temperature regulation of the California leaf¬ nosed bat, Mac roUis culifor/iicus. Amer, ZooL, 4:295, Lyman, C. P. 1970. Thermoregulation and metabolism in bats, Pp. 301-330, in Biology of bats (W, A. Wimsatt, ed.). Academic Press, New York, l:xii+ 1-406. Lyman, C. P., and W. A. Wimsatt, 1966. Temperature regulation in the vampire bat. Desmodus rotnndus. Physiol, Zool., 39:101-109. McFarland, W. N., and W. A. Wjmsatt. 1969. Renal function and its relation to the ecology of the vampire bat. Desmodtis roiundti\. Comp. Biexhem. Physiol., 28; 985-1006. McManus. J. J., and R. J. Esher. 1971. Notes on the biology of the little brow'n hat, Myoiis hibernating in New Jersey. Bull, New Jersey Acad. Sci., 16: 19-24. McManus, J. J., and D. W, Nellis. 1972. Temperature regulation in three species of tropical bats. J. Mamm,, 53:226-227. McNar, B. K, 1969. The economics of temperature regulation in neotropical bats. Comp. Biochem. Physiol., 31:227-268. -—. 1970. Body weight and Ihe energetics of temperature regulation. J. Exper. Biol., 53:329-348. -. 1971. The structure of tropical bat faunas. Ecology. 52:352-358, 292 SPKCIAL PUB 1.1 CAT IONS MUSEUM TEXAS TECH UNIVERSITY --. 1973. EnergetiLSi and ihe disiribuiiun of vampires. J. Mamm,. 54:13}-U4. MokRisoN, P., AMI B. K. MlNais, 1967. Temperarure regulation in some Brazilian phyU lostomid bats. Comp. Rioehem, Physiol., 21:207-221. RASVVtiii.R, J. J,, IV, 1970, The laboratory biology of the long-tongued bat. dossopha^u soridfur. maimenartce procedures, estivation, the menstrual cycle, histophysiology of the oviduct, and intramural implantation. Unpublished Ph.D. dissertation, Cornell Univ. REt.OER, W. G., AND R. B. Cowi ES, 1951. Aspects of thermoregulation in bats. J. Mamm., 32:-38 9-403. Stones. R. C., and J. E. Wieders. 1965. A review of temperature regulation in bats (Chiroptera). Amer, Midland Nat., 74:155-167. Stl’DIF-R, E. H., .and D. E. Wii son. 1970. Temperature regulation in some Neotropical bats. Comp. Biochem. Physiol., 34:251-262. Terth , R, 1972. The effect of behavioura] thermoregulation on the daily metahotism of .4/NJt/eNNo ujirrur/nv (Pallas, 1771). Acta Theriol, 17:295-313. Twente, J- W., jR. 1955. Some aspects of habitat selection and other behavior of cavern- duelling bats. Ecology, 36:706-732. Wai ker. E. P., i't ill. 1968, Mammahs of the world. Johns Hopkin,s Press, Baltimore, l:‘?-(- 1-644. Wuinow, G. C. (ED.). 1971. Comparative physiology of thermoregulation. Academic Press, New York, 2:xi+ 1-410. Whitiow, G. C. 1973. Evolution of thermoregulation. Pp. 201-258, in Comparative physiology of thermoregulation (Ci, C. Whitlow, ed.). Academic Press, New York. 3:,Aii+ 1-278. WiMSA i T, W. A, 1962. Responses of captive common vampires to cold and w arm environ¬ ments. J. Mamm., 43:185-19L (Editors' note: John J. McManus died on 22 August 1975 without having an opportunity to read galley proofs or correct any errors inadvertently incorporated, or overlooked, in bis contribution to this volume.] FEEDING HABITS Alfred L. Gardner There is something fascinating about science. One gets such wholesale returns of conjecture out of such a trifling investment of fact, Mark Twain With few exceptions, knowledge of bat food habits, like that of many other aspects of chiropteran biology, is superficial or w'anting. Nevertheless, the study of bats sometimes has been stimulated because of the interest generated by their unusual or economically important food habits: the sanguivorous diets of vam¬ pires, flower*feeding habits of glossophagtnes, and the earn ivory of some phyl- lostomatines. General knowledge of food habits w'as used in some of the early dassiftcations of bats. Gray (1821:299) divided his class Cheiroptera into tw'o orders, the Fruc- tivorae and the Insectivorae, Along the same lines, Koch (1862-1863:298) erected the two suborders Carpophagen and Entomophagen, and Gill (1872, 1886) separated the Chiroptera into the suborders Animalivora and Frugivora (again representing major differences in food habits). In these examples, the In¬ sectivorae, Entomophagen, or Animalivora included all of the known forms of the Phyllostomatidae. The Fructivorae, Carpophagen, or Frugivora arc equivalent to the Old World fruit bats, the Megachiroptera. Other names applied to members of the Phytlostomatidae reflect known or alleged feeding habits. The generic names Vampymnu Vampyrops, Vampyrodes^ and Vampyressa refer to the alleged blood-teeding habits of vampires. Dtaemus means blood-stained, an appropriate name for a true vampire. Ghssophaga lit¬ erally means to eat with the tongue. Lichonyctens means a bat that licks, also in reference to using the tongue w'hen feeding. Mimmycteris implies an associa¬ tion with banana plants. The name Anoum werckleai' was employed to reflect the feeding relationship of this bat with the plant Wercklea lutea. The trivial name mordax refers to biting {LDnchophylla mordax and Stitrnira tmmULx), Common names often refer to presumed food habits as well. Some of these are the “Cuban Fruit-eating Bat’* {Brachyphyila nana), the “Hairy Fruit-eating Bat” {Anibem hirsiittis)^ and the “Cuban Flower Bat” {PhyiUmyaeris poeyi), A few names suggest diets when, in fact, the foods consumed are not known (for in¬ stance, the “Red Fig-eating Bat," Stemxierma nifum\ the “Brown Flower Bat," Erophylla honibifrons). My principle objective in surveying the food habits of the Phyllostomatidae was to bring together most of the available published information on the diets of these bats in a form that not only will provide an accessible information source, but also will encourage future investigations on this aspect of bat biology, I have reviewed most of the accessible literature with disappointing results. With few exceptions, very little has been recorded on the diets of these animals, and much 293 294 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY of this is superficial and non in formative. The information presented in the species accounts deals almost exclusively with the diets of free-living bats as reported in the literature. A few personal observations on food habits as w-ell as some litera¬ ture references on foods consumed in captivity have been included when con¬ sidered pertinent, although the chapter. Care in C'aptivity, by Greenhall ade¬ quately covers the latter subject. Inferences on diets as suggested by dental and alimentary tract anatomy are equally restricted because the chapters, Oral Biology, by Phillips ci and Ciastro-inteslinal Morphology, by Forman and Rouk, provide ample information on these aspects as w'ell. The diet for each species follow's the scientific name in the food habits accounts below. Table I is a list of those plant genera and species for which parts have been reported as foods consumed by phyllostomatid bats. PROBLiiMS IN Determining Food Habits if we could observe and record the variety and quantity of foods as they are gathered and consumed by bats, the determination of diets w'ould be a relatively simple matter. Because this usually is not possible, the examination of feces or digestive tract contents would appear to be the next best method. However, the comminuted remains of insects and small vertebrates are usually difficult to identify; a problem intensified by the habit of many bats to discard the harder, and often the only diagnostic, parts of their prey. For example, the abdomens of lepidopterans and other large insects are often the only parts consumed, the other parts being discarded. The taxonomy of many of the insects consumed by bats is little know'n and reference collections of insects from areas where the bats were collected are usually not available. Therefore, the determination of the insect order or family may be the only identification possible from fecal or stomach contents. Masticated remains of fruits found in stomachs are almost impossible to identify if associated seeds are not present. Seeds found in stomach contents and feces also are difficult to identify, particularly w ithout the aid of a comprehensive reference collection of seeds. Nevertheless, fruits are often emphasized w'hen diets are reported because when seeds are available, they are usually easier to identify than insects. This is especially true when the fruits come from locally conspicuous and well-known plants. Some items such as seeds, fruit, or bits of sand or gravel found in the stomachs of omnivorous and carnivorous bats can be misleading because they may have been consumed by an animal before it was ingested by the bat itself. Stomach content analyses also can be deceptive if the bats are maintained alive together in small cages or cloth bags subsequent to their capture. Fighting and cannibal¬ ism among hats held under these conditions is common, and finding blood or the remains of bats in the stomachs of these bats should not be considered as indica¬ tive of normal diets unless, of course, the bats are vampires or species with known carnivorous habits. Detailed analyses of stomach contents can be very informative. The excellent study by Alvarez and Gonzalez Q. (1970) demonstrated that analyses of the di¬ gestive tracts of pollinivorous species not only indicate what flowers are being BIOLOGY OF THE PHYLLOSTOMATIDAE 295 Tahie I .—of plu/m fin wfifch priHiiH'ts are knomt to he inciuttiii in the iticfs of phyl- hsuunaiid hat.\. Acacia Achnis sapoia Avnhtas A a'oconiUi aculeata Ascnve scholtii Aihfzzia Aicun gramiiflora A hi ns AfiiH^ardiiim exvelsi/m Anacardinm occidentafc Anamo/nis nnihc'llIIIiferu Andira inermis Annona mnricaia Annona pi son is Atotona sqnanio.sa Anoda Amhtoiatn Arbutus Aristoloihia A riocarpus iutcgrifoiia Baetn's Baiihinia pauiethi fieiischniiedia Berber is Hi mi h a x fioupuinvi!lea spectaldlis Bros imam alicastrum Bur sera By rsouim a sp ica ta Caesalpinia Cidliaudra Cali >ci trpuni matnniosutu Cahipbylluni hrasiitense Calycolptis wuirsze ^ciczianns Car ica papaya Carl udov tea pal mat a Carnepiea gigantea Casi/niroa edit!is Cecrop ia hurean ia/ut Q' crop ia oh I us ifol ia Cecropia peltafa Ceiha pentandra Ceufrttpogon Cere us hexagon us CenH ' iirpus { = Eugenia ) disticus Ch lorophoni tinetoria Chrysaiidocarpus { Areea } lutesveus Ch ry so ha fa > \ us lea co Chrysophyilum ca ini to Citharexyium Coccotoha iiv 'tfera Leguminosae, Mimosoideae Sapotaccae Solanaceae Palmae Amaryllidaceae Leguminosae, Mimosoideae Leguminosae, Papilionoideae Betulaceae Anacardiaceae Anacardiaceae Myxlaceae Leguminosae, Papilionoideae Annonaceae Annonaceae Annonaceae Malvaceae Araceae Ericaceae Arisiolochiaceac Moraceae Palmae Leguminosae, Caesalpinioideae Lauraceae Berberidaceae Bombacaceae Nyciaginaceae Moraceae Burseraeeae Malpighiaceae Leguminosae, Caesalpinioideae Leguminosae, .Mimosoideae Sapolaceae Guuiferae M yrtaceae Caricaceae Cyclanthaceae Caciaceae Rutaceae Moraceae Moraceae Moraceae Bombacaceae Campanulaceae Cactaceae Myriaceae Moraceae Palmae Polygonaceae Sapotaceae Verbenaceae Polygonaceae 296 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNJVERSITY Table 1.— Co/uittneJ. Coicothrituix Coffea Conztifiki Cordiu hicofor Corditi coltoctHTii Coni in JtHleCiifHlni Cniiiiex'ii he/t(iiai>ii Crescentid anuizifniai Crexcetuhi cnjen' Croto/i Cynoffu’tra n'msn DuUtru Dendropnmi.x nrhoreus Dioxpyri>x iliffy-fu! Dii *J py rox eh e i ms! er Diospyros kiik i biospyros tnnhoht Dipieryx odontta Durio zihdhhuts EvhiniHiietifS E!izah e! ini pa me/i se Epertm falcma Epipftyllno} hookei i Erit)ho! ry u japonivn Encnfypin.s Engeniti jittnlufs Edge n HI tsnilurce fix is Eugenki tieskti ica Eugenia n/iijiora Fu ns heiiiami/ui Fii ifS aginijhlia Finis gliihrma Fk tis insipidu Ficus obttisififlki Finis padifolki Ficus mdida Fk ns religkisa Fk ns reiiisn Fliicourtia indica Genipa Gemiaiui Geranium HeisH'fiit Hihisnis Htihenhergkt Hy hcerei ts / e mu iret Hyn I rmea conrbnrii Ingu ipomoeu urhoreu h iarteu exorrhiiu Jumhitsu vtdg(trisi= Engeniu jamhos) Kigeim ueihiopk n Palniae Rubiaceae I.egiiminosae. Caesalpinioideae Boraginaceae Boraginaceae Boraginaceae Capparaceae Btgnoniaceae Bignoniaceac Euphorbiaceae Leguminosae, Cacsalpinioideae Solanaceae Araiiaceae Ebenaceae Ebenaceae Ebenaceae Ebenaceae Leguminosae. Papilionoideae Bombacaceae Cactaceae Leguminosae, Caesalpinioideae Leguminosae, Caesalpinioideae Cactaceae Malaceae Myriaceae M yrtaceae Myrtaceae M yrtaceae Myrtaceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Moraceae Flacourtiaceae Rubiaceac Gentianaceae Geraniaceae Olacaceae Malvaceae Bromeliaceae Cactaceae Legu mi nosae, Caesalpin ioideae Leguminosae, Mimosoideae Convolvulaceae Palniae Myrtaceae Bignoniaccae BIOLOGY OF THE PHYLLOSTOMATIDAE 297 Table L- — Co/uitiued. Liiftianu lA'iyihii Ziihuctijo Lcnutireocereuii Liainia Liviitonn chine IIS is Lonchoearpus Lifctiiftti { = PiiKieria) titimiuf Miufltucii laiifolid M til p ;X' / 1 1(1 g hthrti Mah’ti vise Its (tariftilius .X'fiiinnu’u timericiiiiti Mangifera i/ulica Mtill if k tiru hitienfiiin Mtinilkarti zapmn Maffgrtixha Xfelicocea hijitga Miffiosa M i nut St )ps eleng i Xfonts ( = Cftlorophora) iinciorUi Xfiicuiia a/!(lrearui Muiitiiig in viilahum Mimi piiradisiactt My n • in job o f k -a Im Myr/illoaicfits Oehroiutt iagopus Oenofheni Park id giguti/ticarpti Park id pc/ulufa Partnentiera alaia Passijlora (/mtdratigtiiaris Pcrsea americana Pilocarpus p inmn ifolius Piinentu racemosa Pi tuts Piper amulago Piper aurititm Piper hispiiittm Piper sanctum Piper tiihercniattun Pitcairnia PkityopHtnia Pouisenia urrnata Poiiterki maltijlora Pseadohombax Pse udohned ia oxyphyliaria Psidiutn giudava Psiditim medkerranetint Parpurethi grossa Putranjivora roxhnrghii Quaroribea fitnehris Qiiercus Rheedia edit Us Verbenaceae Lecyihidaceae Cactaceae Chrysobalanaceae Pal mae Leguminosae, PapiMonoideae Sapotaceae Sapotaceae Malpighiaceae Malvaceae Guttiferae Anacardiaceae Sapotaceae Sapotaceae Marcgraviaceae Sapindaceae Leguminosae, Mimosoideae Sapotaceae Moraccae Leguminosae, Papilionoideae Elaeocarpaceae Musaceae Myriaceae Cactaceae Bombacaceae Onagraceae Leguminosae. Mimosoideae Leguminosae, Mimosoideae Bignoniaceae Passifloraceae Lauraceae Ruiaceae Myriaceae Pinaceae Pipcraceae Piperaceae Pipcraceae Piperaceae Piperaceae Bronieleaceae Caciaceae Moraceae Sapotaceae Bombacaceae Moraceae Myrtaceae Myriaceae Melastomataceae Euphorbiaceae Bombacaceae Fagaceae Guttiferae 29S SPECJAl. PUHLlCA'l'tONS MUSEUM TEXAS TECH UNlVEkSJTY Table 1. — Cottii/un'tl. Roupa((! Proteaceae RoyMoneti o!vritvi’ci Palmae Ruhachiii ( = MarliL'n'a] Myrtaceae Self Lx Saliaceae Siilvia l.abiatae Supiiuius stipoiuiriis Sapindaceae Sidcro.xyfon quadrioi uhire Sapotaceae St if at} u ni pa/iiiulaiunt Solanaceae Sp (1 nd fa s cy t /i erea Anacardiaceae Spo ltd ills Isftta Anacardiaccae Spo/itfkis uiiiiitfiifi Anacardiaceae Spo/idius purpurea Anacardiaceae Slemmadetiiti Apocynaccae Sy/nhoiaudius iuiifoisus Gentianaceae Syzi},ditnj (= Eapeitki) jauthos Myrtaceae Tertninal ta cat up pa Combretaceae TheifhroiiU! cacao Sterculiaceae Turps/iks pinnata Staphylaceae Vixmiti fatifofki Hypericaccae Viiis vifii/era Vitaceae yochyskt Vochysiaceae Wercklea (utcu Malvaceae Zca Graniinae Zinpiher Zingiberaceae visited but can provide evidence on the movements of popiiIationSj seasonal changes in diets, and competition for the same foods by sympatric species. The recovery and identification of pollen grains from the fur of bats may indicate the flowers visited as w'ell (Heithaus et af, 1975; Howell and Burch, 1974). Analyses of blood meals in the stomachs of vampires have indicated if mammals, birds, or both w'erc the prey (Villa-R. a al., 1969), When names of the food items of bats are reported (Greenhall, 1956; Goodwin and Greenhall, 1961; Wilson, 1971; Vazquez-Yanez et «/., 1975), they most often are based on identifications of dropped or discarded parts of plants, insects, or vertebrates found associated with bat roosts. Here the problem is to associate the remains with the bats that left them. Correctly associating food remains is difficult if the roost sites w'here the remains were recovered are used by one or more species during the day and by yet other species at night. Observing, photographing, or collecting bats as they feed on flow'ers or fruit arc ways to associate species wdth the foods they eat. Another w-ay is to identify items the bats are carrying when captured in mist nets. Among the Phyllostomati- dae, the commonest focxl items found in nets are fruits; however, Valdez and LaVal (1971) recovered an Anolis leniurinus after it had been carried into a net by a Tmehops cirrhosus. BIOLOGY OF THE PHYLLOSTOMATIDAE 299 Food Habits The PhyJlostomatidae display a wide variety of food preferences, and relatively few species are restricted to a specific dietary regime. Only piscivory, among the types of chiropteran food habits, has not been found in the New' World leaf-nosed bats. The majority of the Phyllosiomatinae are omnivores; however, some have strong tendencies toward carnivory. The only true insectivore in the subfamily (and family) may be Macrophylium macrophyitum, Vampyrum spectrum and Chrotopterus auritus^ both principally carnivorous, prey upon a variety of small vertebrates, including bats. Fruits and flowers are important components in the diets of many phyllostomatines, and some species, such as Phyilosmmus hastatus^ serve an important function in flower pollination (chiropterogamy) and plant dispersal (chiropterochory). Most phyllostomatines have the ability for hovering flight and, as suggested by their relatively large ears and eyes, probably are able to detect and capture prey on the ground or from foliage, tree trunks, and other surfaces. The diets of the Glossophaginae include pollen, nectar, and, occasionally, parts of the corolla of flowers. As specialized flower feeders, these bats also play an important role in chiropterogamy. The majority is known to consume a variety of fruits, and some are suspected to pursue actively insects in addition to eating those captured at now'ers. Wind borne pollen {Alnus and Pinus, see Alvarez and Gonzalez O., 1970) has been found in the stomachs of Glossophaga, Lep- tonycteris^ Choeronycteris, and Anoura. These pollen grains probably were in¬ gested from flowers and watering places where they settled (on pools of water, in bromeliads, and in cavities or depressions in trees). The Carolliinae and Stenoderminae perhaps are best considered as frugivorous and some species indeed may be obligate frugivores (for example, Pygodemw, Ametrula, and Centurio). Many, how-ever, also consume flower parts, ptillen, nectar, and insects, particularly the Carolliinae, which arc known to consume quantities of insects (Fleming et u/., 1972). Stenodermines undoubtedly con¬ sume insects in the course of eating fruit because many fruits contain insect larvae. Frugivorous bats, as pointed out by Ayala and D'AIessandro (1973) and Forman (1973), probably must consume large amounts of fruit because adequate proteins, fats, and minerals are not abundant in this food. Several bats have been caught in fruit-baited traps placed on the ground and their capture supports the observa¬ tions of Jimbo and Schw'assmann (1967) that bats will feed on ripe fruit that has fallen to the ground. Anibeus has been reported to capture insects (Tuttle, 1968) and eat nestling birds (Ruschi, 1953y3; other stenodermines may prove to be in¬ sectivorous or carnivorous. The Phyllonycterinae are an assemblage of fruit, pollen, nectar, and insect¬ eating species restricted in distribution to the Antilles. The limited information available on their food habits indicates a preference for pollen and insects, although soft fruits are eaten as well. The Desmodontinae are obligate sanguivores and, within the Phyllostomatidae, have the most specialized dietary requirements. Insects and bits of flesh have 300 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY been found in the stomachs of Desmodns nuundus (Arata e( al., 1967; Greenhall, 1972). i'hese items, however, were most likely ingested during bite site prepara¬ tion or when feeding on the host. The ectoparasite reported by Arata ei aL {1967) in a Desmodus stomach probably was consumed during grooming activities. Diaemus and, to a lesser extent, Diphyila show' a preference for bird blood. Des- /iiodiis preys on birds as welt, but apparently prefers mammalian blood. Population densities of vampires probably have increased owing to the readily available food source supplied by domestic livestock. Before the availability of livestock and the widespread use of mosquito netting, vampires (particularly Demiodi{s) may have depended on human populations as a major food source. Cashinahua Indians living at Balia, Departamento de Loreto, Peru, an area with very few Desmodus and no domestic livestock, like mosquito netting as much for keeping away vampires as for protecting against insects. These people have been using mosquito netting for relatively few' years and clearly remember having been bitten by bats in the past. Subfamily Phyllostomatinae Genus Micronycteris Gray Micronycteris megalotis A variety of insects and fruits. Gaunter (1917), observing that M. tnegidoiis tlew slowly and near the ground in Yucatan, Mexico, surmised that they ate insects caught close to the ground. Ruschi (1953f/) mentioned insects and the fruits of Musa paradisktea, Psidium gaajava, Jamhosa vulgaris, Cecropia sp., Erioholtya Japan lea, and Salanimi paniculaUim as part of the diet of Brazilian M, megalotis. Guavas (Psidium gimjava) were also reported as a food item of this species in San Luis Potos'i, Mexico, by Dalquest (1953), w'ho believed that they probably feed on many kinds of fruit. He found them feeding on small guavas, which they plucked and carried off to a nearby tree to cat, often dropping and losing much fruit in the prcKCss. Goodwin and Greenhall (1961) classified M. /negalatis as a fniit-eating bat ap¬ parently fond of small ripe guavas, yet noted both insects and yellow' fruit pulp in the stomachs of some specimens from Trinidad. The stomachs of specimens taken during the daytime in Veracruz, Mexico, were empty; how'ever, the stomachs of two collected at night w'ere filled with the remains of insects (Hall and Dalquest, 1963). Howell and Burch (1974) reported that a Costa Rican specimen had con¬ sumed an “unknown green fruit.” The categorization of M. megahris as a nectar¬ eating species by Valdivieso and Tamsitt (1962) appears to be unsupported. Micronycteris schmidtorum Insects and probably fruit. Howell and Burch (1974) reported two Costa Rican specimens of M. schmidior- urn that had consumed Lepidoptera. BIOLOGY OF THE PHYLLOSTOMATIDAE 301 Micronycteris minuta Insects and fruit, Goodwin and Greenhali (1961) believed M. mbuaa to consume fruit or insects or both. Fleming et al. (1972) examined 12 individuals from Costa Rica and Panama and found 76 per cent insect and 24 per cent plant material, by volume, in the stomachs of four. Micronycteris hirsuta A variety of insects and fruits. Goodwin and Greenhali (1961) considered this species to be fruit eating but said it may also consume some insects. Fleming ei al. (1972), reporting on the stomach contents of three Panamanian specimens, found only the remains of insects. Wilson (1971) reported on the food remains he gathered at intervals between January and July from under roosting sites located on Orchid Island in the Panama Canal Zone. The major insect food items found represent the families Blattidae, Tettigoniidae, Scarabaeidae, Cerambycidae, Curculionidac, Cicadidae, Saturniidae, Sphingidae, Aeschnidae, and Formicidae, The insects recovered are winged forms capable of flight; however, they spend much of their time moving about on vegetation at night, suggesting that M. hirsuici may be gleaning them from vegetation as well as taking them in flight. The majority of the insect material, primarily whole wings, pieces of legs, and other hard parts, consisted of cockroach¬ es (Blattidae), katydids (Tettigoniidae), and June beetles (Scarabaeidae). The remains of fruits recovered from the roosts represent Carludovka palmata., Piper sp., Beilschmiedki sp,, Amicardhim exceiaum, Vismia iatifolia, Passijlora sp., Calycolpus warszewiczianas, and Eugenia nealoika. Wilson concluded that M. hirsuta is primarily insectivorous and that the small quantity of fruit eaten is con¬ sumed mainly in the dry season (February to March) when fruits are abundant. How'eil and Burch (1974) reported the remains of Lepidoptera in a Costa Rican specimen. Micronycteris brachyotis Insects and fruit. Goodwin and Greenhali (1961) reported finding fruit pulp, plant fibers, and insects in the stomachs of M. brachyotis from Trinidad. They also reported a “white milky substance” in the stomach of a juvenile. This individual probably was still nursing. Vil!a-R. (1967) noted that this bat feeds on pulpy fruits and insects as do other members of the genus. Howell and Burch (1974) mentioned insect remains (Hymenoptera and Coleoptera) recovered from a Costa Rican M. brachyotis. Micronycteris nicefori Fruit and insects. A diet of fruit and possibly some insects was proposed by Goodwin and Green- hall (1961). 302 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Micronjcteris sylvestri** Fruit and insects. A diet of fruit and possibly some insects was suggested by Goodwin and Green- hall (1961). Summary .—^Apparently there is no information available on the food habits of Micronyacris pusilliU M. hehni, and M. duviesL According to Duke {1967) species of the genus Microuyaerh are primarily insectivorous, although in Panama they may be secondarily frugivorous. Walker et ai. (1964) stated that the molar pattern is indicative of an insectivorous diet. The reports by Wilson (1971), Hall and Dalquest (1963), Fleming e{ ai (1972), and Howell and Burch (1974) support the contention that insects are the primary food source of these omnivorous bats, A variety of fruits are consumed by MU ro- nycteiis\ however, their importance in the diet probably varies seasonally, as W'ilson (1971) found in Panamanian M. hirsuia. Genus Macro rus Gray Macrotus waterhousii Large insects and fruit. Osburn (1865:74) reported finding the wings and legs of large Orthoptera under a roost in a cave in Jamaica and mentioned that bats, presumed to be M, waterhinmi would drop the remains of the fruits of Moras tmetoria^ Brosimuin allcastrum, and Eugenia jambos Uom their night roosts. He also described (p, 75) a female killing a nursling bat (not hers, but one that was placed on her) and consuming its blood. This incident may have prompted Dobson (1878) to in¬ clude small bats along with insects and fruit as food items of M. waterhousii. Dob¬ son said the stomach of one specimen contained a yellowish mass with harder parts of insects, including the remains of orthopierans, Macrotus californicus A large variety of larger night-flying insects, some nonflying insects such as lepidopterous larvae, some fruits, and possibly green vegetative matter. The remains of beetles of the species Ligyrus gibhosus^ Chtaenias serkeus, and Poiyphytla decemiUneata, plus parts of “various species of flies,” were reported by Grinneli (1918:256) as scattered over the floor of a cave inhabited by M. califonucus in southern California. She also cited an incident {p. 257) where a Macrotus was caught in a mouse trap set in the open desert and suggested that this species seeks some of its food on the ground inasmuch as the bat likely was caught while attempting to capture ants or beetles attracted to the trap bait. This account probably was the basis for Sanborn’s (1954) mention of a specimen taken in a mousetrap. Additional reports on the food habits of M. vaiifornicas are also based on material gathered from under roosts in southern California. Howell (1920) found the wings of several diurnal butterflies, as w-ell as parts of moths. Huey (1925) reported finding a willow leaf and the remains of grasshoppers BIOLOGY OP THE BHYLLOSTOMATIDAE 303 {Trifnerotropis sp, and Schi.stocerca sp.), cicadas, beetles (Meloidae), Sphinx moths {Celeiio fi/teaia and Smerinthus cerisyi), a nocluid moth {Perkiroma margiirifosa), and a cossid moth below the roosts. He assumed that some of these insects were diurnal and suggested that they had been taken from their resting places on vegetation (willows) by the bats and then carried to the roost to be eaten. Vaughn (1959) noted the remains of moths, butterflies, and dragonflies found under roosts. He also found fragments of orthopieran insects, ncKtuid moths, caterpillars, and beetles (Scarabaeidae and Carabidae) in the stomachs of several M. caUfornicus and concluded that this species w'as totally insectivorous, a supposition echoed by Novick (1963), Villa-R. (1967), Anderson (1969), and Barbour and Davis (1969). Supported by information in earlier reports and by finding the remains of caterpillars in stomachs, Vaughn (1959) also contended that Macrolus mainly lakes insects that are on sparsely foliated vegetation or on the ground. Burt (1938) reported on the stomach contents of five Macrofus taken in Sonora, Mexico, Two had fruit and insects in their stomachs and three contained only fruit. Park and Hall (1951) treated Macrotus as a frugivore in their re¬ port on the tongue and stomach anatomy of several New' World bats, Ross (1967: 214) cited observations which mentioned that M. califomkus feeds on various cactus fruits. He also repo^ried on the insect remains gathered beneath night roosts and on his analyses of 41 digestive tracts, mostly from bats collected in the vicinity of Tucson, Pima County, Arizona, but including a few from Sonora, Mexico. The insect remains associated with roosts represented desert short- horned grasshoppers (Acrid idae: Trlmerotrapis sp.), long-homed grasshoppers (Tettigoniidae: Microcentrum californk'ux, Schistocen a vaga^ and other species), long-horned beetles (Cerambycidae: Derohrachns geminaius), Sphinx moths (Sphingidae; Celerio lineafa), and underwing moths (Phalaenidae: Catocaia sp,). The stomach contents varied from purely insect remains to purely vegetable matter. Some stomachs from winter-taken M. caUfornicus contained w-hat ap¬ peared to be green vegetative parts of plants. Macrotus cxtlifornicus feeds primarily on the abdomens of larger night-flying insects within an average size range of 40 to 60 millimeters in length and, to a lesser extent, on lepidopterous larvae and other small insects approximately 20 millimeters in length, such as short-horned grasshoppers (Acrididae) and June beetles (Scarabaeidae) (Ross, 1967:211), Ross, disagreeing with the conclusions expressed in earlier reports, claimed that most, if not all, of the insects preyed upon by this bat are nocturnally active forms. He also asserted that no truly ground dwelling forms of insects w'ere found in any of the digestive tracts he examined. Vaughn’s (1959:34) observation that Macrotus regularly forages close to the ground, seems to hover easily, and is able to hover for several seconds at a time, suggests that these bats may glean insect prey from the ground and from vegeta¬ tion, as well as capture flying insects. Therefore, M. caUfornicus probably does include some flightless, ground-dwelling, or diurnal insects in its diet, as appears obvious from finding caterpillars in digestive tracts. 304 SPRCIAI- PUB1,ICAT10NS MUSEUM TEXAS TECH UNIVERSITY Genus Loncuoriiina Tomes LonchorJitna aiirita Insects and plant material. Ruschi (1953 p) stated that aNriia eats insects exclusively. An examination of the stomach contents of two specimens from Trinidad only revealed the re¬ mains of insects (Goodwin and Greenhall, 1961). Duke (1967), citing unpub¬ lished information from Edwin Tyson, stated that this species probably eats nectar, some insects, and overripe fruit in Panama. Fleming et al. (1972) examined tw'o stomachs of Panamanian L. auriia. One contained about equal quantities of fruit pulp and insect remains. Nevertheless, Fleming e( ai. (1972: 560) considered T. aurifa to be primarily insectivorous. Howell and Burch (1974) agreed after finding the remains of Lepidoptera in a Costa Rican specimen, Lonchorhina orinoeensis The food habits are unknowm, but the diet is probably similar to that of aurim. Genus Macrophyllum Gray MacrophyJlum ]tiacT<»phylliiiii A variety of insects. Oueich (1892) believed M. maaophyihim to consume insects; how'ever, on the basis of the large incisors, he also suggested that this species may supplement its diet w'ith blood. Insects and fruit were mentioned by Ruschi (1953/) as foods eaten by Brazilian M. mavraphyllum. Davis ef al. (1964:378-379), commenting on the foraging behavior, body weight, and proportional size of the feet of this species in Nicaragua, suggested that aquatic insects or small fish w-ere included in its diet. The stomachs of the specimens they collected w'ere empty. Duke (1967) stated that Edw'in Tyson thinks Macrophyllum eats swimming insects. Harrison and Pendleton (1974:691) reported finding the stomachs of four Salva¬ doran Mmrophyllum “full of dark brownish, finely masticated material." Wing fragments, w'hich they suggested represented lepidopterans and dipterans, as well as many lepidopteran wing scales w'ere found among the stomach contents. The stomach contents of tw^o Macrophyiluai that I examined from Panama pri¬ marily consisted of the remains of water striders (Hemiptera, Gerridae; cf. Trepohafes), which also appeared as browmish, finely chewed insect remains. Genus Tonatia Gray Tonatia bidens Fruit and insects. The diet probably includes fruit and insects as w'as reported by Ruschi (1953^?) for this species in Brazil. Goodwin and Greenhall (1961) stated that tt eats fruit. BIOLOGY OF THE PHYLLOSTOMATIDAE 305 Tonufia brasiljensis Probably fruit and insects, A diet of fruit and insects was proposed by Ruschi (1953t'); however, the identification of his specimens is open to question because the "^Touatia brasiHeusi!;" illustrated clearly is a CaroUia, Tonal ia sylvkola Fruit and insects. Only 1 1 stomachs of the 22 Panamanian T. sylvicoki reported on by Fleming et «/. (1972) contained food, all of which was the remains of insects. How'ell and Burch (1974) recovered legume pollen and the remains of fruit {Stemmadeniu) from two Costa Rican representatives of this species. Summary .—The diets of Tomuki vaniker!., T. nicaraguae, and 7. veuezutdac are not known, I suspect that species of Tomitkt consume a large variety of arthropods, both flying insects and those gleaned from vegetation and other sub¬ strates. Tyson (quoted by Duke, 1967:8) believed Tomuia to be insectivorous, and thought that it probably gleans insects “off twdgs about w'hich they hover,“ Toruuia may consume a variety of fruits as well (see How'ell and Burch, 1974) and probably has food habits similar to those of Micronyaeris. Genus Mimon Gray Mimon bennellii Fruit and insects. The diet is reportedly insects and fruit (Ruschi, 1953c). Minion co/umelae Plant material and various arthropods. Dalquest (1957:46), commenting on several M. cozumelae he saw' flying around half-spoiled fruit in an orange grove in southern Veracruz, Mexico, sug¬ gested “they may have been eating the fruit, fermented juice, or insects stupefied by the juice.“ Hall and Dalquest (1963), perhaps reporting on the same incident, staled that M. tozufnelae ate only very ripe, sometimes spoiled oranges, or pos¬ sibly the insects that were feeding on the overripe fruit. They commented that the white droppings littering the floor in caves inhabited by M. cozumelae and Travhops cirrhosus resembled the droppings of haw'ks and owls, and concluded that both genera of bats are probably somewhat carnivorous, Villa-R. (1967) reported that M. cozumelae apparently eats fruit. Mimon crenulatum Insects. Dobson (1878) reported finding portions of small coleopterous insects in the mouth and throat of a specimen. This information was apparently repeated with- out citation by Walker et ai (1964). 306 SPECIAL PUBI ICATIONS MUSEUM TEXAS TECH UNIVERSITY Siffnnuiry. —There is no informaiion on the food habits of Minujn k(K^pckt'cie. Species of the genus Mimou probably consume a variety of arthropods and fruits. Genus Phyllostomus Lacepede Phyllostonius diseulor Insects, fruit, pollen, nectar, and vegetative parts of flowers. Van der Fiji (1957:294, citing correspondence from Heinz Felten) noted that remnants of the fruit of Spondias purpurea were commonly found under the roosts of P. discolor in caves in El Salvador, Observations on bats visiting flowers in the Parque do Museu Goeldi, Belem, Brazil, reported by de Carvalho (1960, 1961), revealed that P. discolttr consumes droplets of nectar secreted by the Bowers of Parkia gigautocarpa and P, peiuiida as well as the pollen and vegeta¬ tive flower parts of these species and of Ceiba peunuidni. Digestive tracts ex¬ amined by de Carvalho contained Bower parts, pollen, fruit, nectar, and insects. Goodwin and Greenhall (1961 ;238) staled: “This is a fruit-eating bat; in captivity . , , will not eat flesh. It has a long extendible tongue, with a deep groove on the upper surface which is used to scoop out fruit pulp.” Valdivieso and Tamsitl (1962), misinterpreting Goodwin and Greenhall (1961), included small verte¬ brates among the foods eaten by this species, Tamsitl and Valdivieso (1965) con¬ sidered P, discolor to be frugivorous although they had once reported it to be a consumer of both flow-ers and fruit (Tamsitl and Valdivieso, 1961). Villa-R. (1967) remarked that this species was a frugtvore in Mexico and included Ficus sp., Diospyros ehenaster^ and Achras sapola among those fruits consumed. The stomach of a Colombian specimen contained plant material and insects (Arata cf aL, 1967), The stomach contents of 128 Costa Rican and Panamanian P. discolor were reported on by Fleming et ai, (1972). They found 73 containing approximately one per cent fruit and 99 per cent insect remains, by volume. Only one kind of seed was noted in the plant material suggesting that a single fruit type had been consumed, i’he stomachs of the remaining 55 bats were empty. Heithaus a id. (1974) found P. dm-o/or carrying Bouhinia and Crescenda pollen on their fur and observed this species feeding at the flowers of Bauhinia paidcfia in Costa Rica. Later, Heithaus et al. (1975) reported the recovery of Ceiha pemandra. Crescentia spp., Ochroma lagopus, Pseudobonjbax seprinatuni, Manilkara zapoia, and iiyfnenaca courbaril pollen from the fur of Costa Rican P. discolor. They concluded that this species was primarily nectarivorous, at least during the dry season, and utilized a broad range of potential floral resources (79 per cent of the pollen loads were mixed). Fleming et al. (1972) w'ere cited as the authority for including insects in the diet; nevertheless, Heithaus ei al. (1975) found no remains of insects or fruit seeds and pulp in the feces. The diets of Costa Rican P. discolor reported on by Howell and Burch (1974) included vesi¬ culate plant material, fruit (Piper, Acnisms, and Musa), pollen (Hymenaea and Ceiba), and insects (Coleoptcra, Hymenoptera, Diptera, and Lepidoptera), McNab (1969) stated that P. discolor is a fruiteater; but in captivity, requires a small, but regular intake of meat. Power and Tamsitt (1973) remarked that this species is known to feed on fruit, insects, pollen, and nectar. HtOl.OGV OF THE PHYLLOSTOMATIDAE 307 Phyllosfomtjs hast;)tiis A variety of insects, small vertebrates, and plant material including fruit, pol¬ len, nectar, and flower parts. Authors of early accounts on South American bats often confused this species with Vampyrurn spectrum as well as attributing to it the “blood-sucking” habits of vampire bats. Bat species are difficult to recognize in some of these early narratives. Husson (1962:126) interpreted Waterton’s (1825) and Quelch’s (1892) observations on the habits of large Guianan bats they identified as Vampyrwn as being correctly ascribed. Waterton (1825:175), while discussing Vampyrurn^ stated: “He does not always live on blood. When the moon shone bright, and the fruit of the Banana-tree w'as ripe, I could see him approach and eat it. He would also bring into the ioft, from the forest, a green round fruit, something like the wild Guava, and about the size of a nutmeg. There was some¬ thing also, in the blossom of the Sawarri nut-tree, w-hich w-as grateful to him; for on coming up Waratilla Creek, in a moonlight night, I saw several Vampires fluttering round the top of a Sawarri tree, and every now' and then the blossoms, w'hich they had broken off, fell into the water, they certainly did not drop off naturally, for on examining several of them, they appeared quite fresh and bloom¬ ing. So I concluded the Vampires pulled them from the tree, either to get at the incipient fruit, or to catch the insects which often take up their abode in flow-ers.” Ouelch (1892:99), also relating observations on bats he believed to be Vampyrurn, reported: “It had been tantilising the evening before to W'ilness a continuous stream of these great winged creatures pouring out of one central hole high up in the trunk, and darting and w'heeling, fluttering and hovering, about the fruit trees around the house, and helping themselves, no doubt, to the ripest fruits on the small branches, as they listed; but it was infinitely more tantilising to know that the same stream would issue undiminished next evening, after our departure. “Though these bats are to a great extent insectivorous, yet from their size they must devour a large quantity of the mangoes, star-apples, sapodillas and other soft fruits w'here they occur, since their stomachs, when full, contain a con¬ siderable amount of pulpy matter. And indeed their great canine teeth, as in our bats generally, seem especially adapted for piercing and tearing open the skin, rind and fleshy parts of fruits, the power for the tear being derived from the force of their flight after they have seized the fruit with their teeth.” When these ac¬ counts by Waterton and Ouelch are critically examined, however, they obviously apply to Phyllcfsiomus hastatus, and perhaps to Ariibeus liiuratus as well, but not to Vampyrurn spectrum. Bates (1875:338) observed bats, which he called vampires, at Ega on the upper Amazon in Brazil. He discussed their habits and referred to their large numbers, frugivorous diet, and blackish and reddish color phases (he considered each color pattern to represent a distinct species)—characters identifying them as P. hastatus. Bates opened the stomachs of several and found a few remains of in¬ sects intermingled with masses of fruit pulp and seeds. Alston (1879-1882:39) and Goldman (1920:189), perhaps misled by Bates’ (1875:337) reference to their large size, assumed that these bats represented the species Vampyrurn spectrum. SPECIAL PUBt. I CATIONS MUSEUM TEXAS TECH UN I VERS f TV Dobson (1878) noted that the stomach of a P. hastadis was filled with the remains of insects. However, he also assumed that they occasionally fed on bats and other small mammals, Ruschi (1953/?) claimed that some of the feces of P. hcisfams were like those of vampires and, therefore, presumed them to feed on blood. Nonetheless, Ruschi (I953f/) gave the diet of this species in Brazil as in¬ sects, small birds and mammals (including bats), and the fruits of Musa pum- {iisiacay Carica papaya, Pskiiitu} pifajava, Eriohorrya Japtntica, Cecropiu sp., Solanu/n panicttlatam, Tcrntinalia cauippa, Livisknw chinensis, MangiUra iiuHca, Achras sapoia, Luaumi caintito, Eugenia unijlora, Myrcia Jaboticaba, Viiis vinlfera. Pa sstflora quadrangularis, Annona murkata. Pilocarpus pinnaiifolius, Artocarpus iniegrifolia, Ruhachia glomerata, Dhspyros kaki, '‘etc.” In Belem, Brazil, de Carvalho (1960, 1961) found P. hastaius feeding on the intlorcscences of Parkia gigatitocarpa, P. pcndula, and Cciba pentamlra. He reported finding agglutinated masses of pollen, anthers, parts of the corolla, and a yellowish clear liquid, possibly nectar, in the stomach. De Carvalho described the diet of this species as insects, fruit, birds, other bats, blood, and flow'er parts. The inclusion of blood in the diet may have been prompted by Ruschi's (1953^?) comments on the desmodontinclike feces found in roosts. Goodwin (1946) stated that the diet of Costa Rican P. hasiatus included vari¬ ous kinds of fruit, birds, small bats, mice, and insects. This is essentially the same diet suggested by Williams ct al. (1966) and Duke (1967). Goixlwin and Green- halt (1961) noted the remains of fruit, fur, and feathers at the bases of roosts in Trinidad, and the inclusion of both fruit and tlesh in the stomach contents. They mentioned that P. hasratus eats the fleshy funiculus of the Sapucaia nut (LiTyth/s zabucajo), a habit reported on in greater detail by Greenhall (1965). Greenhall (1966) reported P. hastatus feeding on ripe V'alencia oranges in Trinidad. De la Torre (1961:37) remarked that several P. hastatus, captured as they attempted to enter a cave, were carrying large guava fruit. Bloedel (1955) reported on a group of about 30 P. hasiatus that he observed several limes at twilight following late afternoon rains as they fed on swarming termites at Juan Mina in the Panama Cana! Zone. A Costa Rican specimen examined by Starrett and de la Torre (1964) contained fruit pulp, insect remains, a few bird feathers, and a partially digested tick in its stomach, The latter w-as probably consumed with its verte¬ brate host or was gleaned during grooming. Arata e( al. (1967) listed six stomachs of P. hasiatus from Colombia as con¬ taining plant material and three with insect remains out of seven they examined. Fleming et al. (1972) gave the stomach contents of 19 of the 25 Costa Rican and Panamanian specimens they examined as 4 per cent plant material and 96 per cent insect remains, by volume; the remaining digestive tracts were empty. Tuttle (1970), reporting on Peruvian bats, mentioned that netted specimens were frequently dusted with pollen. I have noted this in P. hasiaitis netted in Costa Rica and eastern Peru, HowtII and Burch (1974) did not report pollen from Costa Rican P. hasratus, but they did find the remains of fruit {Cecropia and Piper) and insects (Coleoptcra, Hemiptera, Lepidoplera, tmd Diptera including Culicidae) in the feces and stomach contents. HIOLOGY OK THE PHVLLOSTOMATIDAE 309 Summary. —The food habits of Phyllostomus ehngaius and P. latifolius are not known. Their diets, however, likely include flower parts, fruits, insects, and small vertebrates such as anoles and geckos gleaned from vegetation. As noted for P. hastaius, Tuttle (1970) frequently found the heads of netted P. elougatiis covQVQd wdth yellow pollen. Bats of the genus Phyllostomus omnivorous. Both P. iiiscolorand P. hastaius feed on animal matter, but in the former this is probably restricted to insects, w'hcrcas P. hastatus preys on a variety of small vertebrates as well. Fruits, pollen, nectar, and insects caught in flow'ers probably are the major food items of P. dis¬ color. The inclusion of blood in the diet of P. hastaius is w'ithout basis. The only blood consumed by this species is that of its vertebrate prey. Genus Phyllooerma Peters Phylloderma stenops Plant material and insects. The only reference to the food habits of this species is that by Jeanne (1970) who captured a male in the act of eating the larvae and pupae from an active nest of a social wasp (Polybia sericea) near Santarem, Para, Brazil. The stomach of this bat contained the well-masticated remains of both larvae and pupae, but no evidence of adults. Genus Trachoi^s Gray Trachops cirrhosus Insects, small vertebrates, and possibly some fruit. Ruschi (1953t’) recorded a diet of fruits, insects, and small reptiles for T. cirrlmsus in Brazil. Burt and Stirton (1961) staled that the stomachs of several specimens collected in El Salvador contained hair and flesh, Goodwin and Green- hall (1961) reported finding flesh and small sharp bones in the stomach of a T. cirrhosiis from Trinidad and commented on finding the remains of a gecko { The- cadactylus rapidicaudus) in the stomach of a specimen from Panama. Duke (1967) noted that Edw'in Tyson had observed Trachops hovering up and down tree trunks in a manner suggestive of an insect gleaner. In Honduras, Valdez and La- Val (1971) found a freshly killed ancle (AuolLs lemurinus) in the same net pocket containing a T. clrrhosus. They suggested that Trachops feeds on a variety of lizards- Only two of the eight stomachs of Panamanian T clrrhosus reported on by Fleming et ai (1972) contained food. The contents of both consisted entirely of insect remains, Howell and Burch (1974) recovered a mixture of Lepidopiera and bat hair from each of four Costa Rican specimens. 1 found T. cirrhasits commonly entering the houses of Cashinahua Indians at Balta, Departamenlo de Loreto, Peru, to feed on cockroaches during the evening. The bats were considered a nuisance because the sound of their flight as they moved along the walls and roof, the chewing noise as they consumed their prey, and the rain of urine, feces, and insect parts falling upon the mosquito nets below, disturbed the Cashinahua in their sleep. 310 SPECrAL PUHLlCA'llONS MUSEUM TEXAS TECH UNIVERSITY Genus Cuko i X)]> [ erus Peiers Chrotopterus auritus Small vertebrates, insects, and fruit. Goodwin (1946) was among the first to comment that Chrotoptenis is probably carnivorous. Ruschi {1953/;) repr^rted finding many bird vertebrae, solanaceous seeds {Sokinuml), and blood in the feces as well as fragments of fruit, fruit seeds, and scattered vertebrae under a cave roost of C aunius in Brazil. He also claimed to have witnessed a Chwtoptems land and commence feeding on the back of a calf. This observation was alluded to by Ruschi and Bauer {1957:41). Ruschi (1953/) listed the diet of Chnnopients as small mammals, young birds, fruit, insects, and blood.The bat on the calf most likely was a Desmodus, and the only blood in the diet of C. aurltus probably is that of the small birds and mammals preyed on by this bat. At least some of the seeds in the feces mentioned by Ruschi may have been in the stomachs and crops of birds eaten by Chrotoptems. Hall and Dalquest (1963) commented that the while stains beneath the roosts of these bats in Veracruz, Mexico, resembled those left by the excreta of hawks and owls. They also presumed C auritns to he carnivorous, an opinion repealed by Villa-R. (1967) and McNab (1969). Villa-R. and Villa Cornejo (1969, 1971) reported finding the fragments of skeletons, skin, and hair below a roost of C aurhus in a mine in northern Argentina, Their suggestion that these fragments were the re¬ mains of Cienomys is unlikely because of the fossorial habits of these rodents. Tuttle (1967) reported finding the remains of a gecko {Thecadaciylus mpidi- citudus) in the stomach of a Venezuelan specimen. Olrog (1973) reported finding the remains of a mouse opossum {Marmosa) and a bird among the stomach con¬ tents of Argentinian Chrotoptenis, Because the bird was being eaten in a mist net, Olrog concluded that Chrotoptenis had been eating hts mist-netted bats and birds. Genus Vampvrum Rafinesque VainpjTum spectrum Birds, bats, rodents, and possibly some fruits and insects. Vampyrmn spectrum figured prominently in many of the early narratives on the South American fauna because of its awesome proportions and erroneously ascribed blotxlTeeding habits. Husson (1962:14, 122-126) discussed those ac¬ counts dealing w'ith Guianan bats. Many of the early travelers, however, confused V. speetrum with Phyliostomus hastatiis account of P. hastatus) and possibly W'ith Artibeits lit unit its, Dobson (1878:471) remarked on finding “some vegetable matter of rather firm consistence, apparently |a] portion of the rind of some large fruif’ in the stomach of a V. spectrum. His remark (p. 471) that this species has been “shown by the observations of modem lra%'eltcrs to be mainly frugivorous” may have been influenced by Bates {1875, see acoimt of Phyliostomus hastatus), Goodw'in (1946) reported the diet of Costa Rican Vampyrum to be small birds, rodents, smaller bats, some fruit, and probably insects. Wehekind (1956:20) presented in- BIOLOGY OF THE PHYLLOSTOMATIDAE 311 formation on the food habits of K spectrutn in Trinidad. He found fur, feathers, and bone in the stomachs of three he collected and the remains of “blue birds,” doves, and rodents at the base of a roost in a silk cotton tree {Cetba penianclra). Two Costa Rican K specfrum were reported on by Cascbcer c/ ai. (1963) who found the remains of a passerine bird in the digestive tract of one, Brosset (1966:54) and Goodwin and Greenhall (1961) noted that Vampynitn is largely, if not entirely, carnivorous, and the latter mentioned finding fur and feath¬ ers in the stomach of this bat and bat bones at the base of a roost. Greenhall ( 1968) also said these bats were carnivorous and mentioned that a variety of fruits offered to V. speefrum in captivity w'ere never eaten. Duke (1967), however, listed Ana- canlium sp, and Psidium sp. as examples of fruits eaten by this species in Panama in addition to bats, rodents, birds, and insects. Peterson and Kirmse (1969) re¬ ported on finding the remains of bats and an oryzomyine rodent in the stomach of a Panamanian yampynmi. They surmised that this bat had been eating other bats caught in mist nets without having become caught itself. How-ever, their statement (p. 140) "‘that these bats w^ere virtually eaten in siiu in the net negated any evidence for birds of prey being responsible” is not necessarily correct. In Chiapas, Mexico, 1 saw a barred forest-falcon {Micnisiur mficoHis) in the act of eating small bats in a mist net at daybreak. The hawk hit a bat in the net, rebounded, and ate parts of it w'hile hanging nearly upside down and free from the net. As 1 approached, the hawk released the bat (a Stnntim I ilium) and flew off. The net contained other bats including some for which heads and upper parts of the body were missing. Later in the day, the hawk returned to the net and w'as captured while hanging and feeding on a small bird. Subfamily Glossdphaginae Genus Glossopkaga E. Geoffroy St. -Hilaire Glossophaga soridna Insects, fruits, pollen, nectar, and flow'er parts. This species, w'hich figured prominently in many early accounts of tropical American bats, was assumed to feed on blood. For example, Quelch (1892:101) Slated: “It seems likely . . , that these bats supplement their ordinary insect diet, with the blood of the domestic animals.” He noted, however, that the tongue seem¬ ed to be modified to lick out the pulpy matter of fruits. Gaumer (1917) stated that C. soridna feeds on insects and small or soft fruits in Yucatan, Mexico, and mentioned the fruit of Corciia dociecandra. He remarked (p, 297) that they open holes in the fruit and lick the juice, and “aiinque son vampiros nunca chupan sangre.” One of the earliest reports on the flower-feeding habits of G. soridna was by Porsch (1931), who observed this species at the flowers of Crescemki and Parmemiera alata in Costa Rica. Vogel (1958) reported that they feed on flowers of Ki^€dia aethiopka and noted finding the pollen of Maregravia sp. on the heads of some specimens. Baker (1970) presented information on flowers visited by bats at the botanical gardens of the Tela Rail road Company, Lancetilla, Honduras, 312 ■SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY and at Finca Lornessa, Santa Ana, Costa Rica. Even though the bats were not identified, examination of his photographs (figs. 1-4, 6-11) show G. soricina vh\t- ing the flowers of Dttrio ziht'dunus in Honduras. Those bats visiting the flow'ers of Miicuna andreana in Costa Rica are almost certainly this species as w'ell, in¬ asmuch as this is the only glossophagine I have ever netted at Ftnca Lornessa despite extensive collecting, and the species proved to be common there. Heithaus ei cii, (1974) reported capturing G. soticina dusted with Bauiiinia and Cre\ceniia pollen and mentioned observing this species at the flow'ers of Bauhiniu pattleiia near Cahas, Costa Rica. Heithaus e/ aL (1975) noted that 59.6 per cent of the 146 6\ soricifia they examined from the same region in Costa Rica carried pollen on the fur. The six most common plants represented by the pollen were Ochroma kigopn,s^ P.seKdoho/ttbax sepiinatiini, Ci'iha petuafidra, flyfneuaea cotirharil, Manilkam zapoui, and Crescetuia sp. They also found that these bats had fed on the fruits of Piper luherculaiti/u, Muniingia calahuray Sidanum sp., and Ficus sp. as well as on other fruits, the remains of which they could not identify. When contrasting frugivory and nectarivory in this species, Heithaus et ai (1975) re¬ marked that G. sork imi was primarily nectarivorous in both the wet and dry sea¬ sons in Costa Rica. The insect-eating habits of G. sorkina were acknow'ledged, how'cver; they cited Fleming ei ai (1972) as the source for this information. Test (1934) reported O’, soridua feeding on ripening bananas in Honduras, and rarnsitt and Valdivieso (1965) considered this species to be frugivorous. Glossophaga sorkina was thought to be nectarivorous by Park and Hall (1951), Wille (1954), Tamsitt and Valdivieso (1961, 1963), and McNab (1969). Goodw in (1946) also thought it w'as a nectar feeder and mentioned flowers of the calabash tree and night-blooming cacti as food sources, Piccinini (1971) noted that Brazil¬ ian G. soridua eat pollen and nectar. Dalquesl (1953), reporting on mammals from San Luis Potosi, Mexico, suggested that G. soridna feeds principally on nectar but consumes fruit juices and pulp as well. Hall and Dalqnest (1963) stated that it W'as a nectar and fruit eater and mentioned catching a specimen in a banana- bated snap trap placed in a tree in Veracruz, Mexico. A nectar and fruit diet was cited for G. soridua by Novick (1963) and Hall and Kelson (1959), Duke (1967), relating Edw'in Tyson’s information on Panamanian G. soridmu stated that they feed on overripe bananas and guavas and drink from the flowers of Musaceae, Bignoniaceae, and Bombacaceae. Pollen, nectar, and soft fruits w'ere noted in the diet of this species in Mexico by Villa-R. (1967), w'ho also reported finding some specimens with their heads covered with Ipomoea arhorea pollen. W'ied-Neuw'ied (1826) reported finding insects in the stomach of a G. soridna from Brazil. Alston (1879-1882:43) also stated that G. soridna preys on insects but mentioned that this species “feeds largely on fruits, lapping up the Juices and soft pulp w'iih their extensile tongues.” Brosset (1965), on the basis of tongue structure, suggested that G. soridna may capture insects at flowers as well as drink the nectar. Felten (1956) repvirtcd this species feeding on flowers (especially Cresceuda) in El Salvador; how'ever, on the basts of observations on captive individuals, he said they prefer insects. Fruit jutce.s and small insects were found in the stomachs of Trinidadan G, soridna (Goodwin and Green hall, BIOLOGY OF THE BHYLLOSTOMATIDAE 313 1961). Ruschi (1953^>) listed insects, fruits, nectar, and pollen in the diet of Brazil¬ ian G. Stfrkina. He noted that this species consumes nectar from many kinds of flowers, including Crescenka cujefe and Vochysia sp., as well as eating the fruits of Musa panidisiaca, Carica papaya^ and Solanuni paniculatam. De Carvalho (1960) reported on the stomach contents of G. soricitia from Belem, Brazil. One stomach contained the remains of insects, a large quantity of reddish fragments, presumably flower parts, and yellowish and whitish masses, probably pollen. Another contained scales of lepidoplerous insects and a gelatinous mass of protein¬ aceous material, presumably pollen. According to de Carvalho (1961) G. soricina eats fruit, flowers, nectar, and insects. Based on observations in the Parque do Museu Cioeldi, Belem, he cited a variety of flowering plants of which the nectar, pollen, and sometimes the flower parts are eaten: Cresccntia cajere, Crescentia aniazouica, Ak’.xa grandiflonu Hynienaea coarharil. Bougainvillea speaahilis, Cmfaeva bemfuwjiy and Eiizahelha paraense. Also cited were the fruits of Cccw/^/tr bureanianay Cecropia sp., Piper sp., and Achras sapota as foods consumed during periods of annual fruiting maxima (December to January), Starrett and de la Torre (1964) reported on C. soricina from El Salvador, Honduras, Nicaragua, and Costa Rica. They wrote {p, 57): “Fruit ‘pulp’ and seeds of a number of different kinds of plants were present to some extent in the digestive tracts of every Glossophaga. No pollen w'as found in any individual. Eight., . had insect remains in their digestive tracts. In tw'o cases the insect parts made up the hulk of the contents of the tract. The insects had been finely chew-ed but lepidopteran scales w'ere readily recognizable, as w'ell as portions of the wings of Diptera and Hymenoptera,'' They also observed G, soricifia feeding on ripe bananas at Los Diamantes, Costa Rica. Fleming ef al. (1972) classified G. soricina as an omnivore after examining the stomachs of 217 specimens from Costa Rica and Panama. Of these, only 38 stomachs contained food—^34 per cent plant material (including 5 per cent pollen) and 66 per cent insect remains, by volume. Two col¬ lected in January and one in May contained insects exclusively. One taken in March only contained pollen, and two caught in September only held fruit. Tw'O captured in February and six from December contained insects, fruit, and pollen, whereas the stomachs of bats in June, July, August, and October, contained fruit and insects. At least seven seed types were found in the plant matter recovered from these stomachs. Howell and Burch (1974) reported finding the pollen and nectar of Cresemiay Inga, Hymenaea, Mucuna, Musa, Pitcairnia, and an unidentified Bombacaceae in Costa Rican G. soricina. They also found the remains of fruit [Acnistus, Muntifigia, Musa, and an unidentified Melastomaceae) and Lepidop' tera in the feces or stomach contents. Lepidopterous insects were the only food items recovered from 24 of the 62 G. .soricina they examined, Alvarez and Gonzalez Q. (1970) presented information on their analyses of 174 G. soricina from the Mexican states of Veracruz, Oaxaca, Guerrero, and Morelos. Of these, 107 (61.6 per cent) did not contain pollen in their digestive tracts. Nevertheless, G. soricina contained the greatest diversity of pollen grains (at least 34 species recognized—see Table 2) of any of the other glossophagines studied {Anoura geoffroyi, Choeronycieris mexicana, Leptonycteris sanborni, 314 SPECIAL PUBLICATIONS MUSEUM TEXAS IECH UNIVERSITY Pah I E 2. — Pia/u.', kii’/uifial hy fn/lfcti firains in the Mtwftivh.'y •>/ Glossophaga sorici na, Anoura geoffroyi, Choeronycieris mexicaaa, Leptonyctcris sanborni, L. Nivalis, (tnd Hylonycieris underwoodi fro/n Mi'xico (modified from kihSe 5 in Alvarez rind Gmiznlez Q„ y970;/6.5), Podtn G, s^3rU^^llt^ -l. fnnt/frityi C. tui’suiiuti sutthifrni L. isi\ A etic hi + -f + + Acanlhaceae + Af^me a a a tu b iU Alhizziti + AlfUis + + + Amaranthaceac + Anoda + Apocynaceae + Arhatus + Arisfolochui + Haidiinia + + + fierherts + Bofuhtix + + + + a. i Bitrsem + Caexalpiniti + CtfKuindrn + a + a Ceihd h m b a a Citharexylum + Conipositae + a a + a Conztifihi + Conlia + + C re seen! in + + Crolon + Ei'/dninactnx + Eiictiiyptux + a + a Fit as + + + + + Gen (hi mi Ceraniimi + Graminae A + + + a Hibiscus + ipomoea a a a a Labiaieae + Luntanii + + Leguminosae + + Liliaceae + Lonvhocarpns Malvaceae + + + + Mimosa Musa + + Myrtillocaf 1 us^ a, h ii, b f;, h a Oenothera + + Onagraceae + + Pinas a a a a Plafyopuntta + + + Querens + Rouptda + BIOLOGY OF THE PHYLLOSTOMATIDAE 31 ^ Table 2.- — Coftiimteti. Salix + Sul via + + + Solanaceac + Urlicaceae + Zett + + Zingiber + Symbols: n, occorrini; in over 20 per cent of ii|l siomachs conlaining pollen l h, exceed iiii^ 2S per cent of loiirl volume of pollen; c, 9^.8 per cenl of slomach conients; d, includes Lemairencereus. L nivaiis, and Hyhtjycteris underwoodi), Alvarez and Gonzalez Q. (1970) con¬ sidered G. soricitm to be an opportunistic omnivore that utilizes pollen as a food source whenever nectar, fruits, or insects are not readily available. Their ob¬ servations on this species extended from February to September, during which time the commonest pollen ingested varied seasonally from Cordia, Acacia, Conzatiia, and Albizzki in the spring to Ceiba, Ipomoea, MyrtUiocactus, and Agave in the summer, and Cofjzadia in September. The pollen grains in the diet were correlated with each habitat. For example, three samples of G. sonciaa collected in May from different localities on the Pacific versant of Michoacari demonstrated different pollen profiles. Specimens from the Tepalcatepec-Balsas Basin south of Nueva Italia principally contained the pollen of LA'maireoceren.i and Ecb/nocacfas. In the vicinity of Arteaga, the most abundant pollen found were Raupaia, Agave, and Ceiba; however, in the MeJehor Ocampo area on the coastal plain, the primary pollen grains consumed w'ere Ceiba and Cordia, Arata ef al. (1967) presented data on the stomach contents of 16 Colombian G. soricina. They found that 15 contained plant material, and 6 contained the remains of insects. On the basis of a stomach containing matted hair, claws, and flesh, they ascribed carnivorous habits to this species. I suggest, however, that this observation not be interpreted as reflecting a normal aspect of the diet. As the authors noted (p. 653), the bats were collected at night and kept alive for prwes- sing the next day. According to Arata (personal communication), the bats were individually held in small cloth bags. Nevertheless, bats kept under these conditions sometimes will chew on themselves and, if pregnant, often abort and eat parts of the fetus. The finding of claw^s and flesh in the stomach of a G. soricina probably represents cannibalism induced by the treatment the bat received between the time it was caught and its death. Unfortunately, the sex and reproductive state of the bat was not presented. The information presented by Arata ef al. (1967) may have prompted Phillips (1971) to include meat in the diet of G. soricina, Goodwin (1934:9), commenting on G. soricina from Guatemala, said, “The single specimen taken at Barrillas was caught in a mousetrap hanging over a pile of raw sugar. Whether the bats were after insects drawn by the sugar or were there for the sweets, I cannot say, but I lean toward the former idea. The natives insist that bats eat the sugar.” The owners of Finca Lornessa, Santa Ana, Costa Rica, often found dead bats hanging on the edge or floating in the large vats used to concentrate sap from sugarcane if these pots were left filled and uncovered during 316 SPEClAl- PUBLICATIONS MUSEUM l EXAS TECH UNIVERSITY the night. The Finca Lornessa sugar mill houses a large colony of G. soricina, and the owners believe the bats drink the concentrated sugar cane sap. Glossophaga cominissarisi Insects, fruit, pollen, and nectar. The remains of Lepidoptera, fruit and the pollen and nectar of Musa and Mucuna were recovered by Howell and Burch (1974) from Costa Rican C. cofntuissarisi. Glossophaga longirostris Insects, fmit, pollen, nectar, and possibly other flow'er parts. Wille (1954) and Valdivieso and Tamsitt (1962) considered G. longirosiris to be nectarivorous. Goodw in and Greenhall (1961) said it feeds on fruit pulp and fruit juices, occasional insects, and some nectar. Pirlot (1964) said G. rosiris is nectarivorous and frugivorous. He also cited correspondence from Good- w'in and Novick, w'ho suggested that insects found among stomach contents have come from the fruit these bats have eaten. Summary, —The diet of Glossophaga alltcola is not knowm; however, its food habits likely arc similar to those of G. soricina and G commissarisi. Villa-R. (1967) reported the diet of G. morenoi as nectar, pollen, and pulpy fruits. I am uncertain to which of the three species of Glossophaga occurring in Mexico he was referring. The diet of Glossophaga includes a variety of plants and insects. Many of the insects may be consumed in conjunction with the flower- feeding habits of these species; however, some insects likely are caught aw'ay from flowers- Genus Monophyllljs Leach MonophyJlus redmani Nectar and fruit. Osburn (1865) reported finding yellow' pulp in the intestines of tw'o Monophyl- lus reihnauf taken in Jamaica. Wille (1954) reported that this species w'as “nectar- eating” on the basis of tongue structure. Tamsitt and Valdivieso (1970) also considered M. redmaui to be nectarivorous. Sioumary .—Nothing has been reported on the diet of Motwphyllus plethodou. According to Walker et al. (1964), Mouophyllus spp. are known to feed on the juices and pulp of fruits and presumably include insects in their diet. Genus Leptonycteris Lydckker lA'ptonycteris nivalis Fruit, pollen, nectar, and insects. Park and Hall (1951), on the basis of tongue and stomach anatomy, considered L. nivalis to be nectarivorous. Dalquest (1953:28) found L. nivalis, captured in BlOl.OGY OF THE PHYLLOSTOMATIDAE 317 rooms of Hacienda Capulin, San Luis Potosi, Mexico, with their stomachs filled “with thick, brilliant red fruit juice . ., almost certainly the Juice of the fruit of the organ cactus.** Novick (1963) believed that this species feeds on dowers and fruits. Barbour and Davis (1969) remarked that L. nivalis feeds on nectar and pollen. Phillips et ai. (1969) mentioned nectar, pollen, and soft fruit in the diet of L. nivalis in their report on the macronyssid miles inhabiting the oral mucosa of these bats. The mites were found in L. nivalis but not in L. sanhorni, even when both species were found in the same cave. They implied that the presence of the mites in L, nivalis indicated a diet differing from that of L. Sanborni and suggested that abrasive diets of insects or plant fibers might pre¬ vent the mites from inhabiting the oral cavity of L. sanborni. Alvarez and Gonzalez O. (1970) reported on pollen found in the stomach contents of 13 specimens from the Mexican states of Michoacan and Hidalgo. Pollen grains representing 22 kinds of plants were found in 12 stomachs (Table 2); one stomach did not contain pollen. Lepionycferis nivalis consumed the pollen of Ipomoea, Ceiba, and Mynillocacms in about the same proportions as did L. sanborni. Alvarez and Gonzalez Q. remarked on not finding any significant differences in the diets of the two species. Leploiiycteris sanbomi Fruit, nectar, pollen, and insects. Wille (1954), on the basis of the throat musculature of specimens from Jalisco, Mexico, considered L. sanborni to be nectarivorous. Hoffmeister and Good- paster (1954) presumed L. sanborni to feed heavily on pollen and nectar after observing that nearly every specimen they collected in the vicinity of the Huachuca Mountains in southern Arizona had yellow pollen covering the head. An analysis of the stomach contents of six specimens revealed an average of 92 per cent pollen and 8 per cent insect remains. Hoffmeister and Goodpasier (1954) surmised that the pollen came from Jimsonweeds {Datura)^ w'hich have yellov^ pollen, are open at night, and are abundant in the area. Alcorn et al. (1961) and McGregor et ai. (1962) reported on experiments they conducted in southern Arizona with caged L. sanborni exposed to flowering saguaros {Carnegiea gigantea) and century plants (Agave schottii). They found 62 per cent of the saguaro flowers setting seed when pollinated by L, sanborni as opposed to 52 per cent for bees and 45 per cent for white-winged doves. Hayward and Cockrum (1971) presented information on analyses of digestive tracts col¬ lected from L. sanborni between 15 May and 2 September over a four-year period in southeastern Arizona. They found the tracts to contain 100 per cent saguari> pollen in mid-May. The pollen content shifted whth increasing percentages of Agave pollen beginning in late May to early July and thereafter to early Septem¬ ber, w'hen the pollen content w-as 100 per cent Agave. The stomach of one bat taken on 7 November near Carbo, Sonora, Mexico, contained a few grains of saguaro pollen. Hayward and Cockrum (1971) expressed the opinion that L. sanborni is a nectarivorous species in the United States and accidentally ingests pollen while feeding at flowers. They believed pollen to comprise the major pro¬ teinaceous portion of the diet and mentioned that, when nectar is not available. SPEC[AL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY 3JK L. sanborni eats soft juicy fruits. Barbour and Davis (1969) gave a diet of nectar, pollen, insects, and fruit, Alvarez and Gonzalez 0. (1970) reported on the pollen found in the stomachs of L. sanbomi over a six-month period in the Mexican states of Hidalgo and Guerrero. Fecal samples were collected in a cave in Xoxafi, Hidalgo, from Feb¬ ruary to September. Of the 279 stomachs examined, 249 contained identifi¬ able pollen grains representing 28 kinds of plants (Table 2). The results from the fecal analyses duplicated the information obtained from the stomach contents, Lcptonycteris sanbomi first arrived at the cave in Xoxafi in February and, since their stomachs contained pollen grains of BoinbcLX and Ipomoea, were presumed to have come from subtropical habitats. The pollens found in the stomachs of these bats reflected the plants that were flowering at the time as well as the changes in the flowering times of the flora from one season to another. For example, in the vicinity of Juxtlahuaca, a subtropical locality in Guerrero, bats contained large amounts of the pollen of Bomhax, Ipomoea, Ceiba, and A^ave on 3 February, as well as very small quantities of the pollen of Myriillocaaus, On 20 July, however, L. sanbomi stomachs contained nearly 90 percent MyrtiUocacfns pollen, no Botnbax pollen, and greatly reduced amounts of Ceiba, fpomoea, and Agave pollens. Comparisons of pollens found in the digestive tracts of L. san- borni taken in late July from Xoxafi and Juxtlahuaca demonstrated differences in the foods available in these two contrasting habitats. Agave pollen predomi¬ nated (98.7 per cent) in the stomachs from Xoxafi, w'hereas the pollen of Lemaireocereus was commonest (87.7 per cent) in bats from Juxtlahuaca. No significant differences were noted in the pollens consumed by L. sanbomi and L. nivalis. Howell (1974) reported finding fragments of thrips {CarpophUus) and a bee ( Haliaus) in some stomachs of L sanbomi from southern Arizona. However, she suggested that, because these insects are associated wdth batflowers, they were consumed incidental to nectar feeding and are not actively pursued. Thirty stomachs contained an average of 4 grams of material each, of which about 25 per cent was pollen; the remainder, nectar, Howell’s thesis is that L. sanbomi is nectarivorous, prefers Agave and Carnegiea flowers as food sources while in Arizona, and that pollen supplies all of the dietary proteins. She sup¬ ports her contention regarding the dietary role of pollen by pointing out the higher nitrogen content of pollen from chiroptcrophilous plants (when compared against anemophilous and entomophilous pollen) and demonstrating the array of “essentiar’ amino acids present in Agave and Carnegiea pollen, two of which (proline and tyrosine) are recommended as being of special importance to bats. Summary. —-The food habits of U^ptonycteris curasoae are unknown; however the diet likely is similar to those of L. nivalis and L. sanbomi. There have been several reports concerning the food habits of Lepfonyaeris; however, it is nearly impossible at this time to determine to which North Ameri¬ can species this information applies. Duges (1906) reported on finding an Ichnoglossa (= Lepkmycteris) in Guanajuato, Mexico, the fur of which was covered and stomach filled with the pollen of Malvavisvus acerifolius. BiOl.OGY OF THE PHYIT.OSTOMATIDAE 319 Palmer (1954) stated that Li’ptonycferis probably feeds on flowers. Sanborn (1954) maintained that these bats feed on insects from night-blooming flowers because cactus pollen has been found in some of the stomachs examined. Hoff- meister (1957) gave the diet as nectar, pollen, and insects. According to Walker et aL (1964), LQ'piouyaens is known to visit the flowers of Malvaviscus ami perhaps jimsonweed {Datura) and to eat the fruits of cacti. Genus Lonchophylla Thomas LonchophyNa rnordax Insects, fruits, nectar, and pollen. Ruscht (1953/) recorded the diet as insects, succulent fruit, nectar, and pol¬ len. Lonchophylla concava Pollen, nectar, and insects. How'ell and Burch (1974) reported the following food materials recovered from six Costa Rican specimens of L, coucava: one with nectar and Mucuna pollen, two with nectar and Musa pollen, and three with the remains of Lepidop- tera. Lonchophylla robusfa Pollen, nectar, fniit, and insects. Wille (1954) considered L. robusta to be a nectar-eating bat. Fleming et ai. (1972) examined the stomachs of 17 specimens from Costa Rica and Panama, Ten per cent plant materia! and 90 per cent insect remains were found in the only stomach containing food items. Howell and Burch (1974) did not find any plant material in their three L, rohusta from Costa Rica; however, they did find the remains of Lepidoptera, Coleoptera, and Streblidae, 5//m/nary.—Nothing has been published on the food habits of Uynchophylia hesperia and L. thomasi. Walker et at. (1964) remarked that Lonchophylla feeds on flowers and the diet may consist of nectar, pollen, insects, and fruit. Duke’s (1967) mention of nectar and possibly overripe fruit, pollen, and insects in the diet of Panamanian species of Lonchophylla may apply to all species in the genus. Goodwin (1946:312) wrote that Umchophylia. , . is to some extent a nectar feeder, and uses its long tongue to lap up the honeyed liquid from the large night- blooming flowers.” I have seen L. thomasi feeding at banana flowers in eastern Peni and many of these bats had their heads and shoulders dusted with pollen. Genus Lionycteris Thomas Lionycteris spurrelti The diet is unknown; however, the food habits of Lionycieris likely are similar to those of Umchophylla. 320 SPECIAL PU13I.1CATIONS MUSEUM TEXAS TECH UNIVERSITY Genus Anoljra Gray Afioura neoffroyi Fruit, pollen, nectar, and insects. Perhaps the earliest account providing information on the food habits of A. Si'ojfroyi is that of Tschudi (1844:73), who recorded the remains of Diptera in the stomach of a specimen from Peru. Ortiz de la Puente (1951) also found in¬ sects in the stomachs of Peruvian A. fieojfroyi. According to him (p. 12), an ex¬ amination of stomach contents revealed the remains of two species of small coleopterans, one of w'hich is a member of the family Nitidulidae. Knuth (1906:73) related J, H, Hart's observations {in lirf.) on visits by G!os- sonycieris Geoffroyi A. geoffmyi) to the flowers of Eperua falcani in Trinidad. This information was mentioned by Baker and Harris (1957:449) and given without citation by Walker et al. (1964). According to Goodwin (1946:312), ''Anoura is in part a nectar feeder, and its long longue is adapted for reaching into the corolla of various night-blooming tropical flowers. It is known also to visit blossoms w here there is no secretion of nectar, and it may be supposed that there they are attracted by the insects drawn in by the perfume of the flow'ers.” Ruschi (1953^) mentioned visits to Bowers of Vodtysia sp. by A. geojfroyi in Brazil and listed insects, fruit, nectar, and pollen in the diet of this bat. W'ille (1954) considered this species as ncciarivorous. Vogel (1958) noted that A, geoffmyi visits the flowers of Symbokinthus taiifolius and Purpiirelki groHsa. Goodwin and Greenhall (1961) reported that Anoimi feeds on nectar and the soft pulp of ripe fruit. Villa-R. (1967) gave nectar and pollen as the foods of A. geoffroyi in Mexico. Duke (1967) stated that this species is a nectar feeder in Panama that also eats overripe fruit. Goodwin (1934) remarked on catching an A. geoffroyi in Guatemala in a mousetrap hanging over a pile of raw' sugar. The high incidence of insect remains and the numerous stomachs without pollen prompted Alvarez and Gonzalez Q. (!97()) to suggest that A, geoffroyi is a facultative pollen eater. Alvarez and Gonzalez Q. reported on 69 A. geoffroyi from the Mexican states of Michoacan, Mexico, Guerrero, and Oaxaca. Of these bats, 34 contained identifiable pollen grains, representing 20 kinds of plants, in their stomachs (Table 2), The kinds of pollen present were similar to those found in other species and reflected the flora in the different habitats w'here the bats were found. The major differences noted betw'een A, geoffroyi and the other species Alvarez and Gonzalez Q. (1970) studied were the increased representa¬ tion of entomophilus plant pollen (for example, Compositae) and the high fre¬ quency of insects in the stomach contents. According to them, A, geoffroyi be¬ haves tike an insectivorous species with a partiality for pollen. This observation is supported by Howell and Burch (1974) w'ho found only Lepidoptera in the specimen they examined from Costa Rica. Anoura caudifer Fruit, nectar, pollen, and insects. Wied-Neuw'ied (1826:217) mentioned finding the remains of insects in the stomach of an Anoura ecaudata ( = A. caudifer) from Brazil. Ruschi (1953y;) tllOLOGY OF THE PHVLLOSTOMATOAE 321 claimed that Brazilian A. caiHiifer eat insects, soft Juicy fniits, nectar, and pollen. Anoura cult rata Insects, pollen, and nectar. I'hc six A. cullrata examined by Howell and Burch (1974) contained only the remains of Lepidoptera. However, I collected several A. cuhra/a in Costa Rica, the heads and shoulders of which were dusted with pollen. Anuura wcrckJeae Pollen and probably nectar, fruit, and insects. Starrett (1969) reported A, werckleae visiting the Bowsers of Werckica lufea, as he determined by finding iVercklea pollen on the fur of the head and shoulders Sttnimary .—-Very little is known of the food habits of Atu^ura werckleac and A. cult rata; the diet of A. brevi rostrum is unknown. However, these species probably have diets similar to that of A, geoffroyk which is a highly insectivorous glossophagine. Genus ScLERONYCTEKis Thomas Scleronycteris ega Probably fruit, pollen, nectar, and insects. Nothing has been reported on the diet of this species. Genus Lichonycteris Thomas Lichonycteris obscura Pollen, nectar, and insects. Goodwin (1946:315) wrote: '^Lichonycteris is probably a nectar feeder as is indicated by its weak teeth and absence of tower incisors, to give the long tongue free play,” Tamsitt and Valdivieso (1961) included L. ohscuru among species that consume flow'ers and fruits. Carter et al. (1966) reported on two specimens netted near a plant bearing night-blooming flow'ers in Guatemala. They noted pol¬ len on the rump and uropatagium of these bats. Summary .—Nothing is knowm of the diet of Lichonycteris degener and little is known of the food habits of L ohscura other than the fact that these bats visii flow'ers. Genus Hylonycteris Thomas Hylonycteris underwoodi Insects, pollen, and nectar, Goodwin (1946) believed Hylonycteris to be a flower visitor and Hall and Kelson (1959) stated that it is a nectar feeder. Hall and Daiquest (1963:228) re- 322 SP£C]A[- PUBLJCATIONS MUSEUM TEXAS TECH UNlVERSiTY porting on this species in Veracruz, Mexico, wrote: “Beneath their resting place was a pile of guano about three inches high by six in diameter. There were several pits of jobo plums \Sp(Huiias lutea] on the pile, showing that some of this fruit is taken to the cave to be eaten/’ Carter er aL (1966) reported a speci¬ men with pollen grains on the rump and iiropatagiiim netted near night-blooming flow^ers in Guatemala. Viila-R, (1967) stated that Hykmycieris feeds on nectar and pollen. He cited a specimen from Tabasco, Mexico, with pollen that he suspected to be cacao ( Theobroma cucdif) on the vibrissae and hairs around the mouth. Walker e! al. (1964) reported the diet as probably nectar, fruit, and some insects, Alvarez and Gonzalez O- (1970) reported on the pollen found in the Stomachs of two specimens from Chiapas, Mexico. The stomach contents w'ere composed almost entirely of the pollen of Lofidiocarpus (99.8 per cent). Pollen of and Finns were also present but in minute amounts (Table 2). Apparent¬ ly, Howell and Burch (1974) were the first to demonstrate insectivory in H. iimkr- woodi. They recovered the remains of Lepidoptera from a specimen in Costa Rica. Genus Platalina Thomas Platalina getiovensium Probably p<.illen, nectar, and insects. The food habits are unknown. Genus CHOiiRONtscus I’homas Choeroniscus gudmani Probably pollen, nectar, and insects. Gotxlwin (1946:313), commenting on Costa Rican C, godnunib wrote: “Tip of tongue has numerous thread-like papillae forming a bnish, especially adapted for reaching the nectar at the base of the corolla in large blossoms.” Villa-R. (1967), while discussing C. ^pidmani in Mexico, cited the stomach contents given by Goodwin and Greenhall (1961) for C intermedins from Trinidad, Choeronisciis intcrmcdius Pollen, nectar, and insects, Goodw'in and Greenhall (1961:248) stated: “Microscopical examination of the stomach contents of one specimen [from Trinidad], however, revealed some minute particles that are possibly honey or fruit juice, many fragments of a coleop¬ terous insect, and numerous brown and white, hair-like strands, probably cither from insects or from fruit. This specimen, at least, had fed to a large extent on in¬ sects. ” Nothing is know'll of the fotxl habits of Choeroniscus minor, C incety and C periosus. Their diets, how^ever, probably include pollen, nectar, insects, and small juicy fruits. BIOLOGY OK THE PHYLLOSTOMATtDAE 323 Genus Choeronycteris Tschudi Choeronycteris inexicaiiii Fruits, pollen, nectar, and probably insects. Dalquest (1953) expressed the opinion that in San Luis Potosi, Mexico, C mexk ana feeds principally on the nectar of desert flowers, probably of cacti. Park and Hall (1951), Wille (1954), and Hall and Kelson (1959) considered C nwxi- cam to be nectarivorous. Hoffmeister and Goodpaster (1954) noted several specimens from the Huachuca Mountains in southern Arizona that had yellow^ pollen on the fur around the face. Huey (1954) mentioned C me.Kkatm from San Diego, California, with yellow' matter in their stomachs and pollen on their faces. Sanborn (1954) suggested that this species may have the same feeding habits as Li'ptonyaefis nivalis. Villa-R. (1967:263) reported capturing four C mexicatui in Bahia de San Carlos, Sonora, Mexico, the mouths of w'hich contained remains of the fruit of pitahayas i Lanaireocereus) or garambullas {Myrfillocacfus). He also mentioned observing this species flying w'ilh Li^ptonycteris around fruiting cacti. In Guerrero, Villa-R. (1967) caught five specimens that had their heads and shoulders covered with Ipomoea arhorea pollen. Walker et al. {1 964) suggested a diet of nectar, pollen, fruit juices, and insects for this species. Barbour and Davis (1969) stated that C mexicana probably feeds on nectar and mentioned individ¬ uals for which the heads and faces were covered with pollen when captured. Alvarez and Gonzalez Q. (1970) reported on the pollen found in the stomachs of 16 C mexicana from the Mexican states of Hidalgo, Guerrero, and Morelos. All stomachs contained pollen grains, and 17 pollen types w'ere identified (Table 2). Noting that the major percentages of pollen were from Lemaireocereas, Ceiha, Ipomoea, Agave, and Myrtillocactus (plants that are especially attractive to pol- lenophagus bats), Alvarez and Gonzalez O. (1970:156) expressed the opinion that C, mexicana is an obligate pollen feeder. Genus Musonycteris Schaldach and McLaughlin Musoiiycteris hairisoni Probably pollen, nectar, and insects. Schaldach and McLaughlin (1960) remarked on M. harrisoni feeding on the nectar of banana flowers in Colima, Mexico. I noted pollen on the heads and faces of several M. harrisoni caught in a small banana grove in Colima (some of these specimens were included in the report by Schaldach and McLaughlin, I960). Villa-R. (1967) stated that this species feeds on pollen, nectar, and insects found in banana flowers. Subfamily Carolliinae Carollia castanea Genus Carollia Gray A variety of fruits and insects. ?24 Sf^EClAL 1>UHI ICATJONS MUSEUM TEXAS TECH UNIVERSITY Because C castanea has been confused with C\ subnifu and C. brevicaiuia by many investigatiors, it is difficult to ascribe correctly the information on food habits in many of the accounts on Carollia to this species even w'hen the name C castanea was cited. A notable exception is the report by Fleming ei al, (1972) on bats from Costa Rica and Panama. Sixty-nine of the 102 stomachs of C. ais- icmea they examined were empty. The other 33 stomachs contained approxi¬ mately 92 per cent plant material (fruit) and 8 per cent insect remains. Included among the plant matter were 10 kinds of seeds. I’he stomachs of 28 C castanea from Costa Rica and Panama collected during all months of the year, except July and September, contained plant material exclusively. The stomach of a bat col¬ lected in July contained only insect-matter. No information was given on the stomach contents for September=caught C casfanetL The eight May-caughl C. castanea from Costa Rica reported on by How'ell and Burch (1974) had been feeding on Piper, They were able to identify Piper aurifum in three of the bats. Car(»Uia subrufa Probably fruit, flowers, and insects. Many investigators have confused C. perspicillata, C. hrevicaiuia, and C. castanea with this species. Information, however, in the accounts by Sanborn (1936) and Starred and de la l orre (1964) probably apply to this species. Star- rett and de la Torre (1964:58-59) stated: “Several types and colors of fruit pulp w'ere taken from the digestive tracts of both specimens [from El Salvador], along with bat hairs. A small stalked inflorescence was also found in the small intestine of one, and a segment of an insect leg in the tract of the other." Sanborn (1936) related catching a C sahrufa in a steel trap set on a bunch of bananas in Escobas, Guatemala. Carotlia brcvicauda A variety of fruits and probably insects as well. According to Dalquest (1953:30), in San Luis Potos'i, Mexico, "'Carollia per- spicilUita I — C. brevicauda —‘See Pine, 1972:35, 38] feeds entirely on fruit. It does some damage to stored bananas, but wild figs and other wild fruits con¬ stitute its principal food." Hall and Dalquest (1963:231) repK>rted that the stomachs of C perspicilkita C, brevkaiida —see Pine, 1972:35, 38) taken in Veracruz, Mexico, “held a semi-liquid mass of yellow' pulp, probably of the w'ild sweet- lemon or wild orange." They also referred to four occasions when C brevicauda were caught in banana-baited snap traps. Three w-ere taken in traps on the ground and the fourth was caught in a trap suspended above a steni of ripe bananas hang¬ ing in a tree. Villa-R, (1967-269) reported that he observed this species in San Luis Potos'i, Mexico, eating cakes of raw' sugar that were hanging high up in the eaves of a house. Carollia perspicillata A variety of fruits, flowers, and insects. BIOLOGY OP [ HE PHYLLOSTOMATIDAE 325 Many reports on C perspkiHaia simply call it a frugivore (Park and Hall, 1951; Goodwin, 1946; Valdivieso and Tamsitt, 1962; Tamsitt and Valdivieso, 1965; McNab, 1969). Davis (1945) recorded instances in Brazil where C perspiciilata were caught in banana-baited snap traps set on the ground. Puttie (1970) also reported catching several of this species in rat traps (baited with bananas) set on the ground and mentioned that in Peru these bats entered Indian houses to eat bananas. Ruschi (1953/) gave the diet of Brazilian C. perspicidaw as fruit and insects. Goodwin and Green hall (1961) and Green halt (1956, 1957) cited the fruits of 23 species of plants consumed by this species in Trinidad (see Table 3). Goodw-in and Greenhall (1961:250) also stated: “If the fruit is large, the bat eats it while hanging in the tree; if small, the fruit is plucked and carried by the bat to a temporary roost, called ‘digesting place’ to be eaten.. . . Some fruit is carried to the regular daytime roost." Starreit and de la Torre (1964:58) described the contents of the digestive tracts of four C. perspicilUifa from Honduras and Costa Rica as “several types of fruit pulp, seeds and vegetable fibers." Arata ei al. (1967) reported on the stomach contents of 74 Colombian C perspicilUtHL They found 71 w-ith plant material, 16 containing insects, and 6 with matted hair, claws, and flesh. The remains of bats (claw's and flesh) in the stomachs could be the result of holding the bats together for a pcrkKl of time after they were caught and may not rep¬ resent normal food items for this species (see account of Glossophaga soridna for the discussion of a similar situation). Howell and Burch (1974) reported re¬ covering the following food items from Costa Rican C, perspk iikita: the fruits ot Piper, Cecropia, Heisteria, Licania, Acnisms, Soianum, Mangifera, an unidenti¬ fied large-seeded Solanaceae, and the remains of unidentified insects as w^el! as those of Coleopiera. Summary ,—Many of the reports on Carol I ia confused the species C. casntuea, C. subrufa, C, perspicillaia, and C. brevicamia. In some instances, species can be recognized because only one is knowm to occur in the region discussed; for example, C. brevicamia in San Luis Potosi, Mexico (Pine, 1972:38), Villa-R. (1967:269) slated that Caroilia eat Musa sp., Dhspyros ebenasU'r, Achras sapcKa, Casimiroa edulis. Ficus spp,, and pitahayas {Lemaireocereu.s) ir Mexico. Duke (1967) remarked that Caroilia eat almost any fruit in Panama, and he cited, as examples, Cecropia^ Ficus, Mangifera, Musa, Piper, and Pskiium, Fleming ei ai (1972) acknow'ledged that their sample of Carollic from Costa Rica and Panama included C. brevicaiula and C, perspicilUua, The> examined 760 stomachs and found 272 with food items that consisted of about 87 per cent plant matter and 13 per cent insect remains, by volume. The plant material included a wide variety of fruits as determined from the 22 kinds of seeds present in the stomachs analyzed. Stomachs either contained all plant matter or a combination of plant and insect remains, Heithaus a al. (1975) also pointed out that their C ""perspicillata'^ from Costa Rica probably included more than one species. Nevertheless, they concluded that these bats were primarily frugivorous but utilized nectar during the dr}' SPEClAl, l>UHLICATIONS MUSEU\f TEXAS TECH UNIVERSITY Taivi.e 3 ,—Pliifitx utUizi'il in tiw f>/ Carollia perspicillata. Ariibeus jamaicensis, and Artiheus liiuraiiis. Plani ipecit-'s I'ari cjiiL-n* References" C pci't’ffk'nfutii A. Jitnntifi‘Hai.% A. iiturafm Achrtis xapafti SP 2. 3. 4 2, 3, 4, 5, 6 1. 2, 3, 4 Avnixiiis sp. fruil 7 Acrtnontia aadetita P 4 4 Aitiiini»fus umhi'lhdifi'i'n P 2,3,4 2, 3.4 A>idif-(t i tier ft} IX SP 2, 3, 4 2, 3, 4 Aiituitia fiuit'imut P 2, 3, 4 2, 3, 4 An no fit! pixotiix fruit 1 Annotui xqnatnoxa P 2, 3, 4 2, 3. 4 A/uhitriiifft sp. fruit y A rfociirpits itinprijHYLLOSTOMATiDAE ni TAflLt 3.-— Cf>nti!uu'd, Ficus t)hiitsift}iUi Ficus raduht Ficus reiiiiiostt Fk'its rcsitsu Fisuc ^ipp. FUu'Ouriia indica Gcnipu sp. HeistcrUt sp. HyUscen its ivnutini Hymetuica courburil iyiarfea cxorrhizu Jumhosii vtdgufis Lcuutireoceri’us sp. Lk'itnhi sp. Livisumu chine us is Luvittiu! ciiififitti MiidhtiCit iiuifotia Mill p is,' hkt ^ fa hr a Mciiittuea iimerkafiu Mari}'if cm indica Ma tiilkaro bide a (at a Manifkara zap of a Mclicffiva bijuffu XJituusops eie/ipi Mismiuffia calahura .\'ftisa pa rad is ia ca ■Vfiisa spp. Myrcia jaboficaha Ochtwiiu iaut^pas Fassijlora iiuadranffaiaris Persca amcricana Fifneufa racc/nosa Piper afualiif'o Piper aurittiai Piper bispidu/a Piper saacfuftt Piper taberculafuui Piper spp. Poidsenia ar/nafa Pouteria mullijlora Pscudohouihax sepiinafu/u Pseudo!media oxyphyllaria Pskiium puajavu Ps id sum m ed iferrtitieum Putraajivora roxburph ii Quaruribea funehris Rheedia edufts Roystsmea oleracea Suip only’, Sp, skin and pulp; F, flowers (includes pollen andneciar; 1., leaves; fruit, no informal ion on ptirl offrtiil consumed, ♦♦References: I) Riischt, I'J?.!/; 2 ) Greenhall, Grcenhall. !^57; 4) Gotxlwin and Greenhall. IS61; 5) Carvalho. 1961; 6) Villa-k., 1967; 7) Howell and Uurch. 1974; S) lleilhaiis I'l tti. 1975; 9) Vii/gueZ' Yanesrr (d., 1975. season. The six most common ptillens found on these bats were Of7/ro/?j<:i lagopus, Hyffu^tuiea courharil, Pseiulohonthax sc’pti/mfum, Crescentia sp., Manilkara zapota, and Ceiba pemundra. Identifiable fruit remains recovered in the feces of these bats included Ficus sp., Mimtingia caiabum, Sulammi sp., and Piper !uhercukinim. They reported 38,2 per cent of i 86 bats with pollen, 32.4 per cent of which carried two or more species of pollen; 44,9 per cent of 316 bats w'ith seeds in their feces; and 13,0 per cent of 272 bats had consumed insects (per¬ centage by volume; data from Fleming et id., 1972). Klite (1965) reptirted on the transit time through the digestive tract of dyed fruits in three Neotropical bat species from Panama including three individuals he identified as C. perspiediauu When India ink was used as a marker, two of the three CaroHhi passed stools containing the ink after a time lapse of 30 minutes. These results indicate that some frugivorous species arc able to extract the nutri¬ tive components of their food in a very short time and may consume several times more fruit in a single night than the holding capacity of the stomach would suggest. Genus Rhinophyj.la Peters Rhinophylla piimilto Presumedly fruit. McNab (1969) considered R. pumilio to be frugivorous. Tuttle (1970) re¬ corded capturing a male in a banana-baited trap set on the ground beneath ferns in dense mature forest in Peru. Summary ,— The fot^d habits of Rhinophyiki alethiua and R. fischerae are not known. Bats of this genus are probably all frugivores, although they may con¬ sume insects as well. BIOLOGY OF THL PHYLLOSTOMATEOAE m Sdirtlira Ulitim Subfamily Stenodermjnae Genus Sturnira Gray A variety of fruits and possibly pollen and nectar as well. Most accounts on S. lilium simply state that the species is frugivorous (Go(k1- win, 1946; Tamsitt and Valdivieso, 1961; Villa-R,, 1967; McNab, 1969). Sev¬ eral investigators have mentioned finding the remains of fruit in the digestive tracts of these bats (Dalquest, 1953; Goodw in and Greenhail, 1961; Starrett anti de la Torre, 1964; Arata et uL, 1967; Fleming e( al., 1972). Cockrum and Brad¬ shaw' (1963) reported on a S, liliufu shot from among several bats observed feed¬ ing on w'ild figs (Ficus) growing along the Rio Cuchajaqui in southern Sonora, Mexico. Villa-R. and Villa Cornejo (1969, 1971) remarked that S. liliuni take the fruit of the date palm and are attracted to ripe bananas in northern Argentina. Sanborn (1936) referred to a specimen caught in a steel trap placed on a bunch of bananas in Escobas, Guatemala. Gaunter (1917) reported that S. Ulium in Yucatan, Mexico, eat insects, although he said their principal food was fruit. Ruschi (1953A) also gave the diet of 5. liliiun in Brazil as fruit and insects. I have collected S. lilium at Balta on the Ri'o Curanja, Departamento de Loreto, in eastern Peru, the feces of which contained the seeds of Cecropia sp. and Piper sp. One entered a mist net while carrying a catkin of Cecropia sp. in its mouth. Heithaus ei al. (1974) reported recovering Bauhinia pauleiia pollen from a specimen near Cahas, Costa Rica, and mentioned (p. 418) that "'S. Ulium visited other flowers in the study region.” The latter observation was substantiated by Heithaus ei al. (1975) who reported finding pollen on 4L8 per cent of 110 Costa Rican S, Ulium of w'hich 47.8 per cent carried two or more species of pollen. The six most common pollens they recovered were Crescemla sp,, Pseudohomhajc septinalum, Marillkara zupom, Hymenaea courharil, Ochrcm.a lagopus, and Ceiha pentandra. Most of the fruit remains found in the feces weie unknow'n; however, they were able to identify the seeds of Piper luberculaiurp MiuUingia calahura, and Solauum sp. How'dl and Burch (1974) reported the fol¬ lowing fotxl items recovered from S. Ulium in Costa Rica: insects (Lepidoptera), pollen (Celha)^ and fruit {Piper, Licania^ Mumingia, Acnistus^ Solamom mel- aslomaceous fruit, and large-seeded solanaceous fruit). Sturnira tildae Fruit. Goodwdn and Greenhail (1961) reported finding purplish fruit juice in the stomach of a S. fildae from Trinidad. Sturnira mordax Fruit. 330 SPEC[AL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Howell and Burch (1974) reported recovering the identifiable remains of the fruits of Ci-'iuropogon, Anthuriwu^ Musa, and Cevropki from S. tnordux in Costa Rica. Sturnira ludovici Fruit. Dalquesi (1953) wrote that he observed S. hulovki feeding on tree fruits in San Luis Potosi, Mexico. 'I’schudi (1844) mentioned that 5. oporophUiitn (= S. (udovivi) eats fruit, but he also believed that this species feeds on blixid. Starretl and de la Torre (1964) remarked on S. /ndov/r/ from Costa Rica that had fruit pulp in their digestive tracts, Howell and Burch (1974) reported another speci¬ men from Costa Rica that had consumed fruit. Sturntra erythronios Fruit. The only account containing food habits information is that by Tschudi (1844), who stated that this species feeds on fruit. In addition, he (1844:67) related an incident where a bat he identified as this species bit a sleeping drunken Indian on the nose and became so engorged with blood that it could not fly. The bat was captured and taken back to Europe as a specimen. This bat was undoubtedly a desmodontine and not a Siurfdra, and, inasmuch as Tschudi described the species S. erythromos in this publication (the holotypc was not the same speci* men mentioned above), he may not have w'itncssed the incident personally. Summary. —-No information has been published on the food habits of Smrnira ihomasi, S. magtm, S. bidens, S. maui, and 5. arauuhomasi; nevertheless, the diets of these species most likely include a wide variety of fruits. Duke (1967), relating information from Edwin Tyson, said the foods eaten by Sianura in Panama ‘‘consist mainly of fruits, e.g. Piper, PsidianL"' Gaumer (1917) and Ruschi (I953A:) mentioned insects in the diet of S. Udam-y however, the actual role of insects as food items of Siurnira is unknowm. Genus Uroderma Peters Uroderma bilobutum Various kinds of fruit and insects. Most references that allude to the food habits of U. biiohanim simply state that this bat eats fruit or is a frugivore (Goodwin, 1946; Tamsitt and Valdivteso, 1965; Villa-R., 1967; Duke, 1967; Walker er ai, 1964; McNab, 1969). Bloedel (1955) reported U. hdohanmi eating the pericarp of small unidentified palm fruits in Panama. Fruits, particularly of three species of Ficus, w'ere recorded by dc Carvalho (1961) as food items for this species in Brazil. Goodwin and Green- hall (1961) mentioned finding the remains of Psidium guajava in the stomachs of two U. hilohatum in Trinidad. Fleming, ei ai (1972) cited 405 stomachs of U. hilohatam they examined from Costa Rica and Panama. Of these, 320 contained BIOLOGY OF THE PHYLLOSTOMATfDAE :?3i food remains consisting of approximately 76 per cent plant matter, 13 per cent insects, and 11 per cent unclassified material, by volume. Howell and Buref (1974) reported finding B/osimufn in one and an unidentifiable green fruit in the other of the tw'o V. hilohatu/n they examined in Costa Rica. The diet of Urodemui magninfsnitm has not been reported; how ¬ ever, it probably includes fruit, flower products, and insects. I collected several U. niagfitrostnun at Balia on the Rio Curanja, Departamento de Loreto, Peru, the fur of which was stained yellow' from flow'cr pollen or the heads and shoulders of w'hich were dusted wdtb pollen. Both species of Uroderma likely are frugivorous but many also consume quantities of pollen, nectar, and insects found in flowers and fruit as well. Genus Vampyroi^s Peters Vampyrops vittatus Fruit. Tuttle (1970) reported that he netted several V. vlttams in Peru that wer; carrying large figs (F/rus)- Howell and Burch (1974) listed Cecropia and Acnisfits as food items eaten by this species in Costa Rica. Vampyrops dorsalis Fruit and insects. Arala et al. (1967) reported on the stomach contents of four Colombian Vnnipyrops identified as V. dorsalis. Three of the stomachs contained plant material and one contained insect remains. Their paper does not indicate whether the stomach containing insects was one of the three with fruit or w'as the fourth they examined. Vampyrops helleri Fruit. Goodwin (1946), reporting on Costa Rican V. helleri^ stated that it is a fruit- eating species. Villa-R. (1967) also reported that Mexican F. helleri are frugi¬ vorous. The remains of fruit have been noted in most analyses of stomach contents (Goodwin and Greenhall, 1961; Starrett and de la Torre, 1964; Arata e! a(., 1967; Fleming er al., 1972); however, Howell and Burch (1974) reported two Costa Rican V. helleri that had eaten both fruit (Cecropia) and insects (Lepidop- lera). The other eight they examined had been feeding on the fruit of Acnistus. Vampyrops lineatus A variety of fruits and insects. Ruschi (1953jf) recorded the foods of Brazilian V. lineatus as various fruits and insects {especially lepidopterans of the family Sphingidae). McNab (1969) l e- ported the diet as fruit. SPECIAL PUIiLJ CAT JONS MUSEUM TEXAS lECH UN I VERS HV 3:12 Summary.^ —Nothing has been reported on the food habits of Vampyrops iti- fasciis, K awariHS^ V. ni^eilus, V. hrachycephalas, and K recifittus. T he diets of these and other species of Vampyrops probably consist of a variety of fruits, some insects, and possibly some tlower products. Walker ef aL (1964) and Duke (1967) presumed Vampyrops lo be frugivorous. Genus Vami’yrodes Thomas Vampyrode.s earacdoloi Fruit. Vampyrodcs caravcioloi is considered to be a frugivore (Goodwin, 1946; Walker ei «/., 1964; Duke, 1967). Goodwin and Green hall (1961) and Fleming ei af. (1972) reported the contents of the stomachs they examined as consisting entirely of the remains of fruit. Genus VAMtn RPSSA Thomas Vampyressa pusitia Fruit. Goodwin (1946) considered Costa Rican K minafa (=K pusilia) to be frugivorous. Starrett and de la Torre (1964) reported finding a small amount of fruit pulp in the digestive tract of a K fhyone (= V. pusilla) from Nicaragua. Fleming ef al. (1972) reported the stomach contents of one V. pusilla out of the eight they examined from Panama as 100 per cent plant material. The others apparently were empty. Howell and Burch (1974) listed five from Costa Rica that had fed on the fruit of Actiistas. Sitmmary\^Thc diets of Vcmipyressa melissa, V. nymphaeay K hrocki, and V, hhiens are not known. These species probably subsist primarily on fruits as was suggested for the genus by Walker e( aL (1964) and Duke (1967), Genus Cfurodkrma Peters Chirodemia villosum Fruit. This species is presumed to be frugivorous (Goodwin, 1946; Goodwin and Greenhall, 1961; Villa-R., 1967), Chirodemia salvini Fruit. Fruit-eating habits were reported by Goodwin (1946), Jones ei aL (1972) implied that C. saivinl eats figs inasmuch as they mentioned catching one along with An the us and Siurnira in a net under a fig tree replete with ripe fruit in Sinaloa, Mexico. BIOLOGY OF THE PHYLLOSTOMAT[DAE 333 Chirodcrma trinitatiiin Fruit. A diet of fruit was suggested by Goodwin and Grcenhall (1961). SitmmaryK —Nothing has been reported on the diets of Chiroderma dorkie and C. improvssum. Although the fruit diets of C. villosum^ C saivinh and C\ fhniiatiim are based only on conjecture, these species probably do subsist primarily on fruits as suggested by Walker et ai. (1964) and Duke (1967). Genus Ectophylla H. Allen Eetophytla alba Presumedly fruit. Casebeer ei al. (1963) reported, finding small amounts of unidentified green vegetable matter in the lower intestine of E. alba from Costa Rica. I also found similar material in the digestive tracts of five Costa Rican specimens. The food habits of EcWphyila macconneUi are not known; how¬ ever, this species most likely is frugivorous. Duke (1967) mentioned that ths food habits were not known for species of Ectophyila and attributed to Edwin Tyson the opinion that Panamanian species are insectivorous. Genus Aktibeus Leach Artibeus cinereiis Fruit and insects. Goodwin and Greenhatl (1961) stated that A. c'mereus eats a variety of fruits in Trinidad, and Piccinini (1971) mentioned that this species is frugivorous in Brazil. Arata et ai (1967) noted that the stomachs of five Colombian specimens contained plant material and one of these held insect remains as well, Artibeus wafsotii Fruit. Fleming et ai (1972) reported only finding plant matter in two of the f3 stomachs of A. watsoni they examined from Costa Rica and Panama, The other 51 stomachs were empty, Howell and Burch (1974) were able to identify Cecropia as the fruit eaten by the two A. watsoni they examined in Costa Rica. Artibeus phaeotis Fruit. Villa-R. (1966) reported that A. turpis ( — A^ phaeotis) is frugivorous in Mexico. Fleming et ai (1972) examined the stomach contents of 90 A. phaeoAs from Costa Rica and Panama. Of these, only two contained food, which was 100 per cent fruit pulp in each case. Heithaus et al. (1975) determined that 40 per 334 SFECIAJ- l»UH[JCATtONS MUSEUM TEXAS TECH UNIVERSITY cent of 15 A. pimeotis in Costa Rica were carrying pollen when captured. Of these, 33.3 per cent carried two or more species of pollen. They tbund seeds in the feces of 8 per cent of 25 of these bats but did not identify any of the fruits con¬ sumed, I’he five most common pollens recovered were Cciha penia/icira, Cre- sveniia sp., Ocfiroma (a^fopu^, Pseiuloho/?}bax septincrntnu and liynienaea COurban I. Artibeiis foltecus Fruit. Villa-R, (1967) reported observing A. tolfectis eating the fruits of “amate prieto" (Fk'tts pmiifolia) in Mexico. Cecrt^pia was listed by Howell and Burch (1974) as the food eaten by the six they examined in Costa Rica. Artibeus hirsutus Presumedly fruit. Jones £7 aL (1972:13) wrote: “A specimen from . . . [Sinaloa, Mexico] was shot as it sought food in a strangler fig [FtVn,v coi in (folia].'* Villa-R. (1967) suggested that the food habits are similar to those of A. jafnaicensis. Artibeus jamaicensis Insects and a variety of plant materials such as fruits, flower products, and leaves. Osburn (1865) reported finding the kernels of Hrosimum strewn on the floor of a cave in Jamaica inhabited by Ardiheus carpolegus ( = AniheusJamaicensis): Some of the nuts (p, 64) had “germinated into young blanched trees on the thick deposit of dung.” In other Jamaican caves used by this species he found dried seeds, berries of Cordia collocovca, and husks that included gnaw'cd fragments of unripe mangoes and the fruit of the rose-apple (Eugenia jamlhys). He (1865:66) also mentioned finding yellow juice and small seeds that he suspected were those of the fustic (Monts n'nctoria) in the digestive tract of a specimen. Ortiz dc la Puente (1951) related finding the male inflorescences of maize and, oc¬ casionally, seeds of Erk^botrya japonica under roosts of A. janiaicensis in caves in w'estern Perii. Van dcr Fiji (1957:294) referred to Heinz Felten’s observations (personal communication) on regularly finding remnants of Spondias purpurea under colonies in caves in El Salvador, Bond and Seaman (1958) remarked that seeds and partly eaten fruits of mango, East Indian almond, hogplum, and other easily recognized food items were abundant under A. Jamaicensis roosts in the Virgin Islands. Goodwin (1970:575) stated that the presence of A. Jamaicensis in caves in Jamaica is usually indicated by a “garden" of pale, spindly, seedling plants growing on the floor beneath the roosting site. He identified the plants from two caves as Andira inermis and observ'cd that the fruits of this tree are a staple food of Anibeus. Allen (1939) mentioned the presence of sprouted nuts of Acroconua in a cave in Puerto Rico and stated that A. jamaicensis was fond of the thin layer of pulp surrounding the small nullike fruits of this palm. Tuttle H[0[,OGY Ot"‘ [HE PHYLLOSrOMATfDAE 335 (1968) remarked on finding the remains of several kinds of fruit on a large pile of guano beneath an A. Januiiccn,sis roost in Chiapas, Mexico. The remains in¬ cluded many hard nuts, each of which had been chewed open at one end. Beneath the roost, Tuttle (1968) also found discarded leafy twigs of which many of the leaves w-ere chcw'ed and appeared to have been partly eaten. While conducting his reconnaissance on the roosting site, he recorded the following observations (p. 787): “While I sat quietly a few' feet below' them, the bats began to catch and eat large (about 6 mm long) blackflies. The bats w'ould hang by one foot and rotate in nearly complete circles w'atching the dies. W'henever a tly tlew' w ithin reach of one of the bats, the bat would capture it w ith a rapid thrust of one of its wings. Flies were caught in the wing-tips and were immediately eaten. This be¬ havior was observed repeatedly.” Ouelch (1892:102) described the foraging behavior of A. Jamaicetixis in British Guiana (Guyana): “During the fruiting season, when the sapodillas, star- apples, mangoes, and such like fruit are ready to be gathered, numbers of these large bats are to be observed at sunset, flitting in and out among the leaves and branches, picking out and feeding on the ripest fruit to be found.'They dart up and down repeatedly at the same fruit, remaining momentarily almost stationary while their teeth are applied, and with the force of their tlight they cause either the tearing away of part of the soft pulp, or of the whole fruit, according to its degree of ripeness.” Jimbo and Schwassmann (1967) reported A. Jamakensis feeding on sapodiila plums {Achnis sapofa), guava i Psidiuni gnajava), and figs {Ficus sp.) at Belem, Brazil. The fruits, some weighing as much as 50 grams, were carried off. If the fruit w as dropped, the bat sometimes w'ould drop to the ground and eat part of it before flying away. Tamsitt and Valdivieso (1961) mentioned two ck- casions in Costa Rica w'hen A, ja/uaicensis enteral mist nets while carrying pomar- rosa fruit {Syzigium Jambos) in their mouths. One of the fruits measured 34.8 millimeters in diameter. Tuttle (1970) related catching an A^Jamaicensis in Peru that was carrying a large (about 30 millimeters in diameter) wild fig in its mouth. I recovered poniarrosa and guava fruit carried into a mist net by A, Jamaicensis in Villaviccncio, Colombia. One of the larger guavas measured 42 millimeters in its greatest diameter and weighed 35 grams. A second measured 48 by 42 millimeters and weighed 50 grams. The largest was not weighed but measured 64 by 50 millimeters. Jones et aL (1972:13) recorded observations on A.janiai- censis in Sinaloa, Mexico, and stated, “individuals of this species w'ere seen emerg¬ ing from a hollow limb of a fig tree. They foraged higher in the tree, sometimes returning in approximately 10 minutes with cut green figs to the hollow'." Dalquesl (1953) recorded this bat feeding on fruits such as jobo plums (Spondkis sp.) and green wild figs {Ficus sp.) in San Luis Potosi, Mexico. He noted that the mouths of caves used as day roosts were commonly heaped w'ith cores and seeds of fruit and small pellets of fruit skin and rind, w'hich the bats eject w'hen they eat. Hall and Dalquest (1963) also mentioned jobo plums and wild figs as foods of A. jamaicensis in Veracruz, Mexico. Vazquez-Yanes et al. (1975) reported the kinds and percentages by weight for each month of occurrence of the fruits they re¬ covered from a cave inhabited by A, jamaicensis in the Tuxtlas region of Vera- SPECIAL PUBL1CAT[0NS MUSEUM TEXAS TECH UNlVE.RSiTY nfy cmz, Mexico. The fruits they identified are Cecropia oblusifolia, Sfomiias nwrnbitu Ficus spp., F. glabmui^ F. oblusifaiia, Poulsenki armaia, Cyn mietra retusa, Calophyllum htasiliense, Brosinmm alicasirurn. Piper anriturn, P. his¬ pid urn, P. anialago, P. sancium^ Turpinia piunata^ Sc^Ia/tum spp., Dendropafia.x: arhoreus, Quararihea funehris, AfUhurium sp., Licania sp., Mimfingia ca*ahura, Pseudolmedia oxypfiyllarki, and Rheedia eduiis, Dalquest et aL (1952) reported on the mucous salivary glands opening in the lips at the ventral angle of the lower jaw in A, jamaicensis. They interpreted the function of these glands as supplying the mucous that binds together the pelletized ejected unpalatable portions of the fruit these bats eat. Greenhall (1956, 1957J and Goodwin and Greenhall (1961) presented nearly identical lists of foods eaten by A, Jamaice/isis (Table 3). Their information was based primarily on debris found beneath roosting sites in Trinidad. De Carvalho (1961) described a number of fruits utilized as food by A, jauiaicettsis in Brazil (Table 3). Villa-R. (1967) noted a number of fruits in the diet of Mexican A, Jamaicensis as w'ell as pollen and nectar from the flow'ers of Ceiba penfandra (Table 3). He also related having observed A. Jamaicensis entering a small house in San Luis Potosi, Mexico, to eat cakes of sugar that were stored near the ceiling. Goldman (1920) commented on catching several of this species at Gatun, Panama, in traps placed about a bunch of ripening bananas. Starrett and de la Torre (1964) gave the stomach contents of A. Jamaicensis from Nicaragua and Costa RJca as fruit pulp, plant fibers, and bat hairs. They also noted (p. 61) that a specimen ‘Troni Costa Rica also had an ant (Formicidae: Ponerinac) embedded in a reddish amber-like substance in its intestine.” Arata et ai (1967) related finding only plant material in the stomach of a specimen they examined from Colombia. Fleming et al. (1972) reported on the 23 stomachs containing food material among the 916 digestive tracts they examined from Costa Rica and Panama. The Stomach contents, by volume, consisted of about 66 per cent plant matter, 25 per cent insect remains, and 9 per cent unclassified material. They aiso ex¬ pressed the opinion that figs (Ficus insipida) were a favorite food of A. Jamai- censis in the Panama Canal Zone. Howell and Burch (1974) identified insects (Coleoptera), pollen (Hymenaea, Ceiba, and Bombax), and fruit (Ucania, Genipa, Munfingia, Brosimum, Ficus, Cecropia, and melastomaceous fruit) as the food items they recovered from A. Jamaicensis in Costa Rica. Palmer (in Miller, 1904:347), in reporting the habits of A. Jamaicensis in Cuba, wrote that “they evidently capture much of their food among flowering trees, as their fur often contains pollen and parts of flowers. These are also found abundantly on the floors of caves where the bats roost.” Silva Taboada and Pine (1969) implied that either Palmer’s observations were unusual or were in error as they had never found flower parts in the fur of Cuban Artibeas. However, Piccinini (1971) noted that several A, Jamaicensis collected in Brazil during October were stained yellow by the pollen of Atiacardium occidentaie and he assumed that these bats, although primarily frugivorous, eat pollen when fruits are not available. Heithaus et ai (1974) recovered Bauhinia pauietia ptdleri from the fur of tw'o A. jamaicensis near Cahas, Costa Rica. Heithaus et ai (1975) BIOLOGY OF THE PHYLLOSTOMATIDAE 337 reported recovering pollen from the fur of 54.1 per cent of 477 individuals in Costa Rica of which 43 per cent were carrying two or more species of pollen. Only 8.6 per cent of 617 A.Jamaicensis had seeds in their feces. The six most common pollens they recorded were Crcsce/uui sp., Maniikara zapoia, Hymenaea coitr- haril, Pseudobomhm septimitum, Ocirronut lagopus^ and Ceiha peniandnL The identifiable fruit remains they recorded were Piper tuheraiUuum, Soianum spp., Miinlingia calabura, and Ficus sp. Hall and Kelson (1959) described the food of A, Jcvuaicensis as mainly ripe fruits, the small kinds of which are plucked and car¬ ried to the feeding sites. McNab (1969) and Tamsitt and Valdivieso (1970) merely stated that the species is a frugtvore. Artlbeus lituratus Insects and a variety of plant matter including fruit, flowers, and leaves. Valdivieso and Tamsitt (1962), Tamsitt and Valdivieso (1965), and McNab (1969) considered A. litunifus to be frugivorous. Dafquest (1953) reported on finding the gound beneath the roosts of this species in San Luis Potosi, Mexico, littered with the small, brown pellets of rind and skin of fruit ejected by the bats as they fed. Van der Pijl (1957:294) quoted Heinz Feltcn (personal communica¬ tion) who told him of regularly finding remnants of Ficus sp. under colonies in caves in El Salvador, Bloedel (1955:235), writing about Panamanian bats, men¬ tioned that A. lituratus'^droppfdd many spave beans below their habitual roosting place.*’ Villa-R. and Villa Cornejo (1969, 1971) reported that they observed numbers of A. liturams taking either ripe palm fruit or pollen from flowers in northern Argentina. I'amsitt and Valdivieso (1963) commented on twice netting A. lituraius thal were carrying ripe almond fruits in their mouths. Green- hall (1956, 195?) and Goodwin and Greenhall (1961) listed a number of plant species utilized in the diet of this species (Table 3). The plant fcx>ds listed by Goodwin and Greenhall (1961) were cited as foods for both A, Hturatus and A, jamaicensis. Villa-R, (1967) also stated that these two species have similar food habits in Mexico. Ruschi (1953u) presented information on the stomach contents of A. Jamaicensis Uturaiitsi — A, liiunutisl) from Brazil. He claimed (p. 3) to have found coagulat¬ ed blood in addition to fragments of fruit in the stomachs of these bats. Ruschi (1953y), in addition to listing a number of the principal fruits eaten by these bats (Table 3), elaborated on his earlier report on finding blood among the stomach contents. He referred to capturing several A. lituraius alive in a palm tree and later finding blood in their stomachs. I recommend, however, that blood not be considered a normal food for Artibeus; the blood in the stomachs may be explained as having come from the cleansing of wounds acquired during fighting among the bats as they were held together subsequent to their capture. Nevertheless, in support of his opinion on the alleged blood-feeding habits of this species, Ruschi men¬ tioned (p. 7) having surprised an A, lituradis in the act of eating nestling robins (Turdus rufiventris) and noted that these bats accepted blood, in addition to fruit and insects, as food in captivity, Ruschi also observed A. lifuranis pursue and capture sphingid moths. SPECIAL PUBL1CAT[0NS MUSEUM TEXAS TECH UNlV.ERStTV Starrett and de la Torre (I964;6( ) remarked on finding “fruit pulp of several colors and types, plant fibers and bat hairs . . . [as well as] a few smail scattered insect remains” in the digestive tracts of A. fiturams from El Salvador, Niciragua, and Costa Rica. Arala er aL (1967) reported on four Colombian A. they examined; three stomachs contained plant materia! and one contained insect remains. Fleming e/ aL (1972) reported the contents of seven out ot the 93 stomachs of A. /tVwmtuY they examined from Costa Rica and Panama as 75 per cent plant matter and 25 per cent insect remains, by volume, Howell and Burch (1974) listed a large-seeded Piper as having been consumed by a Costa Rican A, liiuraiNS. Heiihaus et ai. ( 1975) identified the pollen of Crescent in sp., Ochronui lagopits, Ceiha penwndra, and Manilkara zapotn taken from the fur of representa¬ tives of this species in Costa Rtca. Summary. —The diets of Artiheus glaiwus, A. aztecus, A. inopmatus, and A. coticolor are not known; however, these species probably have feeding habits similar to those of the other species of Aniheus previously mentioned. These bats are primarily frugivorous but consume pollen, nectar, flower parts, and inf ects us well. Duke (1967) cited Acntcamia, Aiiacarditim^ Brosimumy Cecropiu, Cortliay Pugetiiay FicitSy Mangiferuy MusOy and Persea as plants of which fruit is utilized by Panamanian Anibeus. Howell and Burch (1974) reported six small Artibeits sp. that had fed on Cecropia fruit in Costa Rica. The finding of relatively large volumes of insect remains (25 per cent) in the stomachs of Costa Rican anc Pana¬ manian A. Jamakensis and A. lituratus {FiQming et «/., 1972) indicates that in¬ sects are an important food source for these bats. The utilization of bloi>d and small vertebrates as foods by A. ///mY//ns (Ruschi, 1953ri; 19537) is to be considered atypical. Genus Enchisthenes Anderson Enchisthenes hartii Fruit. Goodwin (1946) and Goodwin and Green hall (1961) stated that E. h'^nii is frugivorous. From observations on this species in the vicinity of Ciudad Guz¬ man, Jalisco, Mexico, de la Torre (1955:700) wrote: “The fruit [Ficus sp. eaten in the area is small, about a centimeter in diameter. It is quickly snipped from the tree in flight, and carried to a convenient branch where it is eaten,” Villa-R. (1967:319), reporting on two specimens from the same area in Jalisco, claimed also to have observed E. hartii plucking ripe figs. Genus Ardors Miller Ardops nichollsi Presumably fruit. Walker et al. (1964) commented that A. nicholisi was presumed to damage cacao {Theobroma cacao) by eating the fruits. Additional information on its food habits is lacking. BEOLOGY OF THE PHYLLOSlOMATlDAE 3.^9 Genus pKYi.LOPS Peters Phyllops fakatus and PhyLlops haklensis Presumably wild figs. Hall and Kelson (1959) employed the common names “Cuban Fig-eating Bat” (P. faicaftis) and “Dominican Fig-eating Bat” (P. haitiensis) for these species. Allen (1942) also considered them to be frugivoroiis; however, detailed informa¬ tion on their food habits is lacking. Genus Arj t ELfS Gray Ariteus flavescens Fruits and insects. Gosse and Hill (1851) reported A. achrmi(}phi!ns ( = A. fiavesa^ns) eating the fruit of the naseberry tree {Achras sapoia) in Jamaica. The bats either fed on the fruit in the tree or carried large pieces away to be eaten elsewhere. They also mentioned that this species feeds on the rose-apple {Eugenia jamhos). Walker et al. (1964) repeated this information, and remarked that A. Jlavescens eats insects as w-ell. Hall and Kelson (1959) referred to A. flavescctt.s ii& the “Jamaican Fig-eating Bat.” Genus Si ENODERMA E. Geoffroy St,-Hilaire Sktioderma rufum Fruit. Although considered to be a frugivore (Allen, 1942; Tamsitt and Valdivieso, 1970; Genoways and Baker, 1972) the diet is unknown. According to Thomas (1894:132), S. rufum is “said to do much damage to the cacao plantations” on the island of Montserrat, Hall and Kelson (1959) applied the name “Red Fig-eating Bat” to this species. Genus Pygqoerma E. Geoffrey St.-Hilaire Pygoderma biJabiatum Probably fruit. The species is alleged to feed on fruits (Goodwin, 1946; Walker et a!., 1964). Genus Ame i rida Gray Ametrida eenturio Probably fruit. The diet is unknown. Genus SpHAERONYCTERts Peters Sphaeronycteris toxophylEuni Probably fruit. The diet is not known. .140 SPEC[AL PUBLJCATIONS MUSEUM TEXAS TECH UNIVERSITY Genus Centurjo Gray Centiirio senex Fruit, Goodwin and Greenhall (1961) mentioned finding yellow' fruit pulp in stom¬ achs of C. seuex from Trinidad, Walker ei al. {1964) gave the diet as “soft mushy fruits,” Felten (1956) may have meant to imply that C senex feeds on figs {FictfS sp.) in E] Salvador when he mentioned collecting a specimen when it wai flying around a fig tree. Subfamily Phvllonyctekinae Genus Brachyphylla Gray Brachyphylla cavernarum Fruits and insects. Bond and Seaman (1958:151), reporting on a roost of B. cavenuirufn in the Virgin Islands, noted, “an examination w'ith a hand lens of the washed guano show's it to contain a high proportion of insect fragments, and some amorphous material which may or may not be fruit pulp. Seeds and partly eaten fndts are [present] in Ariiheiis guano. No such materia] w-as found in the guano of Brachyphylla, although a few' small seeds of what appears to be a species of Eugenia were seen. These seeds could have passed through the bats or been brought in by mice. These observations could mean either that Brachypnylla is entirely insectivorous, or that it eats fruit but avoids small seeds, and cannot carry larger-seeded fruits back to the roost as does Anibens," Nellis (1971) found B, cavenmrum on St. Croix, Virgin Islands, feeding on the fruits of Manilkara zapoia. Hall and Kelson (1959) and Tamsitl and Valdivieso (1970) referred to this species as the “St. Vincent Fruit-eating Bat.” Brachyphylla nana Fruit, pollen, nectar, and insects, Silva Taboada and Pine (1969) presented information on the contents of 43 stomachs of B. nana from Cuba, All stomachs contained masses of what ap¬ peared to be partially digested pollen grains. One stomach contained lepidepteran scales and another held fragments of a fly (Diptera). In Cuba, Silva Taboada frequently encountered individuals the head, chest, and shoulders of whica were dusted with pollen. Silva Taboada and Pine (1969:15) considered B. mma, as well as Cuban PhyKonycteris and Erophylla, primarily to be “pollen eaters which probably also feed on soft fruit pulp and nectar.” Genus Erophylla Miller Erophylla bombifrons Fruit, pollen, nectar, and insects. Hall and Kelson (1959) referred to E. bombifrons as the “Brown Flow^er Bat”; however, Tamsilt and Valdivieso (1970) said it is frugivorous. BIOLOGY OF THE PHVLLOSTOMATIDAE Ul Erophylla sezekami A variety of fruits, pollen, nectar, and insects. Osburn (1865) reported on finding breadnut kernels and munched berries of the clammy cherry {Omiia colkKocca] associated with cave roosts of bats he referred to as Monophyilus { = Eraphyifa sezekurni) in Jamaica. The stomach of one specimen he examined (p. 82) “w-as filled w'ith a yellowish frothy pulp,” Osburn described the feeding behavior of a captive: “The tongue was rapidly protruded and drawn in again, and the juice and softer pulp cleared away with great rapidity. I noticed he was very particular in cleaning out the bit of loose skin of the berry \Cordia vullococca]''' Osburn also noted (p. 84) the simi¬ larity of the berries eaten by this bat and those found beneath the cave roosts. Silva Taboada and Pine (1969) presented information on the stomach contents of 30 E. sezekorni from Cuba. They found masses of partially digested pollen grains in all stomachs. In addition, three contained seeds of bromeliaceous fruits {Hohenbergki) and four held insect remains identifiable as parts of an elaterid beetle [Conoderiis)^ a roach (Orthoptera, Blattidac), Diptera, Lepidoptera, and Microlepidoptera. Silva Taboada and Pine (1969:15) expressed the opinion that E. sezekorni (along wdth Cuban Brachyphylia and Phydonyaeris) “arc primarily pollen eaters which probably also feed on soft fruit pulp and nectar.” Hall and Kelson (1939) referred to E. sezekorni as the “Buffy Fruit Bat.” Genus Phyllonycteris Gundlach PhyJlonyeleris poeyi Probably a variety of fruits, pt>llen, nectar, and insects. Allen (1942:26-27) commented that P. poeyi have “long protrusible tongues, which are useful in licking up fruit pulp and juices on which they largely feed. Probably pollen and nectar are also eaten.” Silva Taboada and Pine (1969) re¬ ported the stomachs of 42 P. poeyi they examined from Cuba as containing masses of what appeared to be partially digested pollen grains. One stomach also contained lepidopteran scales. They commented (p. 15) “that the Cuban rep¬ resentatives of Brachyphylia, Phylkmyaeris, and Erophylla are primarily pollen eaters which probably also feed on soft fruit pulp and nectar.” Summary .—The diets of Phyllonyaeris major and P. aphylla are not known; however, these “Flower Bats” (Hall and Kelson, 1959) likely have food habits similar to those of P. poeyi and include fruit, pollen, nectar, and insects in their diets. Su B FA M1 1 . Y DE SMOIK)N ! 1N A E Genus Des.moous Wied-Neuwied Desmodus rotu nd us Blood of warm-blooded animals. The folklore surrounding the sanguivorous habits of D, rofimdus, enhanced by the imaginations of the early explorers and naturalists who visited the New World 342 SPECIA). PUBLICATIONS jMUSEUM TEXAS TECH UNIVERSITY tropics, produced fascinating narratives on the bloodsucking vampires. A few of these accounts have been related by Husson (1962:12-18, 192-197). Most of the early naturalists ascribed awesome proportions and properties to vampiies and mistakenly attributed the feeding habits of the desmodontines to Vim.pyrum spectnmu Phyilosumuts hastaius, and Chrotopterus aiiritus^ among other bats. The majority of the reports concerning the prey of D. ronuuius has been limited to man and his domestic animals. Darv^dn (Waterhouse, 1838:2) was apparently one of the first to witness D. roiundus on its prey and, on this subject, Waterhouse observed, '‘Before the introduction of the domesticated quadrupeds, this Vampire Bat probably preyed on the guanaco, or vicugna, for these, to¬ gether with the puma, and man, were the only terrestrial mammalia of large size, which formerly inhabited the northern part of Chile,” Dalquest (1955) did not find scars from vampire bites on any of the white-tailed deer, brockets, collared and white-lipped peccary, and tapir he examined from eastern Mexico. Ap¬ parently the selection of prey varies from one region to another because Dalquest (1955) observed that Desmodus seems to prefer the blood of burros and calves over that of horses and adult cattle, and does not appear to feed on domestic fowl in Veracruz, Mexico. In eastern San Luis Potosi, Mexico, however, Dehmodus extensively preys on domestic turkeys and chickens as well as on adult catllc. On the Mexican Plateau in western San Luis Potosi, horses, cattle, and burros again are favored, but chickens arc rarely molested. Despite clear evidence that children had been bitten by vampire bats in both Veracruz and San Luis Potosi', their parents denied that bats were responsible. Instead, they attributed those bites to bntjas (witches). Dalquest (1955) also stated that children seemed to be bitten more often than adults, and, of the latter, w'omen more often than men. Many authors (see Linhart, 1971, for a partial bibliography of the literature pertaining to vampire bats), have mentioned D, rotundus feeding on doiiestic livestock, poultry, and humans, Tw'o of the more informative accounts are those of Gotxlwin and Greenhatl (1961:268-269) and Villa-R. (1967:30-37, 334- 336). Detailed information, particularly relating to observations on the m^xle of attack, biting, bite sites, and feeding behavior, has been reported by Mann (1951), Greenhall (1972), Greenhall et at. (1969, 1971), and Turner (19751. W'imsatt and Guerriere (1962) and W'imsatt (1969) suggested a conservative daily consumption of 20 milliliters of blood per day or about 7,3 titers per year per individual. They projected a minimum annual consumption of 730 liters of blood for a moderate-sized colony (approximately 100 adults), This is about one-third the estimated consumption per individual postulated by Goodwin and Greenhall (1961:269). Miller (Allen, 1916:603) remarked, “It is said [in Brazil] that blood-sucking bats vary their diet w-ith insects.” Arata ei aL (1967) rej>orted on the stomach contents of 23 D. rotundus from Colombia. Although all stom¬ achs contained blood, four contained insect remains and four contained flesh. They stated (p. 654) that “the insect remains found in Desmodus stomachs con¬ sisted of a whole ingested ectoparasite, .. and well-broken remains of larger insects.” Rouk and Glass (1970:456) stated that “a few insect remains [found] in one specimen ... supports the observation of Arata {et al.^ 1967) [s/c].” BIOLOGY OP THE PHYU-OSTOMAflDAE 343 Greenhall (J972:485), commeniing on these observations, remarked, “1 have examined thousands of Des/rKnUis stomach contents and have also found the remains of some insects as well as flesh 'divots.' Presumably the insects be¬ came trapped in the viscous blood and were thus swallowed.” The insects also could have become caught in the congealing exudate from the wound and ingested with the scab at a later bout of feeding. The ectoparasite w'as probably con¬ sumed cither during grooming activities by the bat or when preparing the bite site on a prey animal. Villa-R. et al. (1969) related the results from analyzing the stomach contents of 79 D. roiumlns from Brazil. They found 58 (73.4 per cent) contained mam¬ mal blood, 8 (10.1 per cent) with mixed bird and mammal blood, and 13 (16.5 per cent) held milk. The 13 containing milk w-ere judged to be juven¬ iles although they w'ere the size of adults. These investigators concluded that D. romndiis tends to prefer the blood of mammals and augmented their observations by noting (p. 296) that Guillermo Mann had communicated to Amelio Malaga Alba information on the predation by Dcsniodi^s on seals (Otariidae) along the Pacific coast of Chile. Genus Diaemljs Miller Diaemus youngU Avian and mammalian blood. Very little is known of the food habits of D. youn^^fl except that it seems to prefer avian blood. Gotxlw'in and Greenhall (1961) stated that this species preys upon poultry, pigeons, and goats in Trinidad. They commented {p. 272), “Usually, when poultry and goats are attacked in an area, to the obvious exclusion of cattle and cc]uines, Diaemus have been collected.” Vilta-R. (1967:341), relating a personal communication from Reaznet Darnell, reported that a D. captured near El Encino, Tamaulipas, Mexico, was feeding on the blood of a chicken. Genus Diphylla Spix Diphylla ecaudata Avian and mammalian blood. Moojen (1939:7) reported D. ecaudaia preying on chickens in Brazil. Ruschi (1951) reported D. ecauchta, also in Brazil, feeding on the blood of mammals and birds. He mentioned (p. 2) domestic chickens, turkeys, Guinea fowl, ducks, and geese as preferred prey, but stated that Diphyfla feeds on pigs, cattle, and equines as well Later, Ruschi (I953^i:7) described a D. ecaiutata feeding on a sleeping man in the state of Bahia, Brazil. Dalquest (1953:40-41) wrote, “Vampire bats, probably of this species, attack chickens near Xilitla [San Luis Potosl, Mexico], biting them on the lower part of the leg where the feathers are scant, Diphylla ecaiuiata probably preys on horses, burrows, and cattle.” This is essential¬ ly the same information given by Walker e/ ai (1964} and Duke (1967). Koopman (1956:548) cited a specimen from San Luis Potosi, that “according to the field 144 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITV tag, was killed while feeding on chickens.” Villa-R. (1967) stated that D, ecaudaia appears to prefer the blood of birds. The stomachs of 18 Brazilian D, ecaiukua examined by Villa-R. et ai (1969) contained bird blood exclusively. Literature Cited A^.cott^, S. M., S. E. McGregor, and G. Olin. 196L Pollination of saguaro ca;lus by doves, nectar-feeding bats, and honey bees. Science, 133r 1594-1595. An EN, G. M. 19.19. Bats. Harvard Univ, Press, Cambridge, x + 368 pp. --. 1942. 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G., and B. J, Harris. 1957. The pollination of Purkki by bais and its attendant evolutionary problems. Evolution, 11:449-460. Barhour, R. W., and W. H. Davis. 1969. Bats of America. Univ. Press Kentucky, Lexington, 286 pp. Bates, H. W. 1875. The naturalist on the River Amazona. London, 3rd ed., x-l-394p.p. Bloedel, P. 1955. Observations on the life histories of Panama bats. J. Mamn., 36; 232-235. Bond, R. M., and G. A. Seaman. 1958. Notes on a colony of Bnn hyphyllu caven timni. i. Mamm., 39:150-151. Brosset, a, 1965. Contribution a Tetude des Chiropteres de TOuest de (’Ecuador. Mammalia, 29:211-227, -. 1966. La biologic des Chiropteres. Masson et Cie„ Paris, vii + 240 pp. Burt, W. H. 1938. Faunal relationships and geographic distribution of mammals in Sonora. Mexico. Misc. Publ. Mus, ZooL, Univ. Michigan, 39:1-77. Burt, W. H., and R. A. Stirton. 1961. The mammals of El Salvador. Misc. Publ Mus. Zook, Univ. Michigan, 117:1-69. Carter, D. C., R. H. Pine, and W. B. Davis. 1966. 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BioL Prof. Mello-Leilao, Sanu Teresa, ZooL, 13:1-10. --. !953(/, Morcegos do Estado do Espirito Santo. XIL FamlJia Phyllostomidae: descrigao das especies: Micnmyaeris mepudons tnegaiotis e PhyUostomus hastuttis husfaius, com algumas observagoes biologicas a respeito. BoL Mus. Biol. Prof. Mello-Leitao. Santa Teresa, Zoo!., 14; I-14. —-. I953e. Morcegos do Esiado do Espfrilo Santo. XllL Familia Phyllostomidae, descrigao das especies Mimon hentteitii e Lonchorhitm uuyiuu com algumas observagoes. Bol. Mus. BioL Prof. Mello-Leitao, Santa Teresa, ZooL, 15:1-11 -, 1953/ Morcegos do Estado do Espirito Santo. XIV. Familia Phyllostomidae. Descrigao das especies: DolivhophyUnm macrophyHitm tWied) e ChrifUtpteiitx aurhiis tuistraUs (Thomas). BoL Mus. Biol. Prof. Mello-Leitao, Santa Teresa, ZooL, 16:1-7. ■ 1953^', Morcegos do Estado do Espirito Santo. XV. Familia Phyllostcmidae— Dcscrigao das especies: Glossophaga sorkinu sorIvina e Aaount geoffmyi geoffroyi, com observagoes biologicas a respeito. Bol, Mus. Biol. Prof. Mello- Leitao, Santa Teresa, ZooL, 17:1-12. -. 1953/). Morcegos do Estado do Espirito Santo. XVL Familia Phyllostomidae. Descrigao das especies: Lonchogio^^iia cattdifera e Lotichogh^su evaiidaki, com algumas observagoes biologicas a respeito. Bol. Mus. Biol. Prof. Mello- Leitao, Santa Teresa, ZooL, 18:1-7, -, 1953). Morcegos do Esiado do Espirito Santo. XVII. Familia Phyllostomidae, Descrigao das especies: LonchophyUa mordax e Hemidetma perspkUUitnttu com algumas observagoes biologicas a respeito. Bol. Mus. Biol. ProL Mello- Leitao, Santa Teresa, ZooL. 19:1-7. -. 1953/ Morcegos do Esiado do Espirito Sanlo. XVIIL Familia Phyllostomidae, Descrigao das especies Anthem jamaicensis rttumtus e Vampyrops iinealiiHy com algumas observagoes, Bol. Mus. BioL Prof. Mello-Leitao, Santa Teresa, ZooL, 20:1-8. -1953/:. Morcegos do Estado do Espirito Santo, XIX, Familia Phyllostomidae, descrigao de Tomukt hidens e Sturnira itliam iiiiunr. Familia Vespertilionidae, descrigao de Epteskm /)f7f;n7 e Eptesicm hntsHienais. Familia Mclossidae, descrigao de Eumops ubnisus ahrasits. Com algumas observagoes a respeito, Bol. Mus, Biol. Prof. Mello-Leitao, Santa Teresa, ZooL, 21:1-5. Rustm, A., AND A. G, Bauer. 1957, Classificagao de quiropteros do Rio Grande do SuL Arq. Inst. Pesq. Vet. "Desiderio Finamor," Porto Alegre, Brazil, 2:38-41. Sanborn, C. C, 1936, Records and measurements of Neotropical bats. ZooL Str., Field Mus. Nat. Hist., 20:93-106. -, 1954. Bats of the United States. Public Health Rep,, 69:17-28. ScHALDACH, W. J., AND C. A. McLauchlin. I960. A new genus and species of glos- sophagine bat from Colima, Mexico. Los Angeles Co. Mus., Contrib. Sci., 37:1-8, Silva Taroada, G., ,and R. H. Pine. 1969, Morphological and behavioral evidence for the relationship between the bat genus £frar/iyp/!>7/fj and the Phyllonycterinae. Biotropica, 1:10-19. SiARREtT, A. 1969. A new species of Anouru (Chiropiera: Pbyllostomatidae) from Costa Rica. Los Angeles Co. Mus., Contrib. Sci., 157:1-9. Starrett, a., and L. de i a Torre 1964. Notes on a collection of bats from Central America, with the third record for Cytkirops ulectu Thomas. Zoologica, 49: 53-63. Tamsitt. J. R., and D, Valdivieso. 1961. Notas sobre actividades nocturnas y estados de reproduccion de algunos quiropteros de Costa Rica. Rev. Biol. Trop., 9:21 9-225. -, 1963. Records and observations on Colombian bats. J. Marnm., 44:168-180, -1965. Reproduction of the female big fruit-eating bat, Artibem iituratus pab nuirum^ in Colombia. Caribbean J. Sci., 5:157-166, BIOLOGY OF THE PHYLLOSrOMATIDAE 349 - —1970. Observations on bats and their ectoparasites. Pp. 123-128, hi A tropical rain forest (H. Odum, ed.), U.S. Atomic Energy Commission, Washington, D.C.. xivd- 1640 pp. Tes i . F, H. 1934. Records of bats from Honduras. J , Mamm., I 5:316. Thomas, O. 1894. Descripton of a new bat of the genus S!i’ttoiierit}(t from Montserrat. Proc. Zool. Soc. London, 1894:1.3 2-133. Torre, L. of, i a. 1955. Bats from Guerrero, Jalisco and Oaxaca, Mexico. Fieldiana: ZooL. .37:695-701. -. 1961. The evolution, variation, and systeniatics of the Neotropical hats of the genus Stiiniira. Unpublished Ph.D. dissertation, Univ. Illinois, iv+ 146 pp. Tschloi, J. J. von. 1844. Therologie. ///. Untersuchungen iiber die Fauna Peruana (J, J. von Tschudi. ed,), .St. Gallen. part 1.262 pp. Turner, D. C. 197.5. The vampire bat. Johns Hopkins Univ. Press. Baltimore. x + 145 pp. Tuttle, M, F). 1967. Predation by Chrotopfi iux atifitus on }. Mamm.. 48:319. -. 1968, Feeding habits of/I r/j[7j£'/o /f;ju(oreu.vo. J. Mamm.. 49:787. --—, 1970. Distribution and zoogeography of Peruvian bats, w iih comments on natural history. Univ. Kansas Sci. Bull., 49:45-86, Valdez, R., and R. K. LaVai., 1971. Records of bats from Honduras and Nicaragua. J. Mamm,, 52:247-250. Vai DiviESO, D., and j, R. Tamsitt. 1962, First records of the pale spear-nosed bat in Colombia, J, Mamm., 43:422-423, VAN DER PiJL, L. 1957. The dispersal of plants by bats (chiropterochory). Acta Bot. Neerlandica, 6:291-315. Vauohan, T. A. 1959. Functional morphology of three bats: Eumops, Myotis, Macroius. Univ. Kansas Ptibl., Mus. Nat, Hist., 12:1-153. Vazquez-Yanes. C., a. Orozco. G. Fra.ncojs, a.nd L, Trejo. 1975. Observations on seed dispersal by bats in a tropical humid region in Veracruz. Mexico, Biotropica, 7:73-76. Villa-R., B, 1967. Los murcielagos de Mexico. Inst. Biol., Univ. Nac. Autonoma Mexico, xvi + 491 pp. [For 1966] Villa-R., B., and M, Vili.a Cornejo. 1969. Algunos murcielagos del node de Argentina. Pp. 407-428, in Contributions In Mammalogy (J. K. Jones, Jr., ed.), Misc. Publ. Mus. Nat. Hist., Univ, Kansas. 51:1-428. —^——. 1971, Observaciones acercade algunos murcielagos del norte de Argentina, especial- meme de la biologia del vampiro Desmodns r. roinndns. An. In.st, Biol., Univ. Nac. Autonoma Mexico, Ser, Zool., 42:107-148. VtLLA-R,, B., N. Moraes Da Silva, and B. Vjlla Cornejo. 1969, Estudio del contenido estomacal de los murcielagos hematofagos Desnitulus rotnndns rotitnJns (GeofTroy) y Dfp/iy//£j ecciudtiM ecaiulnw Spix (Phyllostomatidae, Desmodinae). An. Inst. BioL, Univ. Nac. Autonoma Mexico, Ser. Zool., 40-291-298, Vogel, S. 1958. Fledermausbiumen in Sudamerika. Osierr, Boi. Zeits., 104:419-530, Walker, E. P., F. Warnick, S. E. Hamlet, K. I. Lange, M. A. Davts, H, E. Uirle, and P. F. Wright. 1964. Mammals of the World. Johns Hopkins Press, Baltimore. Lxlviii + 644 pp, Waterhouse, G. R. 1838. The zoology of the voyage of HMS Beagle under the com¬ mand of Captain Fitzroy, during the years 1832 to 1836. Part 2, Mammalia, London, xii+ 97 pp. Waterton, D. 1825. Wanderings in South America, the North-West of the United Stales, and the Antilles, in the years 1812, 1816, 1820, and 1824. With original instructions for the perfect preservation of birds, etc. for cabinets of natural history. London, xii+ 326 pp, Wehekind, L. 1956, Notes on some Trinidad bats, J. Trinidad Field Nat. Club, pp. 18-21. 350 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY Wjed-Neuwied, M. zu. 1826. Beitrage z:ur Naturgeschichte von BrasiJien. Weimar, vol. 2, 622 pp. WiLLE, A. 1954, Muscular adaptaiion of ihe nectar-eating bats (subfamily Glossophagi- nae). Trans. Kansas Acad. Sci., 57:315-325. Williams, T. C., J. M. Williams, ani> D. R. Griffin. 1966. The homing abiliiy of the Neotropical bat PfjyUosionnis hasiafus, with evidence for visual orientation. Anim. Behav., 14:468-473. Wilson, D. E. 1971. Food habits of Microvycieris hirsufu (Chiroptera: Phyllosiomidae). Mammalia, 35:107-110. Wimsatt, W. a. 1969. Transient behavior, nocturnal activity patterns, and feeding efficiency of vampire bats (Dex/notius roiti/uius} under natural conditions. J. Mamm., 50:233-244. Wimsatt, W. A., and A. Guekriere. 1962. Observations on the feeding capacities and excretory functions of captive vampire bats. J. Mamm., 43:1 7-27. MOVEMENTS AND BEHAVIOR M, Brock Fenton and Thomas H. Kunz The published information on movements and behavior of phyllostomatids is limited and mainly anecdotal. However, owing to several technological develop¬ ments, notably the availability of image intensifiers and microcircuits, new ad¬ vances are anticipated in these areas. Both these tools have already been used to good advantage (see Schmidt and Grcenhall, 1972; Williams and Williams, 1970) by providing means of studying, with minimal disturbance, the behavior and movements of bats under natural conditions. Concurrently, the successes of several workers (for example, Racey and Kleiman, 1970; Rasweiler and Ishiyama, 1973; Wimsatt et rt/,, 1973) at maintaining various bats in captivity will encourage comparative studies of specific behavior patterns and responses under controlled conditions. We expect that together these developments will produce a series of important studies of the movements and behavior of phyllostomatids in particular and bats in genereal. The results of such studies, w-hen considered in the context of other work (such as the evolutionary and energetic implications of fruit and nectar feeding—^Morton (1973) and Heinrich and Raven (1972), respectively—will permit observations on bats to be placed in a general biological context. The availability of the aforementioned instruments and successes at maintain¬ ing bats in captivity w'ould not be as significant as they are if it w'ere not for the work that has been done on the systemalics, distribution, and natural history of phyliostomalids. Only when such technological developments can be applied in areas where a good basic knowledge of the bats exists do they assume great im¬ portance. Specific areas that come to mind in this context include Trinidad (Gwxl- win and Grcenhall, 1961; Williams and Williams, 1970), Costa Rica and Panama (Brown, 1968; LaVal, 1970; Fleming ei u/., 1972), and various islands in the West Indies (Goodwin, 1970; Jones and Phillips, 1970). Movements Circadian The roosting and feeding habits of bats govern the frequency and magnitude of daily movements between roosts and feeding grounds. Presently, little detailed information is available concerning foraging movements and territories of bats, but phyllostomatids provide one partial exception to this general situation. Using radio tracking, Williams and Williams (1970) documented the feeding flights of Phyltostonws /uistatus from three caves in Trinidad. Upon leaving the caves (the day roosts), bats flew directly to areas where they alternately roosted and made short flights in the feeding area. Feeding areas ranged from one to five kilometers in straight-line distances from the caves, and some individuals travelled four kilometers to reach a feeding area only three kilometers distant. 25i 352 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSJ I’Y I’his Study appears to be the only instance in which the movements of phyllotso* matids between day roosts and feeding grounds have been documented. Data obtained by banding (LaVal, 1970; Fleming e/ a/,, 1972) indicate that some phyllostomatids have regular feeding grounds. Further evidence of this is provided by the observations of Baker (1973) concerning the visits of some glossophagines and one sienodermine to flowers. Li’pioftycferis sanhorti^ G!os- sophaga sp., Ghxsophaga wricina, and Artiheus Janwicensis havQ been observed to make fleeting visits to flowers (Baker, 1973). Baker (1973) and others (Vogel, 1968-69; Baker ef «/., 1971) have remarked on the “trap lining” nature of these visits, w'hich appears to indicate regular patterns of movements. Phyllostomatids, including Macwius waterhousii(Vaugh'dri, 1959), Lonchorina aurita (Nelson, 1965), and Lepionycteris sanborni (Hayw'ard and C’oekrum, 1971), but especially the Phyllonycterinae and the Desmodontinae, a*e active later in the evening than are many other bats (see Silva Taboada and Pine, 1969; Leen and Novick, 1969; Wimsatt, 1969; Crespo a ai, 1972). In low'hind rain¬ forest in Guyana, one of us (Fenton) made similar observations. Using mist nets and ultrasonic detectors (Fenton et a(., 1973), it was established that embal- lonurids, mormoopids, vespertilionids, and molossids w^ere most active around dusk and dawn, W'hereas phyllostomatids (including Phyflostonms ei(mgatu.% Glossophaga soridnay Carollm perspicillatat Rhinophylla pumilio, Sturtura lilium, Uroder/mi biloboftmi, Vampyrops helleri, Vampyressa hidens, Chirodernui villosum, C. (riniiaium, EctophyUa macconmdli^ Anibeus cinereus^ A, ( otictd<7r, A, lituratiis^ Ameirida cetuurio^ and Desmodus rot and us), based on captures in mist nets, w^ere active later in the evening and throughout the night until about one hour before dawm. Further observations on phyllostomatid activity have recently appeared (Heithaus ei a!., 1974; Tuttle, 1974; Davis and Dixon, 1976). In part, these temporal differences can be accounted for by the seqjence of departures from the day roosts. At Mount Plenty Cave in Jamaica, Leen and Novick (1969) observed that MonophyUus redmani was the first species to depart in the evening, followed by PteronomspsUoiis^ P. parneiUi, Artiheus jamakensis, and Phyilonycteris sp. Whether or not these departures represent differential sensitivity to light, roost locations, or differences in circadian periodicity remains to be determined. Captures of bats at different locations during the night have been used to in¬ dicate activity patterns (Brown, 1968; LaVal, 1970). However, comparison of activity patterns from different areas or seasons is difficult because the basic pat¬ terns of activity reflect, among other things, the proximity of the study si :e to day and night roosts. For example, when the activity patterns of Artiheus Jamaicensis in Costa Rica (Fig. la and lb) are compared with those w^e obtained in Puerto Rico for this species (Fig. 1 c), marked differences are evident. Given that the values provid¬ ed by Brown (1968) are absolute numbers and the other values are pro[»ortions, different levels of bat activity occur. Brown (1968) and LaVal (1970) obtained similar patterns of activity of A, jamaicensis in forests and banana groves, and we studied its activity at the entrance to a large cave system, parts of which were used BIOLOGY OF TWE PHYLLOSTOMATJDAE 353 a b FiC. 1.—Nightly activity of Ariiht’iix juniiihetLsiy. A. Costa Rica. Banana Grove (after Brown, 1968; N= 30); B, Costa Rica. Dry Forest (after LaVaL 1970; A'=59); C. Puerto Rico, Cave entrance (This study, A?= 124; a solid circle indicates per cent adult males). as day roosts by this species. The three patterns indicate that some individuals of this species are active throughout the night. Adult male A. Janiaicensis in Puerto Rico were more active one hour after dark and one hour before dawn, but did show sporadic activity throughout the night (Fig, Ic). Williams and Williams (1970) found that much of the activity of PhyUostomus haslatus in Trinidad occurred in the first few hours after sunset, considering the times w'hen most individuals returned to their daytime refuges. They also noted an additional period of activity just betbre sunrise, although LaVal (1969) failed to observe comparable predaw-n activity for other phyllostornatids in Costa Rica. The disparity of these tw'o reports may reflect differences in behavior of bats as a function of proximity to the day rcx)st, since predawn feeding may be restricted to the immediate vicinity of the day roost. The effect of roost proximity and, of course, season and weather on activity patterns of bats makes detailed comparisons from different areas tenuous. Inas¬ much as we lack detailed analyses of activity patterns of bats from any area }54 Sj>t-:C1AL PUBl.JCAT[ONS MUSEU\[ TEXAS rECH UNJVERSITY (with the possible exception of Nyholm's, 1965, data from some species of Myifth), a comprehensive understanding of the situation is presently unrealistic. Similarly, other than anecdotal observations, there are few data on ti e effects of weather on the activity of Phyllostomatidae. Tamsitt and Valdivieso (1961) reported a strong inhibiting effect of moonlight on bat activity in Costa ^ica, but this was not observed by LaVal (1970), who noted that his mist nets had been set in closed forest, w hereas Tamsitt and Valdivieso (1961) had been working in more open situations. Crespo et ai. (1972) found a strong inhibiting effect of moonlight on the activity of Desmodus roiundns. Other studies have documented the effects of moonlight on bat activity (Erkert, 1974; Turner, 1975), which may be related in some areas to the threat of predation (Fenton and Fleming, 1976; Fenton er «/., n.d.). How ever, responses to possible predators is not a uniformly ter able ex¬ planation of the effects of moonlight on the activity of bats. Wimsatt (1969) suggested that heavy precipitation had a supressing effect on foraging activity of /). rotumius, and pointed out the need for detailed work on the effects of local environmental conditions on the activity of bats. Interpretation of nightly activity patterns and comparisons of activity between sympatric taxa also must consider competitive strategies of resource use. Horizon¬ tal and vertical patchiness of habitat (including food and roost sites) probably are important parameters selecting for a reduction in interspecific competition. Vertical stratification of Neotropical bat faunas has been noted by Handley (1967), McNab (1971), and Fenton (1972). For example, among phyllostomaticis, Vam- pyressa hidens and Ardheus iiiunmis were more commonly taken in canopy .sampling than at ground level, whereas the reverse was true of Carollia suhrufa and C perspiciiiata (Handley, 1967). Before reliable temporal comparisons of different species can be made, vertical sampling must be undertaken in a variety of habitats. The sensitivity of bats to disturbance is the main drawback to studies of bat activity that involve capture and marking of animals (either by banding c»r punch marking—Bonaccorso and Smythe, 1972). This is clearly reflected in the band recoveries reported by LaVal (1970) and Fleming et aL (1972), and further ac¬ centuated by our owm experiences in Puerto Rico. Over four nights in May 1973, a total of 314 phyllostomatids was banded at Aguas Buenas Cave in Puerto Rico (168 ArribeiiS jamaicensis, 40 Mottophylliis redmani, 80 Bniaiyphylla cavtrnarum, and 26 Emphyila hombifrons) and during this same period i total of 55 band recoveries was made (14 J per cent of the total banded). Remote sensing systems have been used to monitor the activity of some bats that use high intensity echolocating cries (Fenton et al., 1973). This approach avoids disturbance to the bats, but is not particularly useful for most phyllosto- matids, which are low-intensity ecliolocators. Photocells, photographic apparat¬ uses, and ihermister sensors may provide means of remote monitoring ^f phyl- loslomatid activity and thus permit analysis of the effects of various environ¬ mental parameters on the activity of these bats without introducing biases re¬ sulting from disturbance. The tendency of some bats to use alternate roosts—as reported for Des~ modus rotundus by Wimsatt (1969) and Erophylta sezekomi by Goodwin (1970) BIOLOGY OF TtlE l>HYt.I.OSTOMATIDAH 355 —further complicates the problem of the impact of disturbance on roost-oriented studies (I'urner, 1975). Knowledge of the location of alternate roosts has definite survival value for bats, because it permits them to vacate roosts that are tem¬ porarily or permanently rendered unsuitable in favor of roosts that have not been jeopardized. Seasonal The seasonal movements (or migrations) of bats long have been of interest to biologists (see Allen, 1939), hut most know ledge about them has been obtained in the temperate regions of the northern hemisphere and concerns rhinolophids, vespertilionids, and a few^ ntolossids. (Allen, 1939; Brosset, 1966; l.een and Novick, 1969; Griffin, 1970). Some Ptcropodidae in various parts of their ranges, but particularly in eastern Africa and in Australia, have been shown to migrate, but the Phyllostomatidae are conspicuous by their absence from the roster of migratory bats. Anderson (1969) suspected migration by Macroiifs uaierhonsil^ (= M, californicus, part), and their seasonal absence from the American Southwest led Barbour and Davis (1969) to suggest migration for Lcpuniyctais nivalis, L. sanbonii, and Choeronycterls inexkana. There is now- evidence that some nectari- vorous species (for example, L. sanhorni) return year after year to the same sum¬ mer colony (Hayward and Cockrum, 1971) and that seasonal movements in these species arc probably in response to the Oowering seasons (Leen and Novick, 1969). Davis (1945) reported declines in numbers of CaroUlapcrsplcUkita, Anoura geoffroyU and Dcstnodus romndas from October through December in Brazil. Greenhall (1956) suggested that similar declines reflect shifts of populations in response to exhaustion of local food supplies. Local migration in response to reduced flow'er availability is characteristic of nectar-feeding birds throughout the world (Wolf, 1970; Keast, 1968) and similar movements can be expected to occur in nectar-feeding phyllostomaiids. Why such movements may be more characteristic of flower feeders than frugivorous kinds is in the ephemeral nature of llowers as compared to fruits (Leek, 1972). The use of multiple roosts also may account for local shifts in the distribution of bats. W'imsalt (1969} pointed out that use of alternate roosts presented an adaptive advantage to Desmodus rotundas because of the restricted w'ater budget of vampires. Local population shifts by this species to areas near food resources would concurrently lower evaporative water loss related to movements to and from the roosts, and, for the same reason, reduce levels of food consumption. We suspect that strategies employed by other phyllostomaiids throughout their ranges will involve local, latitudinal, and altitudinal displacements. The absence of marked migrations by phyllostomatids stands in sharp contrast to the situation as it is know n for some pteropodids, which is obviously a function of at least size and habitats. The pteroptxlids for w hich migration is knowm are targe and tend to form conspicuous “camps," which makes them easy to observe. The generally smaller and more secretive phyllostomatids are considerably less conspicuous. 356 SPhX’lAL PUHLJ CAT JONS MUSEUM TEXAS TECH UNJVERSJTV Perhaps more significant, however, than size and roosting habits, are the differ¬ ences in climate betw'een South and Central America and Africa and Australia. Keast (1969) provided a convenient comparison of these three areas: whereas 32 percent of South and Central America is rainforest, this habitat .recounts for 10 per cent and 5 per cent, respectively, of the area of Africa and Australia. Habitats in which marked seasonal nucluations occur (with resultant seasonally available food sources) are more conducive to the evolution of migratory pat¬ terns than are habitats with less drastic fluctuations. Climatic actual ions also may account for the higher diversity of fruit and nectar-feeding bats in the Neotropics (relative to the Old World tropi:s). The larger size of the Ptcropodidae (relative to the Phyllosiomatidae) may reflect migratory habits because movement over long distances is proportionally less costly (energetically) for larger as opposed to smaller organisms (Schmidt-Nielson, 1972; Thomas and Suthers, 1972; Thomas, 1975). Honline Griffin’s (1970) review of studies of homing by bats included one pfiyllosto- matid. Williams ei aL (1966) and Williams and Williams (1967, 1970) used radio tracking to examine homing by PhyUostontns hastmuK and showed that bats dis¬ placed more than 30 kilometers from their homes were less effective at returning there than those displaced shorter distances. These studies also demonstrated the importance of visual cues to homing in hasmtm. Banding studies have indicated homing by Maemms caUfornk us (Bradshaw-, 1961; Davis, 1966) and Lef)!ouyctens sunhortu (Hayward and Cockrum, 1971). The whole question of homing in bats was succinctly addressed by Wilson and Findley (1972) w ho, after examining the available evidence, including the afore¬ mentioned studies of Williams and colleagues, concluded that no one had demon¬ strated other than random movements by displaced bats. W'e concur v ith this opinion and w-ith the importance of having information concerning the faniiiarity of bats with the area involved (for F, hasiams, up to 20 kilometers from lome— Williams and Williams, 1970). riie size of the familiar area is greatly influenced by the roosting habits of the bats involved and, as indicated by Fleming et aL (1972), by the size of the bat. Future studies involving displacements of bats from their home roosts probably w'ill demonstrate that larger bats and bats that form large colonies will have pro¬ portionally larger spatial areas of familiarity than small or solitary bats. Migra¬ tory species such as Leptonyciei is nivads^ L. sanborui, Choenmyvteris mt’xkana^ and MuiTotiiS valifomk us will have a greater degree of spatial familiar ty than sedentary species of the same size. Using rates of recovery of marked individuals, LaVal (1970) suggested that Phyilostomus (iiscnloi\ CaroKia brevicauda^ and G. castanea (for which he ob¬ tained high recovery rates) may have smaller home ranges than species for which he had low recovery rates, such as Arlibiia Jamaicensis, Glossophaga lammis- sarisi, and Urocierma hiLfhatuni Because body size or colony size (or both) gen¬ erally reHect resource requirements and distribution of resources, it is clear that HlOi.OGY Oi- I HE PHYLLOSTOMATIDAE 3.^7 ihe local and geographic differences in areas of familiarity will in part be a func¬ tion of resource distribution and density. Present agricultural practices and high cattle densities in some areas of the Neotropics may select against a large familiar area for bats using such artificial concentrations of food resources (for example, Diwnodits rot n ml us). Behavior Sen Si lt y The eyes of phyllosioniatids probably serve regular complex visual functions (Chase and Suthers, 1969), such as surveillance for predators (Suthers, 1970), distance orientation (Williams ei 1966), and the location of feeding areas (Williams and Williams, 1970). Suthers (1970) postulated that passive visual surveillance by a resting bat may function to permit it to select visually important events before making a more detailed acoustical investigation. The relative im- potance of visual as opposed to acoustical information in the responses of phyl- lostomatids is not well understood, but probably depends upon light conditions (as it does for Ronseitns sp.) and the general circumstances (Manske and Schmidt, 1976). The importance of vision in surv-eillance for predators also is suggested by some anatomical features such as the transparent dactylopatagium minus of some phyllostomatids (Vaughan, 1970), The hypothesis that vision is important in orientation and feeding i.s supported from experiments conducted by Williams and colleagues (Williams et ai, 1966; Williams and Williams, 1967, 1970) and from theoretical constraints relating to the relatively short etTective range of echolocation (Griffin, 1958, 1971; Suthers, 1970; Fenton, 1974). Well-developed vomeronasal organs and associated olfactory bulbs as reported by Schneider (1957), Mann (1961), and Suthers (1970) and anecdotal observations indicate well-developed olfactory senses in the Phyllostomatidae. P/}yl!(?stonttiS hasiains can locate fruit hidden from view (Mann, 1961) and the sniffing behavior of Dcstnodus rotutulus before licking and biting prey (Green- hall, 1972; Schmidt, 1973) points to the importance of olfaction. The acute odor discrimination showm by D. roinmhis probably permits it to detect dif¬ ferences betw'een breeds of cattle (Schmidt, 1973). Olfaction may be equally im¬ portant for nectar and pollen-feeding bats; Baker (1973) noted that one of the characteristics of flow-ers visited by bats is a sour smell. Recent comparisons of the olfactory systems of some phyllostomatids with those of other bats (Bhatnagar, 1975; Bhatnagar and Kallen, 1974, 1975) further emphasizes the importance of odor in the lives of bats. I tu raspedfic Phyllostomatids show- a variety of roosting habits with respect to numbers of individuals occupying a roost. Estimates of colony size vary considerably and have usually been based on visual counts during emergence or directly in roosts under low' light levels (usually after the bats have been disturbed). Some phyllosto- SF>ECIAJ, EUBIJCATIONS MUSEUM T BXAS TECH UNJVERSfTY mat ids appear to roosl alone or in small groups (for example, Mivnmycieris mega- iofis^ M. niinuta, M. hlrsuta, M. hrachycnis, Lonchonna au/iia^ ['(ffuiiia syivicaia, Tonafia hide/is, PhyllasromHS elo/igafus, and A/tiheus phoeoiis — Goodwin and Grcenhall, 1961; Leen and Novick, 1969; Tuttle, 1970; Gttodwin, 1970). Others are sometimes found in small aggregations or on other occasions in large colonies (for example, Caroilia pe/spivillala —Pine, 1972; Phylk skf/iufs hasktiifs —Williams and Williams, 1970; Artihetisja/iiaieensis —Leen and Novick, 1969), whereas still others appear to occur only in large aggregations (such as li/mhyphylla cme/iiartini in Puerto Rico). The size of the roost may C'cert an important limiting factor on the size of the colony, as is indicated by the occur¬ rence of larger aggregations of individuals of some species in artificial str jctures than are knowvn from natural roosts {Desmodus /rfimidus, for example). .Species that regularly roost in large rooms in caves probably are more commonly en¬ countered in large aggregations than are those that roost in cavities of trees Aside from observations on colony or cluster size, little has been published on intraspecific behavior of bats in colonies. Some evidence is available indicating that there are social units of groups within colonies and that these may play important roles in reproduction, food gathering, and orientation. It seems logical to expect more elaborate social interactions in gregarious than in solitary species (as in some Canidae—Kleiman, 1972), Williams and W'illiams (1970) reported “coherent social groups” foi Phyl- losto/nus hasiaiNs ranging from five to 20 individuals and consisting of groups of both sexes with one or more dominant males. Bradbury (n.d.) has provided more information on the social groups of P/tyUimoi/iits has/atus and P. disi olo/\ Piiyik/siofuns /uiskinis Ibrms large colonies in caves and the population in any roost site consists of harems (25 to 30 females per male) and nonharem juveniles and males. Harem males protect their females and perform elaborate displays when another male approaches. To feed, females leave the harem singly and in twos, whereas the male departs when the number of remaining females is at its lowest, and remains away for only a short time. Removal of a harem male results in his replacement by another male with little or no turnover among the harem females, Phyilosto/nifs discolor establishes colonies in hollow trees and again the pop¬ ulations include harem (one to 12 females per male) and nonharem bats. Ho'vevcr, harem composition in this species is more variable than in P. hasiams, wath some females being regularly present in the harem and others somcw'hat nomadic. Female P. discolor are more aggressive than female P. luiSkiius and arc actively involved in maintaining the integrity of the harem. A bat returning to a aarem group performs elaborate displays, which include tactile, olfactory, and vocal cues, to gain admission to the group. A1 logrooming by members of harems is com¬ mon. In both species, the non harem groups may be quite stable in their composition and tend to be more nomadic than the harems. F. Potter (personal communica¬ tion) has observed harem structures in Caroilia perspicillaia and it seems likely that this situation may be common in phyllostomatids that aggregate in large numbers. B[OLOGY OF THE PHYl.lTXSTOMATIDAF 359 Departures of groups of bats from roosts (for example, Lepiomcicvis san- hvmi —Hayward and Cockrum, 1971; Desmodus rotundas —Wimsatt, 1969, and Greenhall et ai, 1971) also suggest the presence of social groupings. Similar observations have been reported for other bats (Rhinolophidae-—Mohres, 1967; Vespertilionidac^—-Hall and Brenner, 1968, and Dwyer, 1970) and indicate that this behavior may be w'idespread in the Chiroptera, Segregation of females into discrete groups prior to parturition and until the young are w'eaned has been reported for Anihvas jamnk ensis (Lecn and Novick, 1969) and implied by the observations of Jones es cil. (1973) for De.snioda.s romndas. The observations of been and Novick (1969) for A. janadeensis and those of Schmidt (1973) for IX rotundas indicate that olfactory cues may be im¬ portant in social organization and mother-young relationships. How'ever, other information, some of it from phyllostomatids, suggests that vocalizations are im¬ portant in interactions between females and their young (Brown, 1976; Gould. 1975r;, i975/j; Gould ft cd., 1973; Schmidt and Manske, 1973) and in a variety of other intraspecific contexts (for example, Bradbury and Emmons, 1974; Wickler and Siebt, 1976). Evidence from other mammals strongly suggests that bats w ill be show'n to exhibit various patterns of social dominance within groups. Places where these interactions may be expected are roosts and common feeding sites. Laboratory observations of Schmidt and Greenhall (1972) on the interactions of feeding vampires support this suggestion; they suggested that certain '‘dominant” animals in a group feed first and that, while they are feeding, they chase off other Individuals as in some carnivores (Ewer, 1973). Similar interactions will certainly be reported from situations where food rescnirces are localized (for example, concentrations of ripe fruit). However, species that are nectarivorus or carnivorous (including insectivorous) where food resources are diffuse are more apt to demonstrate ter¬ ritorial interactions than dominance hierarchies at feeding sites (see, for example, Baker*s, 1973, observations of trap-lining in some nectar-feeding bats). The w'hole subject of sexual behavior and the details of mother-young inter¬ actions are poorly known, and we were unable to find any published information on this subject for phyllostomatids. Itacrspecifii Various species of bats are know n to share the same roosts, but in some cases, use of a roost by one species w ill result in its being abandoned by another species (for example, Artiinnis Janiaiccasis and Midossas up ,—Lecn and Novick, 1969). Often several species of bats may roost in one structure (tree, cave, building, and so forth) and not come into physical contact with one another except possibly at times of arrival or departure. There is little information on interspecific behavior of bats, although biting is presumed to occur and possibly be involved with the epidemiology of rabies virus (Constantine, 1970). As w'ith intraspecific in¬ teractions, it is likely that interspecific altercations will occur where food resources or roosts arc localized or limited. The work of Colwell (1973) on the interactions of some hummingbirds suggests that similar interspecific behavior patterns may 360 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY be described for nectar-feeding bats, especially in the light of the trap-lining nature of the visits of some of these bats to flowers. Miscellaneous Several species of bats are known to carry their young wdth them away from their roosts. Tamsitt and Valdivicso (1965) observed this for Anlheus hturanis in Colombia, but Fenton {1969) found no evidence of it tor A. Jafnakrnsis in Puerto Rico. In a review' of the literature on this subject, Davis (1970) reported that some phyllostomatids had been found to transport their young after dis¬ turbances in their roosts {Matrotus ccilifornicus, Choeronycfens mexicana, lA’pionyaeris sanhorni)^ whereas others would do this even in the absence of disturbance {Glossophaga soricina and Li‘pionycteris smibornf), H is likely that species that use alternate roosts will be found to transport their young more reg¬ ularly than those that do not, but certainly the presence of disturbance is an im¬ portant consideration in this regard. Tuttle (1970) reported that the vocalizations of a captured Mimon cren,fiaruni attracted other individuals of the same species to the site of capture, and similar effects were elicited by the "distress calls” of several species of bats. At present we have no definite information as to which frequencies of bat cries are important in "distress” or other calls that evoke responses from other individuals. Recent work has indicated that some bats respond preferentially to "distress” calls of conspecifics (Fenton ej aL, 1976). Further Research From the preceeding discussion, it should be obvious that almost any aspect of the movements and behavior of phyllostomatids will provide productive topics for research. Documentation of the daily and seasonal patterns of the activity of these bats with respect to various environmental parameters such as meteorological conditions, lunar cycles, and seasonal changes in the abundances of food (from insects to fruit) should be a primary goal. At the same time, the whole spectrum of intra and interspecific behavior patterns {territoriality, partitioning o( food and roost resources, reproductive, and mother-young behavior) requires close attention. As we have pointed out, many of these subjects may now' be addressee with the aid of electronic equipment (notably for telemetry and observation at low light levels) and a reasonable knowledge of the phyllostomatids that occur in different areas. This situation is reflected by a variety of recent studies ranging from roosting behavior (Timm and Mortimer, 1976), through feeding and ori¬ entation behavior (Howell 1974u, 19746; Fleming et ai, n.d.), to detailed studies of specific bats (Turner, 1975). The programs of recent North American Symposia on Bat Research indicate that work on some of these subjects is in progress for some species. We expect that the next few' years will see the publication of results that will gi eatly advance our knowledge of the movements and behavior of bats in general and the Phyllosiomatidae in particular. BIOLOGY OF THE PHYLLOSTOMATIDAE 361 I.ITERATL RE Cl TED Al l HN, G- M. 1939. Bats. Harvard Univ. Press. Cambridge, 36H pp. Ani>i--kson, S. 1969. 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Functional morphology of three bats; Eumops, Myolis, Macrotus, Univ. Kansas PubL, Mus. Nat. Hist., 12:1-153. -—. 1970. The transparent dactylopatagium minus in phylloslomatid bats. J, Mamm., 51:142-145. VooEL. S. 1968-69, Chiropterophilie in der neotroposchen Flora. Neue Milteilungen 1. [Land HI. Flora, Abi. B, 157:562-602, 158:185-222, 158:289-323. WiCRi ER, W.. AND U, SEtBT. 1976, Field studies on the African fruit bat, Epo/nophomx iviihihetpi (Sundevall), with special reference to male calling. Z. Tierpiychol., 40:345-376. W'lLi.iAMS. T. C.. AND J. M. Wtn lAMS. 1967. Radio tracking of homing bats. Science, 155:1435-1436. -. 1970. Radio tracking of homing and feeding flights of a neotropical bat. Anim. Behav., 18:302-309. Wiiuams, T. C. j. M. Wii-LiA.MS, AND D. R. Griffin. 1966. The homing ability of the Neotropical bat Phy}l{}si(}tni{x hasktHf.y with evidence for visual orientation. Anim. Behav., 14:468-473. Wilson, D, E., and J, .S. Findley. 1972. Randomness in bat homing. Amer. Nat., 106: 418-424, WiMSATT, W. A. 1969. Transient behavior, nocturnal activity patterns, and ceding efficiency of vampire bats {DesnuHins rornminx) under natural conditions. J, Mamm., 50:233-244. WiMSATi, W. A., ,A. Guerriere, and R. Horst. 1973, An improved cage design for maintaining vampires (Dtsinodns} and other huts for experimental purposes. J. Mamm., 54:251-254. Wolf, L. L. 1970. The impact of seasonal flowering on the biology of some fopical hummingbirds. Condor, 72; 1-14. Copies of the following numbers of Special Publications of The Museum may be obtained on an exchange basis from, or purchased through, the Exchange Librarian, Texas Tech University, Lubbock, Texas 79409. No. 9 No. 10 Watkins, L. C., J. K. Jones, Jr., and H. H. Genoways. 1972. Bats of Jalisco. Mexi¬ co, 44 pp., 3 figs. . . , ... $1.00 Krishtalka, L. 1973. Late Paleocene mammals from the Cypress Hills, Alberta, 77 pp., 21 figs. $2.00 West, R. M. 1973. Review of the North American Eocene and Oligocene Apatemy- idae (Mammalia: Insectivora). 42 pp.. 20 figs.$1.00 Gardner, A. L. 1973. The systematics of the genus Didetphis (Marsupialia: Didel- phidae) in North and Middle America, 81 pp., 14 figs.$2.00 Genoways, H. H, 1973. Systematics and evolutionary relationships of spiny pocket mice, genus Liomys, 368 pp.. 66 figs.$7.00 Northington, D. K. 1974. Systematic studies of the genus Pyrrhopappus (Com- posiiae, Cichorieae). 38 pp.. 14 figs...$1.00 King, M, E., and I. R. Traylor, Jr., eds. 1974. Art and environment in native America, 169 pp. $5.00 Pence. D. B. 1975. Keys, species and host list, and bibliography for nasal miles of North American birds (Acarina: Rhinonyssinae, Turbinoptinae, Speleognathi- nae, and Cytoditidae), 148 pp„ 728 figs. . , ....$4.00 Bowles, J- B. 1975. Distribution and biogeography of mammals of Iowa, 184 pp., 62 figs.$5.00 Baker, R. J.. J. K. Jones, Jr., and D. C, Carter, eds. 1976. Biology of bat-s of the New World family Phyllostomatidae. Part L, 218 pp.$6.00 Foster, D. E. 1976. Revision of North American Trichodes (Herbst) (Coleoptera: Cleridae), 86 pp.. 17 figs. .... $2.00 Mitchell, R. W., W. H. Russell, and W. R, Elliott. 1977, Mexican eyeless chara- cin fishes, genus Aslyanax: environment, distribution, and evolution, 89 pp,. 21 figs... $5.00 No. 13 Baker, R. J., J. K, Jones, Jr., and D. C. Carter, eds. 1977, Biology of bats of the New World family Phyllostomatidae. Part II., 364 pp.. $16,00