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SPECIAL  PUBLICATIONS 
THE  MUSEUM 

TEXAS  TECH  UNIVERSITY 


^  s^AC'i^  JcaKK 

T e  ;  si.c-v  , 

MUS.  COMP.  ZOOL. 

I.J  ~IV>  <\  ^Y 


JAN  1  9 1979 

university 


Biology  of  Bats  of  the  New  World  Family 
Phyllostomatidae.  Part  III 


Edited  by 

Robert  J.  Baker ,  J.  Knox  Jones,  Jr.,  and  Dilford  C.  Carter 


January  1979 


TEXAS  TECH  UNIVERSITY 


Cecil  Mackey,  President 


Regents. — Robert  L.  Pfluger  (Chairman),  J.  Fred  Bucy,  Jr.,  Clint  Formby,  Roy  K.  Furr, 
A.  J.  Kemp,  Jr.,  James  L.  Snyder,  Lee  Stafford,  Judson  F.  Williams,  and  Don  R.  Workman. 

Academic  Publications  Policy  Committee. — J.  Knox  Jones,  Jr.  (Chairperson),  Dilford  C. 
Carter  (Executive  Director  and  Managing  Editor),  Robert  J.  Baker,  David  K.  Davies,  Harold 
E.  Dregne,  Leslie  C.  Drew,  Charles  S.  Hardwick,  Ray  C.  Janeway,  Walter  R.  McDonald, 
George  F.  Meenaghan,  Charles  W.  Sargent,  and  J.  Dalton  Tarwater. 


The  Museum 

Special  Publications  No.  16 
441  pp. 

12  January  1979 

$20.00 


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. 


ISSN  0149-1768 
ISBN  0-89672-068-3 


Texas  Tech  Press,  Lubbock,  Texas 


1979 


SPECIAL  PUBLICATIONS 
THE  MUSEUM 
TEXAS  TECH  UNIVERSITY 


Biology  of  Bats  of  the  New  World  Family 
Phyllostomatidae.  Part  III 


Edited  by 

Robert  J.  Baker,  J.  Knox  Jones,  Jr.,  and  Dilford  C.  Carter 


No.  16 


January  1979 


TEXAS  TECH  UNIVERSITY 


Cecil  Mackey,  President 


Regents. — Robert  L.  Pfluger  (Chairman),  J.  Fred  Bucy,  Jr.,  Clint  Formby,  Roy  K.  Furr, 
A.  J.  Kemp,  Jr.,  James  L.  Snyder,  Lee  Stafford,  Judson  F.  Williams,  and  Don  R.  Workman. 

Academic  Publications  Policy  Committee. — J.  Knox  Jones,  Jr.  (Chairperson),  Dilford  C. 
Carter  (Executive  Director  and  Managing  Editor),  Robert  J.  Baker,  David  K.  Davies,  Harold 
E.  Dregne,  Leslie  C.  Drew,  Charles  S.  Hardwick,  Ray  C.  Janeway,  Walter  R.  McDonald, 
George  F.  Meenaghan,  Charles  W.  Sargent,  and  J.  Dalton  Tarwater. 


The  Museum 

Special  Publications  No.  16 
441  pp. 

12  January  1979 
$20.00 


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. 


ISSN  0149-1768 
ISBN  0-89672-068-3 


Texas  Tech  Press,  Lubbock,  Texas 


1979 


CONTENTS 


Introduction  .  5 

Systematic  and  Distributional  Notes .  7 

J.  Knox  Jones,  Jr.,  and  Dilford  C.  Carter,  The  Museum,  Texas  Tech 
University,  Lubbock,  79409. 

Morphometrics  .  13 

Pierre  Swanepoel  and  Hugh  H.  Genoways,  Kaffrarian  Museum,  King 


William’s  Town,  5600,  Republic  of  South  Africa;  Carnegie  Museum  of 
Natural  History,  4400  Forbes  Avenue,  Pittsburgh,  Pennsylvania 
15213. 


Karyology  .  107 

Robert  J.  Baker,  Department  of  Biological  Sciences  and  The  Museum, 
Texas  Tech  University,  Lubbock,  79409. 

Biochemical  Genetics  . .  157 


Donald  O.  Straney,  Michael  H.  Smith,  Ira  F.  Greenbaum,  and  Robert  J. 
Baker,  Museum  of  Vertebrate  Zoology,  University  of  California,  Berkeley, 
94720;  Savannah  River  Ecology  Laboratory,  Aiken,  South  Carolina 
29801;  Department  of  Biology,  Texas  A&M  University,  College 
Station,  77843;  Department  of  Biological  Sciences  and  The  Museum, 

Texas  Tech  University,  Lubbock,  79409. 

Sperm  Morphology .  177 

G.  Lawrence  Forman  and  Hugh  H.  Genoways,  Department  of  Biology, 
Rockford  College,  Rockford,  Illinois  61101;  Carnegie  Museum  of 
Natural  History,  4400  Forbes  Avenue,  Pittsburgh,  Pennsylvania  15213. 

Alimentary  Tract . 205 

G.  Lawrence  Forman,  Carleton  J.  Phillips,  and  C.  Stanley  Rouk,  De¬ 
partment  of  Biology,  Rockford  College,  Rockford,  Illinois  61101;  De¬ 
partment  of  Biology,  Hofstra  University,  Hempstead,  New  York 
1 1550;  Barton  County  Community  College,  Great  Bend,  Kansas  67530. 

Morphometric  Analysis  of  Chiropteran  Wings .  229 

James  Dale  Smith  and  Andrew  Starrett,  Department  of  Biological 
Sciences,  California  State  University,  Fullerton,  92634;  Department  of 
Biological  Sciences,  California  State  University,  Northridge,  91330. 


Reproductive  Patterns  .  317 

Don  E.  Wilson ,  U.S.  Fish  and  Wildlife  Service,  National  Fish  and 
Wildlife  Laboratory,  National  Museum  of  Natural  History, 

Washington,  D.C.  20560. 

Embryology .  379 

William  J.  Bleier,  Department  of  Zoology,  North  Dakota  State 
University,  Fargo,  58102. 

Ontogeny  and  Maternal  Care  .  387 

D.  G.  Kleiman  and  T.  M.  Davis,  National  Zoological  Park, 

Smithsonian  Institution,  Washington,  D.C.  20008. 

General  Physiology  .  403 

John  M.  Burns,  Department  of  Biological  Sciences,  Texas  Tech 
University,  Lubbock,  79409. 

Population  and  Community  Ecology .  409 

Stephen  R.  Humphrey  and  Frank  J.  Bonaccorso,  The  Florida  State 
Museum,  University  of  Florida,  Gainesville,  3261 1;  University 
College,  European  Division,  University  of  Maryland,  im  Bosseldorn 
30,  6900  Heidelberg,  German  Federal  Republic. 


INTRODUCTION 


Because  of  their  adaptive  diversity  and,  in  many  instances,  unique  morphologi¬ 
cal  attributes,  bats  of  the  family  Phyllostomatidae  long  have  fascinated  biologists. 
Known  only  from  the  New  World,  most  species  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  accumulated,  and  it  is  the  purpose  of  this  three-part  pub¬ 
lication,  which  contains  a  total  of  27  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  future  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  volumes,  each  separately  numbered  as  a  Special  Publication  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  trilogy) 
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  from  one  volume  to  the  next  without  the  handicap  of 
conflicting  names  for  the  same  organism. 

Manuscripts  first  were  requested  from  contributors  in  1973  and  most  had 
been  received  by  the  end  of  1974.  Part  I  of  the  series  was  published  in  1976  and 
Part  II  in  1977.  As  editorial  work  progressed,  some  authors  provided  up-dated 
information  and  all  authors  had  the  opportunity  to  insert  limited  materials  at 
the  time  they  received  galley  proofs.  Therefore,  content  is  as  current  as  reasonably 
could  be  anticipated  for  a  project  of  this  kind.  Organization  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  redundancy  has  resulted,  we  thought  it  best  to  have  a  section 
on  the  cited  literature  with  each  contribution.  Citations  to  manuscripts  in  Part 
III  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 


6 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


pp.  7-38;  Zoogeography  (Koopman),  pp.  39-47;  Chiropteran  evolution  (Smith), 
pp.  49-69;  Collecting  techniques  (Tuttle),  pp.  71-88;  Care  in  captivity  (Green- 
hall),  pp.  89-131;  Economics  and  conservation  (C.  Jones),  pp.  133-145;  Brain 
anatomy  (McDaniel),  pp.  147-200;  and  Lactation  and  milk  (Jenness  and 
Studier),  pp.  201-218. 

Following  a  two-page  introduction  by  the  editors,  Part  II  (Spec.  Publ.  Mus., 
Texas  Tech  Univ.,  13:1-364,  1977)  includes:  Endoparasites  (Ubelaker, 

Specian,  and  Duszynski),  pp.  7-56;  Ectoparasites  (Webb  and  Loomis),  pp. 
57-1 19;  Oral  biology  (Phillips,  Grimes,  and  Forman),  pp.  121-246;  Echolocation 
and  communication  (Gould),  pp.  247-279;  Thermoregulation  (McManus), 
pp.  281-292;  Feeding  habits  (Gardner),  pp.  293-350;  and  Movements  and 
behavior  (Fenton  and  Kunz),  pp.  351-364. 


February  1978 


Robert  J.  Baker 
J.  Knox  Jones,  Jr. 
Dilford  C.  Carter 


SYSTEMATIC  AND  DISTRIBUTIONAL  NOTES 


J.  Knox  Jones,  Jr.,  and  Dilford  C.  Carter 


Since  completion  of  the  manuscript  for  an  annotated  checklist  of  phyllostomatid 
bats,  which  appeared  in  the  first  part  of  this  trilogy  (Jones  and  Carter,  1976), 
several  publications  have  come  to  our  attention  that  alter  the  systematic  arrange¬ 
ment  originally  presented  or  extend  the  known  distribution  of  included  species. 
These  papers  are  summarized  here  for  the  convenience  of  those  who  may  not  have 
all  the  recent  literature  available  to  them  and  also  in  order  to  make  the  three- 
volume  set  on  the  biology  of  the  Phyllostomatidae  more  useful  as  a  source  of 
references.  Some  of  this  new  information  also  is  incorporated  in  an  annotated 
checklist  of  the  bats  of  Mexico  and  Central  America  by  Jones  et  al.  (1977). 

Systematics 

In  a  recent  appraisal  of  the  taxonomy  and  zoogeography  of  Macrotus  water- 
housii  in  the  West  Indies,  Buden  (1975)  reached  the  conclusion  that  only  two 
subspecies  should  be  recognized  there:  waterhousii  (jamaicensis  a  synonym) 
on  Jamaica,  Hispaniola,  and  Puerto  Rico,  and  in  the  southern  Bahamas;  minor 
( compressus  a  synonym)  on  Cuba,  Grand  Cayman,  and  in  the  northern  Bahamas. 
Anderson  and  Nelson  (1965)  had  recognized  four  subspecies  in  the  Antillean 
segment  of  the  distribution  of  M.  waterhousii. 

Greenbaum  et  al.  (1975)  convincingly  argued,  on  the  basis  of  karyotypes, 
that  Mesophylla  is  generically  distinct  from  Ectophylla,  a  conclusion  earlier 
reached  on  the  basis  of  morphologic  comparisons  by  Starrett  and  Casebeer 
(1968). 

We  earlier  listed  the  subgenus  Xenoctenes  to  include  Micronycteris  hirsuta. 
Davis  (1976)  provided  evidence  for  abandoning  Xenoctenes  as  valid  and 
returned  M.  hirsuta  to  the  nominate  subgenus. 

Distributional  records  listed  for  Peru  by  Gardner  (1976)  were  taken  into 
account  in  preparation  of  our  checklist,  but  the  publication  arrived  too  late  to 
insert  remarks  relating  to  systematics.  Among  these,  Gardner  suggested  that  all 
species  of  small  Tonatia  ( hrasiliensis ,  venezuelae,  and  minuta )  probably  are 
conspecific  and  that  Lichonycteris  degener  may  be  synonymous  with  L.  obscura. 
He  also  questioned  the  report  of  Lonchophylla  concava  from  Peru. 

Buden  (1976)  studied  the  genus  Erophylla  systematically  and  reduced 
the  then-recognized  two  species,  including  a  total  of  six  subspecies,  to  two  sub¬ 
species  of  a  single  species,  E.  sezekorni,  as  follows:  sezekorni  ( mariguanensis , 
planifrons,  and  syops  synonyms)  from  the  Bahamas,  Cuba,  Jamaica,  and  the 
Cayman  Islands;  bombifrons  ( santacristobalensis  a  synonym)  from  Hispaniola 
and  Puerto  Rico. 

Buden  (1977)  also  reviewed  morphological  variation  in  Brachyphylla  and 
concluded  that  all  extant  populations  should  be  referred  to  the  one  species  B. 


7 


8 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


cavernarum.  Subspecies  recognized  by  Buden  were:  cavernarum  (Puerto  Rico, 
Virgin  Islands,  Lesser  Antilles  south  to  St.  Vincent);  minor  (Barbados);  nana 
(Cuba  and  Grand  Cayman);  and  pumila  (Hispaniola  and  the  Caicos  Islands  in 
the  southern  Bahamas).  Verona  (1974)  earlier  arranged  all  named  taxa  of 
Brachyphylla  as  subspecies  of  the  single  species  cavernarum,  but  gave  no  reasons 
for  having  done  so. 

In  a  paper  on  activity  patterns  of  bats  taken  near  Iquitos,  Peru,  Davis  and 
Dixon  (1976)  used  the  names  “Artibeus  planirostris ”  and  “  Artibeus  fuliginosus ,” 
evidently  based  at  least  in  part  on  information  contained  in  the  unpublished 
doctoral  dissertation  of  Donald  R.  Patten.  They  also  listed  Artibeus  pumilio 
as  a  distinct  species;  we  referred  to  pumilio  as  a  subspecies  of  A.  cinereus. 
Similarly,  Smith  and  Genoways  (1974)  used  the  name  combination  “ Artibeus 
planirostris  trinitatis ”  in  reference  to  a  population  on  Margarita  Island, 
Venezuela.  They  cited  Patten’s  unpublished  dissertation  as  the  basis  for  recognition 
of  specific  status  for  planirostris  (which  we  listed  as  a  subspecies  of  jamaicensis). 
We  have  read  Patten’s  dissertation  and  do  not  believe  he  intended  to  apply  the 
specific  name  planirostris  to  jamaicensis- like  bats  from  the  Caribbean  coastal 
area  of  northern  South  America  and  adjacent  islands;  nevertheless,  we  deplore 
the  use  of  manuscript  names  and  strongly  suggest  that  such  information  not  be 
incorporated  into  the  published  literature  without  appropriate  documentation. 

Handley  (1976)  provided  a  valuable  annotated  checklist  of  Venezuelan 
bats  in  which  there  are  several  departures  from  the  systematic  scheme  we 
employed.  Unfortunately,  none  of  these  departures  is  documented  with  evidence 
or  other  explanation;  rather,  it  is  indicated  that  the  author  will  describe  new 
taxa  and  discuss  nomenclatural  changes  in  another  paper  that  was  “in  press” 
but  which,  to  our  knowledge,  has  not  yet  appeared. 

Finally,  Jones  (1978)  described  a  new  subspecies  of  the  Artibeus  jamaicensis 
complex  from  the  Antillean  island  of  St.  Vincent  ( schwartzi ),  and  Davis  and 
Carter  (1978)  named  as  new  Tonatia  evotis,  which  occupies  a  distribution  from 
Chiapas  southeastward  in  the  Caribbean  versant  of  Central  America  to 
Honduras  within  the  range  earlier  ascribed  to  T.  silvicola  (note  change  in 
spelling).  They  also  described  a  new  subspecies  of  the  latter  (T.  s.  centralis ) 
from  Honduras,  Nicaragua,  and  Costa  Rica,  and  a  second  new  subspecies 
( T.  s.  occidentalis )  from  western  Ecuador  and  Pern,  while  restricting  the  dis¬ 
tribution  of  the  nominate  subspecies  to  the  region  from  Panama  into  South 
America  as  far  as  Amazonian  Brazil,  Bolivia,  and  Peru. 

[Koopman’s  (1978)  important  contribution  on  systematics  and  zoogeography 
of  Peruvian  bats  was  received  after  our  report  was  in  galley  proof.  It  contains 
accounts  for  71  species  of  phyllostomatids.  Among  the  important  systematic 
comments  are  the  following:  Mimon  koepckeae  was  regarded  as  a  subspecies  of 
M.  crenulatum ;  Choeroniscus  inca  was  synonomized  with  C.  minor,  Vampyrops 
nigellus  was  placed  as  a  subspecies  of  V.  lineatus\  Enchisthenes  was  reduced  to 
subgeneric  status  under  Artibeus,  as  has  been  done  by  several  other  authors; 
Artibeus  glaucus  and  A.  watsoni  were  regarded  as  conspecific  with  A. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


9 


cinereus,  but  A.  anderseni  was  recognized  as  a  distinct  species;  Diaemus  was 
considered  congeneric  with  Desmodus.  Additionally,  Koopman  recognized  and 
defined  the  species  Artibeus  fraterculus,  A.  fuliginosus,  and  A.  planirostris  as 
distinct  from  A.  jamaicensis — we  listed  fraterculus  and  planirostris  as  subspecies 
of  A.  jamaicensis ,  and  fuliginosus  represents  the  “underscribed  species”  men¬ 
tioned  in  the  same  account.  ] 

[After  this  paper  was  in  paged  proof,  we  became  aware  of  a  review  of  the 
genus  Lonchorina  by  Hernandez-Camacho  and  Cadena-G.  (Caldesia,  13:1 99-25 1 , 
1978),  which  included  description  of  a  new  species,  Lonchorhina  marinkellei 
(p.  229),  with  type  locality  at  Durania,  near  Mitu,  Colombia.] 

Faunistics 

Starrett  (1976)  and  LaVal  (1977)  recorded  species  of  bats,  including 
phyllostomatids,  new  to  the  fauna  of  Costa  Rica.  The  latter  paper  contains  the 
first  reported  specimen  of  Micronycteris  daviesi  from  North  America  under  the 
generic  (instead  of  subgeneric)  designation  Barticonycteris.  Koopman  (1975) 
summarized  the  bat  fauna  of  the  Virgin  Islands  and  its  zoogeographic  relation¬ 
ships.  In  a  report  on  bats  from  southern  Haiti,  Klingener  et  al.  (1978)  recorded 
the  first  whole  specimens  of  Phyllonycteris  poeyi  obtusa,  previously  known  only 
from  skeletal  remains. 

Greenbaum  and  Jones  (1978)  reported  new  records  of  phyllostomatids 
from  several  Middle  American  countries  and  Carter  and  Jones  (1978)  recorded 
several  new  species  for  the  Mexican  state  of  Hidalgo,  including  the  northeastern- 
most  record  of  Chiroderma  villosum.  Furthermore,  Baker  and  Genoways 
(1978)  summarized  in  a  useful  way  the  zoogeography  of  Antillean  bats,  and 
Baker  et  al.  (1978)  reported  on  bats  from  the  island  of  Guadeloupe. 

In  our  checklist,  we  indicated  that  Vampyrops  dorsalis  was  known  from  Costa 
Rica  eastward  into  South  America.  Our  inclusion  of  Costa  Rica  within  the 
known  distribution  of  this  bat  evidently  was  in  error  as  we  now  can  find  no 
published  accounts  of  this  species  to  the  north  of  Panama.  Regarding  new 
distributional  records,  Belize  and  Costa  Rica  can  be  added  to  the  countries 
previously  listed  as  within  the  known  distribution  of  Phylloderma  stenops, 
Michoacan  included  within  the  known  distribution  of  Musonycteris  harrisoni, 
and  Oaxaca  added  to  that  of  Uroderma  magnirostrum.  Also,  Centurio  senex 
now  is  known  on  the  mainland  of  South  America  from  Venezuela. 

Readers  should  be  aware  of  the  Mammalian  Species  series,  published  by  the 
American  Society  of  Mammalogists,  in  which  useful  summaries  of  the  biology 
of  individual  species  of  mammals  are  published.  More  than  100  accounts  thus 
far  have  been  distributed  or  are  in  press,  of  which  eight  of  those  previously 
published  deal  with  phyllostomatids:  Ardops  nichollsi  (Jones  and  Genoways, 
1973),  Hylonycteris  underwoodi  (Jones  and  Homan,  1974),  Macrophyllum 
macrophyllum  (Harrison,  1975),  Macrotus  waterhousii  (Anderson,  1969), 
Monophyllus  redmani  (Homan  and  Jones,  1975a),  M.  plethodon  (Homan  and 
Jones,  19756),  Stenoderma  rufum  (Genoways  and  Baker,  1972),  and  Sturnira 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


thomasi  (Jones  and  Genoways,  1975).  Also  of  interest  is  a  catalogue  of  type 
specimens  of  bats  in  European  museums  that  was  compiled  by  Carter  and 
Dolan  (1978).  In  this  work,  evidence  was  presented  to  establish  the  correct 
spelling  of  Vampyrodes  caraccioli  (spelled  caraccioloi  in  our  checklist). 

Literature  Cited 

Anderson,  S.  1969.  Macrotus  waterhousii.  Mammalian  Species,  1: 1-4. 

Anderson,  S.,  and  C.  E.  Nelson.  1965.  A  systematic  revision  of  Macrotus  (Chiroptera). 
Amer.  Mus.  Novit.,  2212:1-37. 

Baker,  R.  J.,  and  H.  H.  Genoways.  1978.  Zoogeography  of  Antillean  bats.  Pp.  53-97, 
in  Zoogeography  in  the  Caribbean  (F.  B.  Gill,  ed.),  Spec.  Publ.  Acad.  Nat.  Sci. 
Philadelphia,  1 3 :  iii  +  1-128. 

Baker,  R.  J„  H.  H.  Genoways,  and  J.  C.  Patton.  1978.  Bats  of  Guadeloupe.  Occas. 
Papers  Mus.,  Texas  Tech  Univ.,  50:1-16. 

Buden,  D.  W..  1975.  A  taxonomic  and  zoogeographic  appraisal  of  the  big-eared  bat 

( Macrotus  waterhousii  Gray)  in  the  West  Indies.  J.  Mamm.,  56:758-769. 

- .  1976.  A  review  of  the  bats  of  the  endemic  West  Indian  genus  Erophylla.  Proc. 

Biol.  Soc.  Washington,  89:1-15. 

- .  1977.  First  records  of  bats  of  the  genus  Brachyphylla  from  the  Caicos  Islands, 

with  notes  on  geographic  variation.  J.  Mamm.,  58:221-225. 

Carter,  D.  C.,  and  P.  G.  Dolan.  1978.  Catalogue  of  type  specimens  of  Neotropical  bats 
in  selected  European  museums.  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  15:1-136. 
Carter,  D.  C.,  and  J.  K.  Jones,  Jr.  1978.  Bats  from  the  Mexican  state  of  Hidalgo.  Occas. 
Papers  Mus.,  Texas  Tech  Univ.,  54:1-12. 

Davis,  W.  B.  1976.  Notes  on  the  bats  Saccopteryx  canescens  Thomas  and  Micronycteris 
hirsuta  (Peters).  J.  Mamm.,  57:604-607. 

Davis,  W.  B.,  and  D.  C.  Carter.  1978.  A  review  of  the  round-eared  bats  of  the  Tonatia 
silvicola  complex,  with  descriptions  of  three  new  taxa.  Occas.  Papers  Mus., 
Texas  Tech  Univ.,  53:1-12. 

Davis,  W.  B.,  and  J.  R.  Dixon.  1976.  Activity  of  bats  in  a  small  village  clearing  near 
Iquitcs,  Peru.  J.  Mamm.,  57:747-749. 

Gardner,  A.  L.  1976.  The  distributional  status  of  some  Peruvian  mammals.  Occas. 

Papers  Mus.  Zool.,  Louisiana  State  Univ.,  48:1-18. 

Genoways,  H.  H.,  and  R.  J.  Baker.  1972.  Stenoderma  rufum.  Mammalian  Species, 
18:1-4. 

Greenbaum,  I.  F.,  and  J.  K.  Jones,  Jr.  1978.  New  records  of  bats  from  El  Salvador, 
Honduras,  and  Nicaragua.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  55:1-7. 
Greenbaum,  I.  F.,  R.  J.  Baker,  and  D.  E.  Wilson.  1975.  Evolutionary  implications  of 
the  karyotypes  of  the  stenodermine  genera  Ardops,  Ariteus,  Phyllops,  and 
Ectophylla.  Bull.  S.  California  Acad.  Sci.,  74:156-159. 

Handley,  C.  O.,  Jr.  1976.  Mammals  of  the  Smithsonian  Venezuelan  project.  Sci. 

Bull.  Brigham  Young  Univ.,  Biol.  Ser.,  20(5):  (4)  +  1-89  +  (2). 

Harrison,  D.  L.  1975.  Macrophyllum  macrophyllum.  Mammalian  Species,  62:1-3. 
Homan,  J.  A.,  and  J.  K.  Jones,  Jr.  1975«.  Monophyllus  redmani.  Mammalian  Species, 
57:1-3. 

- .  19756.  Monophyllus  plethodon.  Mammalian  Species,  58:1-2. 

Jones,  J.  K.,  Jr.  1978.  A  new  bat  of  the  genus  Artibeus  from  the  Lesser  Antillean  island 
of  St.  Vincent.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  51:1-6. 

Jones,  J.  K.,  Jr.,  and  D.  C.  Carter.  1976.  Annotated  checklist,  with  keys  to  subfamilies 
and  genera.  Pp.  7-38,  in  Biology  of  bats  of  the  New  World  family  Phyllostomatidae. 
Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.).  Spec.  Publ.  Mus.,  Texas 
Tech  Univ.,  10:1-218. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


11 


Jones,  J.  K.,  Jr.,  and  H.  H.  Genoways.  1973.  Ardops  nichollsi.  Mammalian  Species, 
24:1-2. 

- .  1975.  Sturnira  thomasi.  Mammalian  Species,  68: 1-2. 

Jones,  J.  K.,  Jr.,  and  J.  A.  Homan.  1974.  Hylonycteris  underwoodi.  Mammalian 
Species,  32:1-2. 

Jones,  J.  K.,  Jr.,  P.  Swanepoel,  and  D.  C.  Carter.  1977.  Annotated  checklist  of  the  bats 
of  Mexico  and  Central  America.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  47:1-35. 
Klingener,  D.,  H.  H.  Genoways,  and  R.  J.  Baker.  1978.  Bats  from  southern  Haiti. 
Ann.  Carnegie  Mus.,  47:81-99. 

Koopman,  K.  F.  1975.  Bats  of  the  Virgin  Islands  in  relation  to  those  of  the  Greater  and 
Lesser  Antilles.  Amer.  Mus.  Novit.,  2581:1-7. 

- .  1978.  Zoogeography  of  Peruvian  bats  with  special  emphasis  on  the  role  of  the 

Andes.  Amer.  Mus.  Novit.,  2651:1-33. 

LaVal,  R.  K.  1977.  Notes  on  some  Costa  Rican  bats.  Brenesia  (Museo  Nacional  de 
Costa  Rica),  10/1 1:77-83. 

Smith,  J.  D.,  and  H.  H.  Genoways.  1974.  Bats  of  Margarita  Island,  Venezuela,  with 
zoogeographic  comments.  Bull.  S.  California  Acad.  Sci.,  73:64-79. 

Starrett,  A.  1976.  Comments  on  bats  newly  recorded  from  Costa  Rica.  Contrib. 
Sci.,  Los  Angeles  Co.  Mus.  Nat.  Hist.,  277:1-5. 

Starrett,  A.,  and  R.  S.  Casebeer.  1968.  Records  of  bats  from  Costa  Rica.  Contrib. 

Sci.,  Los  Angeles  Co.  Mus.  Nat.  Hist.,  148:1-21. 

Varona,  L.  S.  1974.  Catalogo  de  los  mamfferos  vivientes  y  extinguidos  de  las  Antillas. 
Acad.  Cien.  Cuba,  viii  +  139  pp. 


MORPHOMETRICS 


Pierre  Swanepoel  and  Hugh  H.  Genoways 


In  this  paper,  we  have  attempted  to  cite  all  relevant  literature  in  which  mensural 
data  pertaining  to  phyllostomatid  bats  has  appeared.  We  are  not  so  naive  as  to 
believe  this  goal  was  reached,  but  we  do  believe  most  pertinent  publications 
are  listed,  including  all  major  works  relating  to  each  species.  This  information 
serves  as  a  summary  of  what  currently  is  known  concerning  morphometries  of 
phyllostomatids  and  hopefully  provides  a  basis  for  future  morphometric  studies 
of  members  of  the  family. 

Early  descriptive  accounts  of  phyllostomatids  were  based  mostly  on  material 
preserved  in  fluid  and  generally  lacked  mensural  data;  most  measurements  that 
were  included  were  of  external  dimensions  only.  In  the  late  1800s  and  1900s, 
cranial  measurements  began  to  appear  in  the  literature  as  did  the  first  systematic 
reviews  of  phyllostomatid  groups,  notably  those  dealing  with  Micronycteris 
(Andersen,  1906a),  Carollia  (Hahn,  1907),  Uroderma  and  Artibeus  (Andersen, 
1908),  and  Glossophaga  (Miller,  1913 b).  Through  the  years,  systematic  studies 
have  become  more  and  more  sophisticated,  involving  substantial  mensural 
data  and  complex  methods  of  analysis,  culminating  in  multivariate  analyses  such 
as  those  of  Davis  and  Baker  (1974),  Baker  et  al.  (1972a),  and  Power  and 
Tamsitt  (1973). 

In  the  following  accounts,  papers  in  which  measurements  have  appeared 
are  listed  for  each  species.  Additionally,  when  appropriate  information  is 
available  in  the  published  record  one  or  more  of  the  following  kinds  of  variation 
are  discussed:  age,  individual,  secondary  sexual,  and  geographic.  Accounts  are 
included  for  all  species  listed  by  Jones  and  Carter  (1976).  Within  each  subfamily, 
genera  and  species  are  listed  alphabetically.  A  standard  set  of  measurements 
for  specimens  of  all  species  of  phyllostomatids  is  given  in  Appendix  1.  One 
external  (length  of  forearm)  and  seven  cranial  measurements  (greatest  length 
of  skull,  condylobasal  length,  zygomatic  breadth,  postorbital  constriction, 
breadth  of  braincase,  length  of  maxillary  toothrow,  breadth  across  upper  molars) 
were  taken  with  dial  calipers  from  each  specimen.  Four  males  and  four  females 
were  measured  for  each  species  except  in  those  instances  when  fewer  specimens 
were  available  to  us. 


Acknowledgments 

We  are  especially  grateful  to  Rina  Swanepoel  for  aiding  us  in  innumerable 
ways  including  typing  early  drafts  of  the  manuscript,  arranging  citations,  and 
reading  proof.  We  also  thank  Catherine  H.  Carter  and  Margaret  Popovich 
for  their  help  in  checking  proof  and  Flora  Gibson  for  clerical  assistance. 

We  acknowledge  the  following  curators  for  allowing  us  to  measure  specimens 
in  their  care:  Karl  F.  Koopman,  American  Museum  of  Natural  History  (AMNH); 


13 


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Albert  Schwartz,  private  collection  (AS);  John  Edwards  Hill,  British  Museum 
(Natural  History)  (BMNH);  Robert  Goodwin,  Colgate  University  (COLU); 
Jerry  R.  Choate,  Fort  Hays  State  University  (FHKS);  Robert  S.  Hoffmann, 
University  of  Kansas  (KU);  Lan  A.  Lester,  Natural  History  Museum  of  Los 
Angeles  County  (LACM);  George  H.  Lowery,  Jr.,  Louisiana  State  University 
(LSU);  Randolph  L.  Peterson,  Royal  Ontario  Museum  (ROM);  David  J. 
Schmidly,  Texas  A&M  University  (TCWC);  Robert  J.  Baker,  Texas  Tech 
University  (TTU);  Charles  O.  Handley,  Jr.,  National  Museum  of  Natural 
History  (USNM). 


Subfamily  Phyllostomatinae 
Chrotopterus  auritus  (Peters,  1857) 

Measurements  of  Chrotopterus  auritus  have  been  recorded  as  follows:  Peters  (1857), 
external  measurements  of  the  holotype  of  Chrotopterus  auritus;  Dobson  (1878a), 
external  measurements  of  one  specimen;  Elliot  (1904),  and  Goodwin  (1942a),  external 
and  cranial  measurements  of  one  specimen;  Elliot  (1917),  external  measurements  of  one 
specimen;  Anthony  (1920),  external  and  cranial  measurements  of  holotype  of  C.  colombianus 
(sex  unknown)  from  Colombia;  Lima  (1926),  external  measurements  of  a  male  from 
Brazil;  Cunha  Vieira  (1942),  external  measurements  of  four  males  and  a  female  and  cranial 
measurements  of  a  male  and  female  from  Brazil;  Goodwin  (1946),  external  and  cranial 
measurements  of  a  male  from  Brazil;  Hall  and  Kelson  (1959),  cranial  measurements  of  a 
male  and  female  from  Veracruz;  Burt  and  Stirton  (1961),  external  and  cranial  measurements 
of  a  specimen  from  El  Salvador;  Villa-R.  (1967),  external  and  cranial  measurements  of  a 
male  from  Mexico;  Rick  (1968),  forearm  and  cranial  measurements  of  three  males  and  a 
female  from  Costa  Rica;  Goodwin  (1969),  forearm  and  cranial  measurements  of  two  males 
(one  subadult)  from  Chiapas;  Villa-R.  and  Villa  Cornejo  (1969),  external  and  cranial 
measurements  of  a  male  and  two  females  from  Argentina;  Taddei  (1975a),  external 
measurements  of  six  specimens  and  cranial  measurements  of  seven  specimens  (mean, 
se,  range)  of  males  and  females  combined  from  Brazil. 

Individual  variation. — Coefficients  of  variation  for  external  (N=  6,  males  and  females 
combined)  and  cranial  measurements  (N—  7,  males  and  females  combined)  of  specimens 
from  Brazil  ranged  from  1.89  to  5.37  in  external  measurements  and  from  0.84  to  4.08 
in  cranial  measurements  (Taddei,  1975a). 

Lonchorhina  aurita  Tomes,  1863 

Measurements  for  Lonchorhina  aurita  have  been  recorded  as  follows:  Tomes  (1863), 
external  and  cranial  measurements  of  the  holotype  of  L.  aurita;  Peters  (18666),  external 
measurements  of  one  specimen;  Dobson  (1878a),  external  measurements  of  the  holotype 
from  Trinidad;  Elliot  (1904),  external  and  cranial  measurements  of  one  specimen;  Miller 
(1912),  external  and  cranial  measurements  of  a  male  and  female  from  Panama;  Anthony 
(1923),  external  and  cranial  measurements  of  the  male  holotype  of  L.  aurita  occidentalis 
from  Ecuador,  and  forearm  measurements  of  three  specimens  and  cranial  measurements 
of  one  specimen  of  L.  aurita  aurita  from  Venezuela;  Cunha  Vieira  (1942),  external 
measurements  of  three  males  and  cranial  measurements  of  two  males  from  Brazil;  Goodwin 
(1942a),  external  measurements  of  a  specimen  from  Honduras;  Goodwin  (1946),  external 
and  cranial  measurements  of  a  male  and  female  from  Panama;  Goodwin  (1953),  external 
and  cranial  measurements  of  the  holotype  of  L.  a.  occidentalis  as  given  by  Anthony  (1923); 
Felten  (1956a),  external  measurements  of  two  males  and  cranial  measurements  of  a  male 
from  El  Salvador;  Hall  and  Kelson  (1959),  external  and  cranial  measurements  of  a  male 
and  female  from  Panama;  Burt  and  Stirton  (1961),  external  measurements  of  two  males 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


15 


and  cranial  measurements  of  a  male  from  El  Salvador;  Goodwin  and  Greenhall  (1961), 
forearm  and  cranial  measurements  of  two  females  and  a  juvenile  male  from  Trinidad; 
Pirlot  (1967),  external  measurements  of  one  specimen;  Villa-R.  (1967),  external 
measurements  of  22  and  cranial  measurements  of  21  males  and  females  combined  (mean, 
sd,  and  range)  from  Mexico;  Goodwin  (1969),  forearm  and  cranial  measurements  of  four 
females  from  Oaxaca;  Tuttle  (1970),  external  measurements  of  a  male  and  two  females 
from  Peru;  Linares  and  Ojasti  (1971),  external  and  cranial  measurements  of  26  specimens 
from  Trinidad  and  Venezuela. 

Lonchorhina  orinocensis  Linares  and  Ojasti,  1971 

Linares  and  Ojasti  (1971)  gave  external  and  cranial  measurements  (mean,  sd,  range) 
of  five  specimens  from  Venezuela,  including  the  female  holotype. 

Macrophyllum  macrophyllum  (Schinz,  1821) 

Measurements  for  Macrophyllum  macrophyllum  have  been  recorded  as  follows:  Dobson 
(1878a),  external  measurements  of  a  specimen  from  Brazil;  Cunha  Vieira  (1942)  external 
measurements  of  two  females  and  cranial  measurements  of  one  female  from  Brazil; 
Goodwin  (1946),  external  and  cranial  measurements  of  a  male  from  Guyana;  Felton 
(1956a),  external  measurements  (mean,  range)  of  five  males  and  cranial  measurements  of 
three  males  from  El  Salvador;  Hall  and  Kelson  (1959),  external  and  cranial  measurements 
of  a  male  from  Guyana;  Hill  and  Bown  (1963),  external  and  cranial  measurements  of  a 
male  and  female  from  Ecuador;  Davis  et  al.  (1964),  external  measurements  of  two  males 
from  Nicaragua;  Hill  (1964),  forearm  and  cranial  measurements  of  a  male  from  Guyana; 
Starrett  and  Casebeer  (1968),  forearm  and  cranial  measurements  of  a  male  and  female 
from  Costa  Rica;  Harrison  and  Pendleton  (1974),  external  and  cranial  measurements 
of  nine  males  and  three  females  from  El  Salvador;  Harrison  (1975),  forearm  and  cranial 
measurements  (range)  for  the  species;  Taddei  (1975a),  external  and  cranial  measurements 
(mean,  sd,  range)  of  eight  males  from  Brazil. 

Individual  variation. — Taddei  (1975a)  gave  coefficients  of  variation  for  external  (0.48- 
8.03)  and  cranial  measurements  (0.27-3.51)  for  eight  males  from  Brazil. 

Macrotus  cal ifornicus  Baird,  1858 

Measurements  of  Macrotus  californicus  have  been  recorded  as  follows:  Baird  (1858), 
external  measurements  of  a  specimen  from  California  in  the  original  description  of  M. 
californicus;  H.  Allen  (1864),  external  measurements  of  eight  specimens;  H.  Allen  (1894a, 
18946),  mean  external  and  cranial  measurements  of  four  individuals  and  external  measure¬ 
ments  of  another  eight  specimens;  Elliot  (1901),  external  measurements  of  one  specimen; 
Elliot  (1904),  external  and  cranial  measurements  of  one  specimen;  Rehn  (1904),  external 
measurements  (mean,  range)  of  five  topotypes  (Imperial  Company,  California),  and 
cranial  measurements  (mean,  range)  of  six  specimens;  Stephens  (1906),  external  measure¬ 
ments  of  one  specimen;  Grinnell  (1918),  external  and  cranial  measurements  of  18  females 
from  California;  Hall  (1946),  external  measurements  of  two  males  and  mean  and  range 
of  nine  females  and  cranial  measurements  of  a  male  and  a  female  from  Nevada;  Anderson 
and  Nelson  (1965),  external  and  cranial  measurements  (mean,  sd,  range)  of  four  samples  from 
throughout  the  geographic  range  of  the  species;  Villa-R.  (1967),  external  measurements 
(mean,  se,  range)  of  five  males  and  eight  females  and  cranial  measurements  (mean,  se, 
range)  of  five  males  and  four  females  from  Mexico;  Anderson  (1972),  external  measure¬ 
ments  of  a  large  sample  and  cranial  measurements  of  one  individual  from  Chihuahua. 

Secondary  sexual  variation. — Anderson  and  Nelson  (1965)  reported  no  secondary 
sexual  dimorphism  in  28  males  and  30  females  from  California. 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Geographic  variation. — According  to  Anderson  and  Nelson  (1965),  there  is  a 
geographic  uniformity  in  characters  of  populations  from  the  southern  end  of  Baja 
California  north  to  California,  Nevada,  and  Arizona  and  then  southward  through  Sonora. 
Consequently,  they  recognized  no  geographic  races  within  the  area  that  is  now  considered 
to  constitute  the  distribution  of  M.  californicus. 

Macrotus  waterhousii  Gray,  1843 

Measurements  of  Macrotus  waterhousii  have  been  recorded  as  follows:  Saussure 
(1860c),  external  measurements  of  one  specimen;  Gundlach  (1872,  1877),  external 

measurements  of  a  Cuban  specimen;  Dobson  (1876),  external  measurements  of  the 
holotype  of  M.  hocourtianus  from  Guatemala;  Dobson  (1878a),  external  measurements 
of  two  specimens;  H.  Allen  (1890a),  external  measurements  of  one  specimen  in  the  original 
description  of  M.  vv.  bulleri  from  Jalisco;  H.  Allen  (1894a),  external  measurements  of  one 
specimen  probably  from  Jalisco;  J.  A.  Allen  (1904),  external  measurements  (mean)  of 
seven  specimens  from  Tehuantepec,  Oaxaca,  compared  to  those  of  one  specimen  from 
Yautepec,  Morelos;  Elliot  (1904),  external  and  cranial  measurements  of  four  specimens; 
Rehn  (1904),  external  and  cranial  measurements  of  the  various  subspecies  (revision  of 
the  genus);  Elliot  (1905),  range  of  external  and  cranial  measurements  of  the  different 
subspecies;  Shamel  (1931),  external  and  cranial  measurements  of  the  male  holotype  of 
M.  w.  herberfolium  from  Providencialis  Island  and  the  measurement  range  of  five 
specimens  ( =  M.  vv.  waterhousii)  Hispaniola;  Martinez  and  Villa-R.  (1938),  external 
measurements  of  three  specimens  and  cranial  measurements  of  two  from  Morelos; 
Martinez  and  Villa-R.  (1940),  external  and  cranial  measurements  (mean,  sd)  of  samples 
of  males  and  females  from  the  Guerrero;  Goodwin  (1942a),  external  and  cranial  measure¬ 
ments  of  one  specimen;  Anderson  and  Nelson  (1965),  external  and  cranial  measure¬ 
ments  (mean,  sd,  range)  of  12  samples  from  throughout  the  geographic  range  of  the  species; 
Choate  and  Birney  (1968),  cranial  measurements  of  subfossil  specimens  from  Puerto 
Rico;  Anderson  (1969),  external  measurements  for  the  genus  as  the  two  species  are 
treated  conspecifically  under  M.  waterhousii-,  Goodwin  (1969),  forearm  and  cranial 
measurements  of  three  males  and  two  females  from  Oaxaca;  Alvarez  and  Ramirez- 
Pulido  (1972),  external  and  cranial  measurements  (mean,  range)  of  11  specimens  from 
Tamaulipas  and  San  Luis  Potosi;  Silva-Taboada  (1974),  measurements  of  fossil 
mandibles  from  Cuba;  Buden  (19756),  external  and  cranial  measurements  (mean,  sd, 
range)  of  large  samples  from  northern  Bahamas,  southern  Bahamas,  Cuba,  Hispaniola, 
Jamaica,  and  means  of  smaller  samples  from  Isle  of  Pines,  Grand  Cayman,  and  Navassa 
for  sexes  combined. 

Individual  variation. — In  specimens  from  Guerrero,  coefficients  of  variation 
(CV)  for  external  measurements  varied  in  males  from  1.93  to  11.16  and  in  females  from 
1.67  to  8.09;  for  cranial  measurements,  in  males  from  1.36  to  3.08  and  in  females  from 
0.65  to  3.90  (Martinez  and  Villa-R.,  1940). 

According  to  Anderson  and  Nelson  (1965),  length  of  skull  proved  to  be  the  least 
variable  character,  and  then  in  order  of  increasing  variability  were  the  breadth  of  brain- 
case,  length  of  bulla,  interorbital  breadth,  and  breadth  at  canines.  External  measure¬ 
ments  were  generally  more  variable  than  cranial  measurements.  The  coefficient  of 
variation  for  total  length,  however,  was  usually  no  greater  than  that  of  the  more  variable 
cranial  measurements. 

Buden  (19756)  showed  in  West  Indian  specimens  that  cranial  (except  breadth  at 
canines)  and  forearm  measurements  were  the  least  variable  measurements,  whereas  tail 
length  generally  showed  extremely  high  CVs.  Forearm  and  cranial  CV  values,  other  than 
that  of  breadth  at  canines,  ranged  from  1.03  to  3.58;  values  for  breadth  at  canines  varied 
from  2.78  to  4.63.  The  coefficient  of  variation  values  observed  in  tail  length  ranged 
from  6.19  to  9.13. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


17 


Geographic  variation. — Anderson  and  Nelson  (1965)  noted  an  increase  in  size  from 
northwest  to  southeast  through  the  range  of  Macrotus  waterhousii.  This  held  true  for  all 
measurements  except  length  of  bulla,  which  increased  in  size  from  southeast  to  northwest. 
Specimens  from  eastern  Cuba  were  larger  than  those  from  the  western  end  of  the  island. 
However,  samples  from  different  parts  of  western  Cuba  and  the  Isle  of  Pines  did  not  differ 
significantly  in  size  (Anderson  and  Nelson,  1965).  Geographic  variation  was  found  within 
Hispaniolan  samples — those  from  Haiti  averaged  larger  than  those  from  the  Dominican 
Republic.  Populations  on  Hispaniola  were  larger  in  size  than  those  on  Cuba  and  the 
southern  Bahamas.  Specimens  from  several  northern  Bahaman  islands  were  not  significantly 
different  in  size  but  averaged  larger  than  those  from  Cuba  (Anderson  and  Nelson,  1965) 
and  smaller  than  those  from  the  southern  Bahamas  and  Hispaniola.  Bats  from  Jamaica, 
according  to  Anderson  and  Nelson  (1965),  were  larger  than  those  from  Cuba,  and  inter¬ 
mediate  in  size  between  Cuban  and  southern  Bahaman  and  Hispaniolan  populations  (Ander¬ 
son  and  Nelson,  1965:21).  Specimens  from  Oaxaca  averaged  significantly  larger  than  those 
from  Morelos  (region  of  the  type  locality)  but  were  not  as  large  as  specimens  from  Hispan¬ 
iola  and  the  southern  Bahamas.  Specimens  from  Oaxaca  averaged  larger  than  the  western 
Cuban  specimens.  A  sample  from  Morelos,  Guerrero,  and  Puebla  were  only  slightly  larger 
in  cranial  size  than  a  sample  from  Jalisco. 

Buden  (19756)  stated  that  the  statistical  data  he  used  were  comparable  to  those  of 
Anderson  and  Nelson  (1965)  but  concluded  that  a  dendrogram,  based  on  levels  of 
morphological  differences,  placed  the  northern  Bahaman  specimens  with  the  Cuban 
ones.  An  increase  in  specimen  size  from  southwest  to  northeast  throughout  the  West  Indies 
(western  to  eastern  Cuba  to  northern  Bahamas;  and  Jamaica,  Hispaniola,  to  southern 
Bahamas)  was  found.  Ear  length,  however,  did  not  show  this  pattern  (Buden,  19756). 
Buden  (19756)  also  described  an  increase  in  size  from  western  Cuba  to  eastern  Cuba 
as  did  Anderson  and  Nelson  (1965).  However,  in  contrast  to  Anderson  and  Nelson, 
Buden  did  not  find  intra-island  variation  on  Hispaniola. 

Davis  and  Baker  (1974)  reported  a  general  trend  of  size  increase  on  the  mainland 
from  north  to  south  in  all  measurements.  Their  multivariate  analyses  showed  that  the 
groups  were  nonclinally  tied  one  to  another  with  respect  to  geography. 

Micronycteris  behni  (Peters,  1865) 

Measurements  of  Micronycteris  behni  have  been  recorded  as  follows:  Peters  (1865  6), 
external  measurements  of  the  holotype  from  Brazil;  Dobson  (1878a),  external  measure¬ 
ments  of  a  specimen;  Andersen  (1906a),  external  measurements  of  two  specimens  and 
cranial  measurements  of  one  specimen  from  Peru;  Sanborn  (1949a),  range  of  forearm 
length  in  the  species. 


Micronycteris  brachyotis  (Dobson,  1878) 

Measurements  of  Micronycteris  brachyotis  have  been  recorded  as  follows:  Dobson 
(18786),  external  measurements  of  the  male  holotype  of  M.  brachyotis  from  Cayenne;  Miller 
(1900c),  forearm  length  for  M.  brachyotis ;  Andersen  (1906a),  external  and  cranial  measure¬ 
ments  of  the  holotype  of  M.  brachyotis  (after  Dobson  18786);  Sanborn  (1949a),  external 
and  cranial  measurements  of  the  holotype  and  two  topotypes  of  M.  platyceps  (=  M. 
brachyotis)  and  external  measurements  of  four  additional  specimens  from  Trinidad;  Hall 
and  Kelson  (1959),  external  and  cranial  measurements  of  the  holotype  of  M.  platyceps,  two 
topotypes,  and  one  female;  Goodwin  and  Greenhall  (1961),  forearm  measurements  (range) 
of  16  specimens  from  Trinidad,  cranial  measurements  of  one  male  and  two  females  in¬ 
cluding  the  holotype  of  M.  platyceps,  and  a  comparison  of  external  and  cranial  measure¬ 
ments  of  a  large  adult  male  from  Trinidad  and  the  holotype  of  M.  brachyotis  from  Cayenne; 
Davis  et  al.  (1964),  external  and  cranial  measurements  of  a  female  from  Chiapas;  Jones 
(1966),  forearm  and  cranial  measurements  of  a  male  from  Guatemala;  Villa-R.  (1967), 


18 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


external  measurements  of  one  specimen  from  Oaxaca;  Rick  (1968),  external  and  cranial 
measurements  of  eight  males  and  one  female  from  Guatemala;  Goodwin  (1969),  forearm 
and  cranial  measurements  of  a  male  from  Oaxaca;  Marinkelle  and  Cadena  (1972),  forearm 
measurement  of  one  male  from  Colombia,  and  external  and  cranial  measurements  of  one 
female  from  Colombia;  Starrett  (1976),  forearm  measurements  of  a  female,  male,  and 
juvenile  male  from  Costa  Rica. 

Geographic  variation. — The  holotype  of  M.  brachyotis  from  Cayenne,  an  old  male 
with  worn  teeth,  was  larger  than  a  series  of  specimens  from  Trinidad  but  not  larger 
than  a  speciment  of  M.  platyceps  from  Nicaragua  (Goodwin  and  Greenhall,  1961). 

Micronycteris  (=  Barticonycteris)  daviesi  (Hill,  1964) 

Measurements  of  Micronycteris  daviesi  have  been  recorded  as  follows:  Hill  (1964), 
external  and  cranial  measurements  of  the  female  holotype  from  Guyana;  Tuttle  (1970), 
external  measurements  of  two  males  and  one  female  from  Peru. 

Micronycteris  hirsuta  (Peters,  1869) 

Measurements  of  Micronycteris  hirsuta  have  been  recorded  as  follows:  Peters  (1869), 
external  measurements  of  the  holotype;  Dobson  (1878a),  external  measurements  of  one 
specimen;  Elliot  (1904),  external  measurements  of  one  specimen  from  Costa  Rica; 
Andersen  (1906a),  external  measurements  of  two  specimens  and  cranial  measure¬ 
ments  of  one  from  Costa  Rica;  Sanborn  (1932),  external  and  cranial  measurements  of  a 
female  from  Colombia;  Goodwin  (1946),  external  and  cranial  measurements  of  a  male 
and  female  from  Costa  Rica;  Hershkovitz  (1949),  external  and  cranial  measurements  of 
two  males  and  one  female  from  northern  Colombia;  Sanborn  (1949a),  range  of  forearm 
and  greatest  length  of  skull  for  the  species;  Hall  and  Kelson  (1959),  external  and  cranial 
measurements  of  a  male  and  female  from  Costa  Rica;  Goodwin  and  Greenhall  (1961), 
forearm  length  (range)  of  12  specimens,  and  cranial  measurements  of  three  males  and 
two  females  from  Trinidad;  Hill  (1964),  forearm  and  cranial  measurements  of  one  female 
from  Guyana;  LaVal  (1969),  external  and  cranial  measurements  of  a  male  and  female 
from  Honduras;  Gardner  et  al.  (1970),  external  and  cranial  measurements  of  one  male 
from  Costa  Rica;  Valdez  and  LaVal  (1971),  external  and  cranial  measurements  of  two 
males  from  Nicaragua;  Baker  et  al.  (1973),  forearm  and  cranial  measurements  (mean, 
se,  range,  CV)  of  two  samples,  one  from  Trinidad  (four  specimens)  and  the  other  from 
Honduras  (one  specimen)  and  Nicaragua  (four  specimens). 

Individual  variation. — Coefficients  of  variation  in  forearm  and  cranial  measurements 
obtained  from  four  specimens  from  Trinidad  revealed  little  variation  (CV,  0.8-2. 3), 
whereas  one  specimen  from  Honduras  and  four  from  Nicaragua  combined  showed  higher 
values  than  those  from  Trinidad  (CV,  1. 2-4.1)  (Baker  et  al.,  1973). 

Geographic  variation. — Valdez  and  LaVal  (1971)  recorded  this  species  for  the  first 
time  from  Nicaragua  and  showed  that  the  two  specimens  obtained  were  smaller  than 
those  from  Costa  Rica  and  other  countries  recorded  by  Goodwin  (1946),  Sanborn 
(1949a),  Goodwin  and  Greenhall  (1961),  and  Gardner  et  al.  (1970).  However,  these 
Nicaraguan  specimens  proved  to  differ  little  from  Honduran  specimens  (LaVal,  1969). 
Forearm  and  cranial  measurements  of  specimens  from  Trinidad  averaged  larger  than 
those  for  specimens  from  Honduras  and  Nicaragua,  but  only  forearm  and  greatest  length 
of  skull  proved  to  be  significantly  different  (Baker  et  al.,  1973). 

Micronycteris  megalotis  (Gray,  1842) 

Measurements  of  Micronycteris  megalotis  have  been  recorded  as  follows:  Dobson 
(1878a),  external  and  cranial  measurements  of  one  specimen;  Miller  (1898),  external 
measurements  for  specimens  from  Nicaragua  (including  the  male  holotype  of  M.  m. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


19 


microtis),  Trinidad  (one  male),  Margarita  (one  male  and  female),  Colombia  (two  males  and 
females),  Honduras  (two  males),  Colima  (four  males  and  three  females),  Jalisco  (two  males 
and  three  females),  and  Oaxaca  (one  female);  Miller  (1900c),  forearm  length  for  M. 
m.  microtis',  Robinson  and  Lyon  (1901),  external  measurements  of  five  males  and  six 
females  from  Venezuela;  Elliot  (1904),  external  and  cranial  measurements  of  one  specimen 
and  external  measurements  of  the  holotype  of  M.  m.  microtis ;  Rehn  (1904),  external 
and  cranial  measurements  of  the  holotype  of  Macrotus  pygmaeus  (=  Micronycteris 
megalot  is)  and  one  male  from  Yucatan;  Andersen  (1906a),  external  measurements  of 
the  holotype  of  M.  m.  microtis  (after  Miller  1898),  external  and  cranial  measurements 
(range)  of  30  (18  cranial)  specimens  from  Brazil,  Peru,  Guyana,  Venezuela,  Trinidad 
and  Tobago,  and  of  10  (nine  cranial)  specimens  from  Colombia,  Guatemala,  Honduras 
and  Mexico;  Lyon  (1906),  ear  measurements  of  the  holotype  of  M.  m.  microtis  and  a 
specimen  from  Venezuela;  Lima  (1926),  external  measurements  of  a  male  from  Brazil; 
Goodwin  (1934),  external  measurements  of  one  specimen  from  Guatemala;  Martinez 
and  Villa-R.  (1938),  external  measurements  of  one  specimen  from  Morelos;  Cunha 
Vieira  (1942),  external  measurements  of  four  males  and  cranial  measurements  of  two 
males  from  Brazil;  Goodwin  (1942a),  forearm  and  cranial  measurements  of  two  specimens 
of  unknown  sex  from  Honduras;  Goodwin  (1946),  external  and  cranial  measurements  of 
two  males  from  Costa  Rica;  Sanborn  (1949a),  range  of  forearm  length  of  three  subspecies; 
Hershkovitz  (1949),  forearm  measurement  of  one  specimen  and  skull  measurements 
of  another,  both  from  Trinidad;  Dalquest  (1953a),  external  measurements  of  eight  males 
and  10  females,  and  cranial  measurements  of  seven  males  and  nine  females  from  San 
Luis  Potosi;  Goodwin  (1953),  external  and  cranial  measurements  of  the  holotype  Macrotus 
pygmaeus  from  Yucatan;  Goodwin  (1954),  external  measurements  of  a  specimen  from 
Tamaulipas;  Felten  (1956a),  external  and  cranial  measurements  of  two  males  from  El 
Salvador;  Felten  (1956 d),  external  measurements  (mean,  range)  of  specimens  from 
El  Salvador;  Goodwin  and  Greenhall  (1961),  forearm  measurements  of  three  specimens 
from  Trinidad  and  three  from  Tobago  (unsexed),  and  cranial  measurements  of  a  male 
from  Trinidad;  Burt  and  Stirton  (1961),  range  of  forearm  and  cranial  measurements  of 
eight  males  and  five  females  combined  from  El  Salvador;  Husson  (1962),  external  and 
cranial  measurements  of  six  males  and  three  females  from  Surinam;  Tamsitt  and 
Valdivieso  (1963a),  mean  and  range  of  external  and  cranial  measurements  of  three 
males  and  four  females  combined  from  Colombia;  Valdivieso  (1964),  mean  and  range 
of  external  and  cranial  measurements  of  specimens  from  Colombia;  Brosset  (1965), 
external  and  cranial  measurements  of  two  males  from  Ecuador;  Villa-R.  (1967),  external 
measurements  of  six  males  and  10  females,  and  cranial  measurements  of  eight  males 
and  seven  females  from  Mexico;  Pirlot  (1968),  forearm  measurement  of  a  male  from 
Peru;  Goodwin  (1969),  forearm  and  cranial  measurements  of  four  males  and  five 
females  from  Oaxaca;  Gardner  et  al.  (1970),  wing  and  cranial  measurements  (mean, 
range)  of  six  males  and  one  female  combined  from  Costa  Rica;  Jones  et  al.  (19716), 
mean  and  range  of  forearm  and  cranial  measurements  of  three  males  and  five  females  from 
westcentral  Nicaragua,  of  three  males  and  three  females  from  Isla  del  Maiz  Grande,  and 
of  three  males  and  three  females  from  Rio  Coco,  and  forearm  and  cranial  measurements 
of  one  male  from  Bonanza,  Nicaragua,  and  cranial  measurements  of  the  M.  m.  microtis 
|  holotype  (male)  from  Greytown,  Nicaragua;  Watkins  et  al.  (1972),  forearm  and  cranial 
measurements  of  two  males  and  females  from  Jalisco;  Jones  et  al.  (1973),  forearm  and 
cranial  measurements  of  three  males  from  the  Yucatan  Peninsula;  Birney  et  al.  (1974), 
forearm  and  cranial  measurements  of  a  female  from  Yucatan;  Smith  and  Genoways 
(1974),  forearm  and  cranial  measurements  of  a  male  and  female  from  Margarita  Island, 
Venezuela;  Taddei  (1975a),  external  and  cranial  measurements  (mean,  se,  range,  CV) 
of  males  and  females  combined  (N=  10)  from  Brazil. 

Individual  variation. — Coefficients  of  variation  for  10  specimens  (sexes  combined)  from 
Brazil  were  given  for  external  and  cranial  measurements  by  Taddei  (1975a).  Cranial 


20 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


measurements  showed  little  variation  (CV,  0.66  to  3.18),  whereas  those  for  external  measure¬ 
ments  were  more  variable  (CV,  1.77  to  5.48). 

Geographic  variation. — Variation  in  size  in  M.  megalotis  between  two  localities  in 
Costa  Rica  (Fila  la  Maquina,  Cordillera  Talamaca,  6600  to  8700  feet;  Rincon  and  Tilaran, 
below  700  feet)  were  discussed  by  Gardner  et  al.  (1970).  Those  from  the  higher  altitude 
proved  to  be  larger  than  those  from  the  lower.  Size  differences  were  particularly  evident 
in  wing  dimensions;  no  difference  in  ear  length  was  observable  (see  also  Jones  et  al.,  19716). 
Although  cranial  measurements  seemed  to  be  more  or  less  equal,  specimens  from  the  higher 
altitude  tended  to  be  larger. 

Jones  et  al.  (1971  b)  concluded  that  specimens  from  westcentral  Nicaragua  and  Isla  del 
Mafz  Grande  were,  on  the  average,  considerably  larger  in  skull  and  forearm  measurements 
than  the  holotype  of  M.  m.  microtis  from  Greytown,  eastern  Nicaragua.  Specimens  from 
Rio  Coco  were  intermediate  between  the  two  morphological  types  leading  these  authors 
to  suggest  that  intergradation  occurred  between  them.  No  difference  in  ear  length  was 
found.  In  the  original  description.  Miller  (1898)  claimed  that  M.  m.  microtis  was 
characterized  by  much  smaller  ears.  Lyon  (1906)  presented  evidence  that  the  ears  of  the 
holotype  were  small  and  not  damaged.  Forearm  measurements  of  four  specimens  previously 
obtained  from  Isla  del  Maiz  Grande  (G.  M.  Allen,  1929)  were  also  relatively  big  according 
to  J ones  et  al.  (19716). 


Micronycteris  minuta  (Gervais,  1856) 

Measurements  of  Micronycteris  minuta  have  been  recorded  as  follows:  Dobson  (1878c/), 
external  measurements  of  one  specimen  from  Brazil;  Thomas  (1901c),  forearm  measure¬ 
ments  of  the  holotype  as  given  by  both  Gervais  and  Dobson;  Andersen  (1906«),  external 
measurements  of  eight  specimens  (range)  and  cranial  measurements  of  six  specimens  (range) 
from  Brazil;  G.  M.  Allen  (1908),  external  and  cranial  measurements  of  one  female  from 
Brazil;  Cunha  Vieira  (1942),  external  measurements  of  a  male  from  Brazil;  Sanborn  (1949c/), 
range  of  forearm  length  in  the  species,  forearm  and  cranial  measurements  of  one  specimen 
from  Colombia;  Goodwin  (1953),  external  measurements  of  the  female  holotype  of  M. 
hypoleuca  ( =  M.  minuta )  from  Colombia;  Goodwin  and  Greenhall  (1961),  range  of 
forearm  length  of  12  specimens  and  cranial  measurements  of  one  male  and  two  females 
from  Trinidad;  Linares  (1969),  external  and  cranial  measurements  of  a  male  and  female 
from  Venezuela;  Gardner  et  al.  (1970),  mean  and  range  of  external  and  cranial  measure¬ 
ments  of  four  specimens  (three  males,  one  female)  from  Costa  Rica;  Valdez  and  LaVal 
(1971),  external  and  cranial  measurements  of  one  male  from  Nicaragua  and  the  range  of 
measurements  of  three  males  and  one  female  from  Costa  Rica. 

Geographic  variation. — According  to  Sanborn  (1949c/),  specimens  from  Brazil  appeared 
to  be  larger  than  specimens  from  Colombia. 

Micronycteris  nicefori  Sanborn,  1949 

Measurements  of  Micronycteris  nicefori  have  been  recorded  as  follows:  Sanborn  (1949c/), 
external  and  cranial  measurements  of  the  male  holotype  and  the  range  of  measurements 
of  four  paratypes  from  Colombia;  Goodwin  and  Greenhall  (1961),  forearm  length  of  the 
holotype,  the  range  of  this  measurement  in  five  specimens  from  Trinidad,  and  cranial 
measurements  of  the  holotype  (male)  and  a  male  and  female  from  Trinidad;  Hill  (1964), 
forearm  (two  males)  and  cranial  measurements  of  one  specimen  from  Guyana;  Baker  and 
Jones  (1975),  external  and  cranial  measurements  of  a  female  from  Nicaragua;  Starred 
(1976),  external  and  cranial  measurements  of  five  males  and  cranial  measurements  of  one 
male  from  Costa  Rica;  LaVal  (1977),  forearm  length,  greatest  length  of  skull,  and  weight  of 
a  male  from  Costa  Rica. 

Geographic  variation. — According  to  Starred  (1976),  his  specimens  from  Costa  Rica 
agreed  closely  in  most  measurements  with  those  given  by  Sanborn  (1949c/)  for  specimens 
from  Colombia. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


21 


Micronycteris  pus  ilia  Sanborn,  1949 

Measurements  of  Micronycteris  pusilla  have  been  recorded  as  follows:  Sanborn  (1949a), 
external  and  cranial  measurements  of  the  male  holotype  from  Brazil;  Goodwin  (1953), 
forearm  and  cranial  measurements  of  the  holotype. 

Micronycteris  schmidtorum  Sanborn,  1935 

Measurements  of  Micronycteris  schmidtorum  have  been  recorded  as  follows:  Sanborn 
(1935),  external  and  cranial  measurements  of  the  holotype  and  paratype  (both  males) 
from  Guatemala;  Goodwin  (1942a),  external  and  cranial  measurements  of  the  holotype 
from  Guatemala;  Sanborn  (1949a),  range  of  forearm  measurements  in  the  species;  Hall 
and  Kelson  (1959),  external  and  cranial  measurements  of  the  holotype  from  Guatemala 
and  one  male;  Davis  et  al.  (1964),  external  and  cranial  measurements  of  a  male  from 
Nicaragua;  Villa-R.  (1967),  external  and  cranial  measurements  of  two  specimens  from 
Yucatan;  Starrett  and  Casebeer  (1968),  forearm  (two  males,  mean  and  range  of  five  females) 
and  cranial  measurements  (two  males,  two  females)  from  Guanacaste,  Costa  Rica;  Jones 
et  al.  (1973),  forearm  and  cranial  measurements  of  one  juvenile  female  from  the  Yucatan 
Peninsula;  Baker  and  Jones  (1975),  external  and  cranial  measurements  of  a  male  from 
Nicaragua. 


Micronycteris  sylvestris  (Thomas,  1896) 

Measurements  of  Micronycteris  sylvestris  have  been  recorded  as  follows:  Thomas 
(1896),  external  and  cranial  measurements  of  the  male  holotype  from  Costa  Rica;  Elliot 
(1904a),  external  and  cranial  measurements  of  one  specimen;  Andersen  (1906a),  external 
and  cranial  measurements  of  the  male  holotype  from  Costa  Rica;  Goodwin  (1946),  external 
and  cranial  measurements  of  the  male  holotype  from  Costa  Rica;  Hall  and  Kelson  (1959), 
cranial  measurements  of  the  holotype  of  M.  sylvestris  and  one  male;  Goodwin  and  Green- 
hall  (1961),  forearm  and  cranial  measurements  (range)  of  four  males  from  Trinidad  and 
four  males  from  Veracruz;  Villa-R.  (1967),  external  measurements  (mean,  range)  of  nine 
specimens  and  cranial  measurements  (mean,  range)  of  five  specimens  from  Colima  and 
Jalisco;  Goodwin  (1969),  forearm  and  cranial  measurements  of  two  females  from  Veracruz; 
Linares  (1969),  external  and  cranial  measurements  of  a  female  from  Venezuela. 

Geographic  variation. — Specimens  from  Trinidad  were  similar  to  Mexican  and  Central 
American  specimens;  however,  skulls  of  the  material  from  Trinidad  were  relatively  shorter 
than  those  from  Mexico  (Goodwin  and  Greenhall,  1961). 

Mimon  bennettii  (Gray,  1838) 

Measurements  of  Mimon  bennettii  have  been  recorded  as  follows:  Saussure  (1860c), 
external  measurements  of  one  specimen  of  Vampirus  auriculas  (=  M.  bennettii)',  Peters 
(18666),  external  measurements  of  a  specimen  from  Brazil;  Dobson  (1878a),  external  measure¬ 
ments  of  one  specimen;  Lima  (1926),  external  measurement  of  a  specimen  from  Brazil; 
Cunha  Vieira  (1942),  external  and  cranial  measurements  of  a  female  from  Brazil;  Dalquest 
(1957),  external  and  cranial  measurements  of  one  specimen  from  Brazil;  Husson  (1962), 
external  and  cranial  measurements  of  two  females  from  Surinam;  Hill  (1964),  forearm  and 
cranial  measurements  of  a  male  from  Brazil. 

Mimon  cozumelae  Goldman,  1914 

Measurements  of  Mimon  cozumelae  have  been  recorded  as  follows:  Goldman  (19146), 
external  and  cranial  measurements  of  the  holotype  from  Cozumel  Island  off  the  east  coast 
of  Yucatan;  Elliot  (1917),  external  and  cranial  measurements  of  the  holotype;  Sanborn 
(1941),  external  measurements  of  two  specimens  from  Yucatan;  Goodwin  (1942a,  1946), 
external  measurements  of  a  male  and  female  from  Yucatan;  Dalquest  (1957),  external 
and  cranial  measurement  (mean)  of  10  specimens  from  Veracruz;  Hall  and  Kelson  (1959), 


22 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


forearm  and  cranial  measurements  of  the  holotype  of  M.  cozumelae.  Carter  et  al.  (1966), 
forearm  measurements  of  a  male  and  female  from  Chiapas;  Villa-R.  (1967),  external 
measurements  of  one  male  and  one  female  from  Yucatan  and  one  male  from  Oaxaca,  and 
cranial  measurements  of  the  male  and  female  from  Yucatan;  Goodwin  (1969),  forearm 
and  cranial  measurements  of  five  males  and  five  females  from  Oaxaca;  Gardner  et  al. 
(1970),  external  and  cranial  measurements  of  one  male  from  Costa  Rica;  Valdez  and 
LaVal  (1971),  external  and  cranial  measurements  of  one  female  and  the  mean  of  two  males 
from  Honduras;  Marinkelle  and  Cadena  (1972),  forearm  measurements  of  one  male  from 
Colombia. 

Geographic  variation. — According  to  Gardner  et  al.  (1970),  their  male  from  Costa  Rica 
closely  resembled  a  male  from  Chiapas  in  cranial  measurements. 

Mimon  crenulatum  (E.  Geoffroy  St. -Hilaire,  1810) 

Measurements  of  Mimon  crenulatum  have  been  recorded  as  follows:  Peters  (1866a), 
external  measurements  of  a  specimen  from  Brazil;  Dobson  (1878a),  external  measure¬ 
ments  of  one  (A/,  longifolium)  from  Brazil,  and  a  specimen  from  an  unknown  locality; 
Thomas  (1903c),  external  and  cranial  measurements  of  the  male  holotype  of  M.  c.  picatum 
from  Brazil;  Cunha  Vieira  (1942),  external  and  cranial  measurements  of  two  specimens 
from  Brazil;  Sanborn  (19496),  forearm  and  cranial  measurements  of  two  males  from  Peru; 
Handley  (1960),  external  and  cranial  measurements  of  five  males  and  four  females  from 
Brazil,  Trinidad,  Venezuela,  Panama,  and  Ecuador  (including  the  holotype  of  M.  c.  keenani)', 
Goodwin  and  Greenhall  (1961),  external  and  cranial  measurements  of  a  male  from  Trinidad; 
Husson  (1962),  external  and  cranial  measurements  of  two  males  from  Surinam;  Hill  (1964), 
forearm  of  two  males  and  females  and  cranial  measurements  of  one  male  from  Guyana; 
Jones  (1964),  external  and  cranial  measurements  of  a  female  from  Campeche  and  measure¬ 
ments  available  from  the  holotype  of  M.  c.  keenani  from  Panama;  Gardner  et  al.  (1970), 
external  and  cranial  measurements  (mean,  range)  of  four  specimens  (two  males  and 
females)  from  Costa  Rica;  Gardner  and  Patton  (1972),  forearm  and  cranial  measurements 
(mean,  range)  of  four  males  and  three  females  from  Peru. 

Miimon  koepckeae  Gardner  and  Patton,  1972 

Gardner  and  Patton  (1972)  recorded  external  and  cranial  measurements  (mean,  range) 
of  two  males  and  one  female  and  the  measurements  of  the  female  holotype  from  Peru. 

Phylloderma  stenops  Peters,  1865 

Measurements  of  Phylloderma  stenops  have  been  recorded  as  follows:  Peters  (18666), 
external  measurements  of  one  specimen  from  Cayenne;  Dobson  (1878a),  external  measure¬ 
ments  of  Guandira  cayanensis  from  Cayenne;  Goodwin  (1940,  1946,  1953),  external  and 
cranial  measurements  of  the  female  holotype  of  P.  stenops  septentrionalis  from  Honduras; 
Goodwin  (1942a),  external  and  cranial  measurements  of  two  specimens  from  Honduras; 
Hall  and  Kelson  (1959),  external  and  cranial  measurements  of  the  P.  septentrionalis  holotype 
and  one  female;  Husson  (1962),  external  and  cranial  measurements  of  the  male  holotype 
from  Cayenne;  Hill  (1964),  external  and  cranial  measurements  of  three  females  from 
Guyana,  one  male  from  Brazil,  and  of  the  holotype  of  Guandira  cayanensis  (=  P.  stenops ); 
Carter  et  al.  (1966),  external  and  cranial  measurements  of  a  male  from  Chiapas;  Gardner 
(1976),  external  and  cranial  measurements  of  a  female  from  Peru;  LaVal  (1977),  forearm 
length  and  weight  of  a  female  from  Costa  Rica. 

Phyllostomus  discolor  (Wagner,  1 843) 

Measurements  of  Phyllostomus  discolor  have  been  recorded  as  follows:  Peters  (18656) 
external  measurements  of  one  specimen  from  Brazil;  Dobson  (1878a),  external  measure- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


23 


merits  of  one  specimen;  Elliot  ( 1 905 A;  1917),  external  and  cranial  measurements  of  the 
holotype  of  P.  verrucossum  from  Oaxaca;  Miller  (1932),  forearm  (range  of  five  specimens) 
and  cranial  measurements  of  a  specimen  from  Barro  Colorado  Island,  Canal  Zone;  Sanborn 
(1936),  forearm  and  condylobasal  length  of  skull  measurements  (range)  of  specimens  from 
Brazil  (discolor),  and  from  Oaxaca,  Veracruz,  and  Guatemala  ( verruscosus)\  Cunha  Vieira 
(1942),  external  measurements  of  a  male  from  Brazil  and  female  from  an  unknown  locality; 
Goodwin  (1942c/),  external  and  cranial  measurements  of  two  males  from  Honduras;  Goodwin 
(1946),  cranial  measurements  of  two  males  from  Honduras;  Dalquest  (1951),  external 
and  cranial  measurements  of  two  males  and  one  female  from  Trinidad;  Felten  (1956a), 
external  measurements  (mean,  range)  of  185  males  and  217  females,  and  cranial  measure¬ 
ments  (mean,  range)  of  35  males  and  39  females  from  El  Salvador;  Burt  and  Stirton  (1961), 
forearm  and  cranial  measurements  (range)  of  15  males  and  12  females  from  El  Salvador; 
Goodwin  and  Greenhall  (1961),  forearm  measurements  (range)  of  four  specimens 
(two  males  and  females)  and  cranial  measurements  of  one  female  from  Trinidad;  Davis 
and  Carter  (1962a),  forearm  and  cranial  measurements  of  one  male  from  Costa  Rica;  Husson 
(1962),  external  and  cranial  measurements  of  eight  males  and  two  females  from  Surinam; 
Valdivieso  and  Tamsitt  (1962),  external  measurements  (range)  of  five  males  and  three 
females  and  cranial  measurements  of  two  specimens  from  Colombia;  Tamsitt  and  Valdivieso 
(1963a),  external  measurements  (mean,  range)  of  11  specimens  (seven  males,  four  females) 
and  cranial  measurements  of  one  male  and  female  from  Colombia;  Pirlot  (1967),  external 
measurements  of  two  specimens;  Villa-R.  (1967),  external  measurements  of  13  specimens 
(mean,  sd,  range)  and  cranial  measurements  (mean,  sd,  range)  of  14  specimens  from 
Mexico;  Goodwin  (1969),  forearm  and  cranial  measurements  of  six  males  and  three  females 
from  Oaxaca;  Power  and  Tamsitt  (1973),  forearm  and  cranial  measurements  (means)  of 
males  and  females  from  various  localities  in  southern  Mexico  to  South  America;  Smith  and 
Genoways  (1974),  external  and  cranial  measurements  of  four  females  (mean,  range)  and 
two  males  (means)  from  Margarita  Island,  Venezuela;  Taddei  (1975a),  external  (30  males, 
30  females)  and  cranial  measurements  (mean,  sd,  range)  of  15  males  and  females 
from  Brazil;  Gardner  (1976),  external  and  cranial  measurements  of  a  male  from  Peru. 

Individual  variation. — Taddei  (1975a)  reported  coefficient  of  variation  values  for  external 
measurements  of  Brazilian  specimens  to  vary  from  2.38  to  6.51,  whereas  CVs  for  cranial 
measurements  varied  from  0.96  to  4.45. 

Secondary  sexual  variation. — Taddei  (1975a)  found  females  averaged  larger  than  males 
in  17  external  measurements  and  significantly  so  in  three  of  these,  length  of  ear,  digit 
Ill-phalanx  2,  digit  V-phalanx  2.  Males  averaged  larger  than  females  in  15  cranial  measure¬ 
ments  and  significantly  so  in  five  of  these,  breadth  across  canines,  breadth  across  molars, 
zygomatic  width,  mastoid  breadth,  cranial  depth.  Power  and  Tamsitt  (1973),  performing 
a  manova,  showed  that  males  were  significantly  bigger  than  females,  and  a  subsequent 
discriminant  function  analysis  revealed  that  mastoid  width  and  zygomatic  width  contri¬ 
buted  greatly  to  the  separation  of  the  sexes. 

Geographic  variation. — In  forearm  and  condylobasal  length  of  skull,  specimens  from 
Barro  Colorado  Island,  Canal  Zone,  were  somewhat  greater  in  size  than  three  topotypes 
of  P.  discolor  from  southern  Mexico  (Miller,  1932).  Dalquest  (1951),  comparing  cranial 
measurements  of  Trinidad  specimens  with  those  from  Venezuela,  found  no  difference, 
whereas  forearm  length  appeared  to  be  slightly  less  than  in  specimens  from  the  mainland. 
Davis  and  Carter  (1962a)  stated  that  the  measurements  considered  to  that  time  as  an 
expression  of  geographic  variation  were  in  reality  due  to  individual  variation.  According 
to  Husson  (1962),  external  and  cranial  measurements  of  Surinam  specimens  agree  well 
with  those  given  by  Sanborn  (1936),  Dalquest  (1951),  and  Goodwin  and  Greenhall  (1961) 
for  specimens  from  Trinidad  and  Venezuela.  When  comparing  these  data  with  those  from 
El  Salvador  (Felten,  1956a),  Husson  (1962)  concluded  that  the  cranial  measurements  were 
larger  in  the  specimens  from  El  Salvador.  Power  and  Tamsitt  (1973)  stated  that  populations 


24 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


west  of  the  Andes  in  southwestern  Ecuador,  those  near  or  within  the  Andes  mountains  in 
central  Colombia,  and  those  east  of  the  Andes  in  eastern  Colombia  were  quite  similar 
and  did  not  warrant  subspecific  recognition.  Smith  and  Genoways  (1974)  found  external 
and  cranial  measurements  of  specimens  from  Margarita  Island,  Venezuela,  comparable 
to  those  given  by  Sanborn  (1936)  for  specimens  from  Brazil,  Venezuela,  and  French  Guiana, 
and  by  Goodwin  and  Greenhall  (1961)  for  material  from  Trinidad. 

Phyllostomus  elongatus(E.  Geoffroy  St. -Hilaire,  1810) 

Measurements  for  Phyllostomus  elongatus  have  been  recorded  as  follows:  Peters  (18656), 
external  measurements  of  a  specimen  from  Brazil;  Dobson  ( 1 878z/),  external  measurements 
of  one  specimen;  Sanborn  (1936),  forearm  and  cranial  measurements  of  a  female  from 
Ecuador;  Cunha  Vieira  (1942),  external  measurements  of  three  males  and  one  female  and 
cranial  measurements  of  one  male  from  Brazil;  Husson  (1962),  external  and  cranial 
measurements  of  four  males  and  two  females  from  Surinam;  Butterworth  and  Starrett 
(1964),  external  and  cranial  measurements  of  a  male  from  Venezuela;  Hill  (1964),  fore¬ 
arm  measurements  of  a  male  and  female  and  cranial  measurements  of  a  female  from  Guyana. 

Geographic  variation. — Measurements  of  six  specimens  from  Surinam  correspond  well 
to  those  given  by  Sanborn  (1951)  for  specimens  from  Peru,  and  by  Husson  (1962)  for 
material  from  Guyana. 


Phyllostomus  hastatus (Pallas,  1767) 

Measurements  for  Phyllostomus  hastatus  have  been  recorded  as  follows:  Dobson 
(1878a),  external  measurements  of  one  specimen;  Flower  and  Lydekker  (1891),  forearm 
length  of  the  species;  Jentink  (1893),  forearm  length  of  a  male  from  Guyana;  Robinson 
and  Lyon  (1901),  external  measurements  of  five  males  and  eight  females  from  Venezuela; 
J.  A.  Allen  (1904),  external  and  cranial  measurements  (range)  of  two  males  and  four 
females  (including  the  female  holotype  of  P.  h.  panamensis)  from  Chiriqui,  Panama, 
external  and  cranial  measurements  of  the  male  holotype  of  P.  h.  caurae  from  Colombia, 
and  cranial  measurements  (mean,  range)  of  two  specimens  from  Trinidad  and  four  from 
eastern  Venezuela;  Elliot  (1904),  external  and  cranial  measurements  of  one  specimen; 
G.  M.  Allen  (1908),  external  measurements  of  three  and  cranial  measurements  of  one 
specimen  from  Brazil,  and  external  measurements  of  five  specimens  from  Costa  Rica; 
Miller  (1912),  external  and  cranial  measurements  of  a  male  from  Panama;  Cabrera  (1917), 
external  and  cranial  measurements  of  the  male  holotype  of  P.  h.  curaca  and  the  range  of 
some  of  these  measurements  in  three  females  from  Ecuador;  Lima  (1926),  external  measure¬ 
ments  of  a  male  from  Brazil;  Cunha  Vieira  (1942),  external  measurements  of  eight  males 
and  three  females  and  cranial  measurements  of  three  males  from  Brazil;  Dalquest  (1951), 
forearm  and  cranial  measurements  (mean)  of  four  specimens  from  Trinidad;  Goodwin 
(1953),  forearm  and  cranial  measurements  of  the  female  holotype  of  P.  h.  panamensis 
from  Panama  and  of  the  holotype  of  P.  h.  caucae  from  Colombia;  Hall  and  Kelson  (1959), 
external  and  cranial  measurements  of  a  male  and  female  from  Costa  Rica;  Goodwin  and 
Greenhall  (1961),  forearm  measurements  (range)  of  five  specimens  (two  males,  three 
females)  and  cranial  measurements  of  one  female  from  Trinidad;  Husson  (1962),  external 
and  cranial  measurements  of  eight  males  and  two  females  from  Surinam;  Taddei  (1975a), 
external  measurements  (mean,  sd,  range)  of  20  males  and  20  females  and  cranial  measure¬ 
ments  (mean,  sd,  range)  of  15  males  and  15  females  from  Brazil. 

Individual  variation. — Taddei  (1975a)  gave  CV  values  for  external  measurements  from 
1.28  to  6.04  and  for  cranial  measurements  from  1.06  to  2.84. 

Secondary  sexual  variation. — In  all  of  the  15  cranial  measurements  taken  by  Taddei 
(1975a),  males  proved  to  be  significantly  larger  than  females,  this  was  also  the  case  in  eight 
of  the  17  external  measurements. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


25 


Geographic  variation. — According  to  J.  A.  Allen  (1904),  specimens  from  Chiriqui, 
Panama,  were  much  larger  than  those  from  Trinidad  and  eastern  Venezuela.  Specimens 
from  Costa  Rica  seemed  to  correspond  fairly  well  with  the  holotype  of  P.  h.  panamensis 
from  Chiriqui  (G.  M.  Allen,  1908). 

Phyllostomus  latifolius  Thomas,  1901 

Measurements  for  Phyllostomus  latifolius  have  been  recorded  as  follows:  Thomas 
(1901  h),  forearm  and  cranial  measurements  of  the  male  holotype  and  external  measure¬ 
ments  of  a  second  male  from  Guyana;  Husson  (1962),  external  and  cranial  measurements 
of  six  paratypes  (four  males,  two  females)  from  Guyana;  Marinkelle  and  Cadena  (1972), 
forearm  and  cranial  measurements  (means)  of  five  females  from  Colombia. 

Tonatia  bidens(Spix,  1823) 

Measurements  for  Tonatia  bidens  have  been  recorded  as  follows:  Dobson  (1878a), 
external  measurements  of  one  specimen  from  Brazil;  Lima  (1926),  external  measure¬ 
ments  of  a  specimen  from  Brazil;  Sanborn  (1936),  external  measurements  (range)  of  three 
males  and  cranial  measurements  of  two  males  from  Brazil;  Cunha  Vieira  (1942),  external 
and  cranial  measurements  of  a  female  from  Brazil;  Goodwin  (19426),  external  and  cranial 
measurements  (range)  of  one  male  and  five  females  from  the  Amazon  basin,  one  male  from 
Venezuela,  and  two  males  and  six  females  from  Costa  Rica;  Goodwin  (1946);  forearm  and 
cranial  measurements  of  a  male  and  female  from  Costa  Rica;  Koopman  and  Williams 
(1951),  cranial  measurements  of  the  holotype  and  paratype  of  Tonatia  bidens  saurophila 
from  Jamaica  and  of  one  specimen  of  T.  b.  bidens  from  Costa  Rica  and  another  from 
Guyana;  Goodwin  (1953),  one  cranial  measurement  of  the  holotype  of  T.  b.  saurophila 
from  Jamaica;  Hall  and  Kelson  (1959),  forearm  and  cranial  measurements  of  a  male  and 
female  from  Costa  Rica;  Goodwin  and  Greenhall  (1961),  forearm  and  cranial  measurements 
of  one  male  and  one  female  from  Trinidad;  Hill  (1964),  forearm  measurements  of  one  male 
and  two  females  and  cranial  measurements  of  one  female  from  Guyana;  Carter  et  al.  (1966), 
external  and  cranial  measurements  of  a  female  from  Guatemala;  Pirlot  (1967),  external 
measurements  of  one  specimen;  Gardner  et  al.  (1970),  forearm  and  cranial  measurements 
of  a  female  from  Costa  Rica;  Valdez  and  LaVal  (1971),  external  and  cranial  measurements 
of  one  male  and  four  females  (mean,  range)  from  Honduras;  Gardner  (1976),  external 
and  cranial  measurements  (mean,  range)  of  seven  specimens  from  Peru. 

Tonatia  brasiliense  (Peters,  1866) 

Measurements  for  Tonatia  brasiliense  have  been  recorded  as  follows:  Peters  (18666), 
external  measurements  of  the  holotype  from  Brazil;  Dobson  (1878«),  external  measure¬ 
ments  of  the  holotype  from  Brazil;  Cunha  Vieira  (1942),  external  measurements  based  on 
Peters  (18666);  Goodwin  (19426),  external  and  cranial  measurements  of  one  male  and 
one  female  from  Brazil  and  Peters’  measurements  of  the  holotype;  Goodwin  and  Green- 
hall  (1961:236),  forearm  and  cranial  measurements  of  the  holotype;  Gardner  (1976), 
external  and  cranial  measurements  of  two  males  from  Peru. 

Tonatia  carrikeri  (J.  A.  Allen,  1910) 

Measurements  for  Tonatia  carrikeri  have  been  recorded  as  follows:  J.  A.  Allen  (1910), 
external  measurements  for  the  male  holotype  and  five  females  and  cranial  measurements 
of  the  holotype  from  Venezuela;  Goodwin  (19426),  external  and  cranial  measurements 
of  one  male  and  one  female  from  Venezuela;  Goodwin  (1953),  external  and  cranial  measure¬ 
ments  of  the  holotype  from  Venezuela;  Husson  (1962),  external  and  cranial  measurements 
of  a  male  from  Surinam;  Gardner  (1976),  external  and  cranial  measurements  of  two  females 
from  Peru. 


26 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Geographic  variation. — Husson  (1962)  noted  that  a  male  from  Surinam  was  smaller 
than  one  reported  by  Goodwin  (19426)  from  Venezuela  and  that  it  compared  more  favorably 
with  a  female  from  Venezuela. 


Tonatia  minuta  Goodwin,  1942 

Measurements  of  Tonatia  minuta  have  been  recorded  as  follows:  Goodwin  (19426), 
external  and  cranial  measurements  of  the  female  holotype  of  T.  nicaraguae  from  Nicaragua, 
and  the  male  holotype  of  T.  minuta  and  two  females  from  Ecuador;  Goodwin  (1946), 
forearm  and  cranial  measurements  of  the  holotype  of  T.  nicaraguae',  Goodwin  (1953), 
external  and  cranial  measurements  of  the  holotype  of  T.  minuta  and  T.  nicaraguae'.  Hall 
and  Kelson  (1959),  forearm  and  cranial  measurements  of  the  holotype  of  T.  nicaraguae 
and  one  female;  Goodwin  and  Greenhall  (1961),  forearm  and  cranial  measurements  of  a 
male,  female,  and  juvenile  from  Trinidad  and  the  holotype  of  T.  minuta',  Davis  and  Carter 
(1962 a),  external  and  cranial  measurements  of  a  male  and  the  female  holotype  of  T. 
nicaraguae  from  Nicaragua;  Davis  et  al.  (1964),  external  and  cranial  measurements  of 
one  female  from  Panama;  LaVal  (1969),  external  and  cranial  measurements  of  one  male 
and  the  mean  of  two  females  from  Honduras;  Gardner  et  al.  (1970),  forearm  and  cranial 
measurements  of  five  males  (mean,  range)  from  Costa  Rica;  Jones  et  al.  (19716),  external 
and  cranial  measurements  of  two  males  from  Nicaragua;  Ojasti  and  Naranjo  (1974), 
external  and  cranial  measurements  of  one  male  from  Venezuela. 

Geographic  variation. — LaVal  (1969)  noted  that  the  three  specimens  (one  male,  two 
females)  he  measured  from  Honduras  were  notably  larger  in  some  measurements  (fore¬ 
arm,  third  metacarpal,  length  of  skull)  than  those  reported  by  Davis  and  Carter  (1962a) 
and  Davis  et  al.  (1964).  According  to  Gardner  et  al.  (1970),  specimens  from  Costa  Rica 
were  smaller  than  those  reported  from  Honduras  by  LaVal  (1969)  but  similar  in  size 
to  those  reported  by  Davis  and  Carter  (1962a)  and  Davis  et  al.  (1964)  from  Nicaragua  and 
Panama.  Jones  et  al.  (19716)  concluded  that  their  specimens  from  Nicaragua  resembled 
material  reported  from  Nicaragua  by  LaVal  (1969)  and  averaged  larger  than  other  published 
measurements  (Goodwin,  19426;  Davis  and  Carter,  1962a;  Davis  et  al.,  1964;  Gardner 
et  al.,  1970).  A  male  collected  in  Venezuela  was,  according  to  Ojasti  and  Naranjo  (1974), 
slightly  larger  than  the  average  size  reported  from  Eucador  (Goodwin  19426),  Honduras 
(LaVal,  1969),  Costa  Rica  (Gardner  et  al.,  1970),  and  Nicaragua  (Jones  et  al.,  19716). 

Tonatia  silvicola  (D'Orbigny,  1836) 

Measurements  of  Tonatia  silvicola  have  been  recorded  as  follows:  Peters  (18656), 
external  measurements  of  a  specimen  from  Brazil;  Dobson  (1878a),  external  measure¬ 
ments  of  one  specimen  from  Brazil;  Elliot  (1904),  external  and  cranial  measurements  of 
one  specimen;  Thomas  (1910),  external  and  cranial  measurements  of  the  holotype  of 
T.  s.  laephotis;  Cabrera  (1917),  external  measurements  of  a  male  and  a  female  (T.  amblyotis) 
from  Ecuador;  Sanborn  (1936),  external  and  cranial  measurements  (range)  of  specimens 
from  Ecuador;  Sanborn  (1941),  forearm  and  cranial  measurements  of  one  female  from 
Peru,  one  specimen  from  British  Honduras,  four  specimens  from  Bolivia,  and  the  range 
of  measurements  of  a  series  from  Ecuador;  Cunha  Vieira  (1942),  external  and  cranial 
measurements  of  a  male  from  Brazil;  Goodwin  (1942a),  forearm  and  cranial  measurements 
(range)  of  the  species  T.  amblyotis  (=7.  silvicola);  Goodwin  (19426),  external  and 
cranial  measurements  (range)  of  T.  amblyotis  from  Bolivia,  Ecuador,  Colombia,  and 
Panama  and  cranial  measurements  of  one  specimen  from  British  Honduras,  and  for  T. 
laephotis,  external  measurements  of  one  male  and  one  female  from  the  lower  Amazon, 
and  range  of  cranial  measurements  of  16  specimens  from  Brazil;  Goodwin  (1946), 
external  and  cranial  measurements  (range)  of  the  species;  Goodwin  (1953),  external  and 
cranial  measurements  of  the  holotype  of  Chrotopterus  columbianus  ( =  T.  silvicola) 
from  Colombia;  Husson  (1962),  external  and  cranial  measurements  of  one  male  and  two 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


27 


females  from  Surinam;  Hill  (1964),  forearm  measurements  of  two  males  and  females 
and  cranial  measurements  of  one  female  from  Guyana;  Jones  (1964),  external  and 
cranial  measurements  of  a  male  from  Campeche;  Carter  et  al.  (1966),  external  and  cranial 
measurements  of  a  female  from  Guatemala;  Villa-R.  (1967),  external  and  cranial  measure¬ 
ments  (range)  of  T.  s.  silvicola  from  Mexico;  Villa-R.  and  Villa  Cornejo  (1969),  external 
measurements  of  one  specimen  from  Argentina;  Jones  et  al.  (1973),  forearm  and  cranial 
measurements  of  a  male  from  Campeche. 

Geographic  variation. — According  to  Carter  et  al.  (1966),  measurements  of  a  female 
from  Guatemala  approximated  those  given  by  Goodwin  (19426)  for  South  American 
specimens  but  were  slightly  larger  than  those  for  a  British  Honduran  specimen  examined 
by  Goodwin.  Sanborn  (1941)  noted  that  forearm  and  total  length  of  skull  of  a  specimen 
from  British  Honduras  were  small  for  the  species. 

Tonatia  venezuelae  (Robinson  and  Lyon,  1901) 

Measurements  of  Tonatia  venezuelae  have  been  recorded  as  follows:  Robinson  and 
Lyon  (1901),  external  measurements  for  the  male  holotype  and  two  additional  males  from 
Venezuela  and  cranial  measurements  of  the  holotype;  Sanborn  (1941),  forearm  measure¬ 
ments  (range)  in  the  original  series;  Goodwin  (19426),  external  and  cranial  measurements 
of  a  male  and  female  from  Venezuela  (including  cranial  measurements  of  the  holotype 
from  Venezuela);  Goodwin  and  Greenhall  (1961:236),  forearm  and  cranial  measurements 
of  a  paratype;  Ojasti  and  Naranjo  (1974),  external  and  cranial  measurements  of  one  specimen 
from  Venezuela. 


Trachops  cirrhosus(Spix,  1823) 

Measurements  of  Trachops  cirrhosus  have  been  recorded  as  follows:  Saussure  (1860c), 
external  measurements  of  one  specimen  of  Tylostoma  mexicana  (  =  T.  cirrhosus)',  Peters 
(1865c),  external  measurements  of  a  specimen  from  Brazil;  Dobson  (1878u),  external 
measurements  of  one  female  from  Bermuda;  Elliot  (1904),  external  measurements  of  one 
specimen;  Goldman  (1925),  external  and  cranial  measurements  of  the  female  holotype 
of  T.  cirrhosus  coffin i  from  Guatemala;  Lima  (1926),  external  measurements  of  a  male 
from  Brazil;  Cunha  Vieira  (1942),  external  measurements  of  three  males  and  three  females 
and  cranial  measurements  of  two  females  from  Brazil;  Goodwin  (1942u),  external  and 
cranial  measurements  of  two  females  from  Honduras  and  the  holotype  of  T.  c.  coffin i 
from  Guatemala;  Goodwin  (1946),  forearm  and  cranial  measurements  of  one  male  from 
Colombia;  Herskovitz  (1949),  external  and  cranial  measurements  (range)  of  20  specimens 
(eight  males,  nine  females,  three  unsexed)  from  northern  Colombia;  Felten  (1956c/),  exter¬ 
nal  and  cranial  measurements  of  a  male  from  El  Salvador;  Felten  (19566),  forearm  and 
cranial  measurements  of  the  female  holotype  and  two  paratypes  (a  male  and  female)  of 
T.  c.  ehrhardti  from  Brazil,  and  range  of  these  measurements  in  two  other  subspecies, 
coffini  (Guatemala,  Honduras,  El  Salvador)  and  cirrhosus  (Colombia);  Burt  and  Stirton 
(1961),  forearm  and  cranial  measurements  (range)  of  five  males  and  17  females  from  El 
Salvador;  Goodwin  and  Greenhall  (1961),  forearm  measurements  (range)  of  two  males  and 
one  female  and  cranial  measurements  of  one  male  and  one  female  from  Trinidad;  Davis 
and  Carter  (1962 a),  forearm  and  cranial  measurements  of  a  female  from  Costa  Rica;  Husson 
(1962),  external  and  cranial  measurements  of  one  male  from  Surinam;  Villa-R.  (1967),  ex¬ 
ternal  and  cranial  measurements  of  five  specimens  from  Mexico;  Starrett  and  Casebeer 
(1968),  forearm  and  cranial  measurements  of  two  females  and  means  and  ranges  of  four 
males  from  Costa  Rica;  Goodwin  (1969),  forearm  and  cranial  measurements  of  four  males 
and  two  females  from  Oaxaca. 

Geographic  variation. — Husson  (1962),  comparing  external  measurements  of  one  male 
from  Surinam  with  20  specimens  from  Colombia  (Hershkovitz,  1949),  concluded  that  the 
Surinam  specimen  was  large.  The  skull  measurements,  however,  did  not  differ  markedly. 


28 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Davis  and  Carter  (1962a)  found  measurements  of  their  one  female  from  Costa  Rica  within 
the  range  of  variation  reported  in  this  species  from  Colombia  (Hershkovitz,  1949).  These 
authors  also  concluded  that  other  published  measurements  (Goldman,  1925;  Felten,  1956c/) 
fell  within  the  range  of  the  Colombian  series  (Hershkovitz,  1949). 

Vampyrum  spectrum  (Linnaeus,  1758) 

Measurements  of  Vampyrum  spectrum  have  been  recorded  as  follows:  Dobson  (1878a), 
external  measurements  of  one  specimen;  Flower  and  Lydekker  (1891),  forearm  length  for 
the  species;  Elliot  (1904),  external  and  cranial  measurements  of  a  specimen;  Goldman 
(1917)  and  Goodwin  (1942a),  external  and  cranial  measurements  of  the  male  holotype 
of  V.  s.  nelson i  from  Veracruz;  Sanborn  (1941),  external  and  cranial  measurements  of  one 
female  from  Trinidad;  Cunha  Vieira  (1942),  external  measurements  from  Dobson  (1878a); 
Goodwin  (1946),  external  and  cranial  measurements  of  one  male  from  Nicaragua  and  of 
the  holotype  of  V.  s.  nelsonr,  Hall  and  Kelson  (1959),  forearm  and  cranial  measurements 
of  the  holotype  of  V.  s.  nelsonr,  Goodwin  and  Greenhall  (1961),  forearm  measurements 
(one  male,  one  female)  and  cranial  measurements  (one  male)  from  Trinidad;  Husson 
(1962),  external  and  cranial  measurements  of  three  males,  two  females,  and  two  unsexed 
specimens  from  Surinam,  one  male  and  one  female  from  Cayenne,  and  one  male  from 
Guyana;  Casebeer  et  al.  (1963),  external  and  cranial  measurements  of  a  male  from  Costa 
Rica;  Hall  and  Dalquest  (1963),  external  and  cranial  measurements  of  the  holotype  from 
Veracruz;  Goodwin  (1969),  forearm  and  cranial  measurements  for  two  males,  one  from 
Veracruz  the  other  from  Nicaragua;  Peterson  and  Kirmse  (1969),  external  and  cranial 
measurements  of  a  female  from  Panama;  Gardner  et  al.  (1970),  external  and  cranial  measure¬ 
ments  of  one  female  from  Costa  Rica. 

Geographic  variation. — Casebeer  et  al.  (1963)  stated  that  their  measurements  corre¬ 
sponded  closely  with  those  given  by  Goldman  (1917)  for  the  male  holotype  of  V.  spectrum 
nelsoni  from  Veracruz  and  were  slightly  smaller  than  measurements  of  specimens  from 
Trinidad  (Goodwin  and  Greenhall,  1961).  Peterson  and  Kirmse  (1969),  comparing  their 
female  specimens  from  Panama  with  those  reported  by  Husson  (1962)  from  the  Guianas, 
found  their  specimen  actually  larger  in  most  measurements  than  the  mean  of  specimens 
from  near  the  type  locality  (Surinam). 

Subfamily  Glossophaginae 
Anoura  brev irostrum  Carter,  1968 

Measurements  of  Anoura  brevirostrum  have  been  recorded  as  follows:  Carter  (1968), 
external  and  cranial  measurements  of  the  female  holotype  from  Peru  and  (mean  and  range) 
of  five  specimens  (one  male,  four  females)  from  Peru;  Gardner  (1976),  external  and  cranial 
measurements  of  a  male  from  Peru. 

Anoura  caudifer(E.  Geoffry  St.-Hilaire,  1818) 

Measurements  of  Anoura  caudifer  have  been  recorded  as  follows:  Saussure  (1860c), 
external  measurements  of  one  specimen  of  A.  ecaudata  ( =  A .  caudifer );  Peters  (1869), 
external  measurements  of  the  holotype  of  Anoura  wiedii  from  Brazil;  Dobson  (1878a), 
external  measurements  of  one  specimen;  Lonnberg  (1921),  external  and  cranial  measure¬ 
ments  of  a  male  from  Ecuador  in  the  original  description  of  A.  c.  aequatoris-,  Lima  (1926), 
external  measurements  of  a  specimen  of  Lonchoglossa  ecaudata  ( A .  caudifer)  from  Brazil; 
Sanborn  (1933),  forearm  and  cranial  measurements  (range)  of  11  specimens  from  Brazil; 
Sanborn  (1938),  external  measurements  of  two  specimens  and  cranial  measurements  of  one 
specimen  from  Venezuela;  Sanborn  (1941),  forearm  measurements  (range)  of  two  males 
from  Venezuela  and  one  male  and  four  females  from  Brazil  combined,  and  the  forearm 
measurement  of  one  male  from  Peru;  Cunha  Vieira  (1942),  external  measurements  of  five 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


29 


males  and  two  females  and  cranial  measurements  of  two  males  and  two  females  from 
Brazil;  Hershkovitz  (1949),  external  and  cranial  measurements  (range)  of  four  males  and 
one  female  combined,  and  these  measurements  for  one  young  adult  from  Colombia;  Husson 
(1962),  external  and  cranial  measurements  of  a  female  from  Surinam;  Tamsitt  and 
Valdivieso  (19666),  external  measurements  of  a  male  and  female,  cranial  measurements  of 
a  male  from  Colombia,  and  mean,  sd,  se,  and  range  in  measurements  of  specimens  from 
Andean  and  Amazonian  populations;  Taddei  (19756),  external  measurements  of  40  males 
and  40  females  and  cranial  measurements  of  15  males  and  15  females  (mean,  se,  range) 
from  Brazil. 

Individual  variation. — In  specimens  from  Brazil,  coefficients  of  variation  for  external 
measurements  varied  in  40  males  from  2.64  to  5.88  and  in  40  females  from  2.09  to  7.44; 
for  cranial  measurements  in  15  males,  CV  values  were  from  1.37  to  4.27  and  in  15  females 
from  1.22  to  3.17  (Taddei,  19756). 

Secondary  sexual  variation. — In  material  from  Brazil,  17  external  measurements  showed 
no  secondary  sexual  differences.  However,  in  three  (breadth  across  canines,  zygomatic 
breadth,  mastoid  breadth)  of  15  cranial  measurements,  males  proved  to  be  significantly 
larger  than  females  (Taddei,  19756). 

Geographic  variation. — Tamsitt  and  Valdivieso  (19666)  found  specimens  from  an  Andean 
population  to  be  generally  larger  in  external  measurements  than  those  from  an  Amazonian 
population — forearm  measurements  proved  to  be  significantly  different.  Cranial 
measurements  were  similar  between  the  two  populations  and  no  geographic  trend  was 
obvious. 


Anoura  cultrata  Handley,  1960 

Measurements  of  Anoura  cultrata  have  been  recorded  as  follows:  Handley  (1960), 
external  and  cranial  measurements  of  the  female  holotype  from  Panama;  Carter  et  al. 
(1966),  external  and  cranial  measurements  of  a  male  from  Costa  Rica;  Carter  (1968), 
external  and  cranial  measurements  (mean,  range)  of  15  specimens  from  Panama  and 
Costa  Rica;  Gardner  et  al.  (1970),  forearm  and  cranial  measurements  (mean,  range)  of 
five  specimens  (four  males,  one  female)  from  Costa  Rica;  LaVal  (1977),  forearm  length 
and  weight  of  a  specimen  from  Costa  Rica. 

Anoura  geoffroyi  Gray,  1838 

Measurements  of  Anoura  geoffroyi  have  been  recorded  as  follows:  Peters  (1868),  external 
measurements  of  the  holotype  of  A.  g.  lasiopyga  from  Mexico;  Dobson  (1878u),  external 
measurements  of  the  holotype  of  Lonchoglossa  wiedii  from  Brazil,  external  measure¬ 
ments  of  the  holotype  of  A.  geoffroyi,  and  those  of  an  immature  specimen;  Elliot  (1904), 
external  and  cranial  measurements  of  one  specimen;  Anthony  (1921),  external  and  cranial 
measurements  of  the  female  holotype  of  A.  g.  antricola  from  Ecuador;  Lima  (1926), 
external  measurements  of  a  male  from  Brazil;  Sanborn  (1933),  external  and  cranial  measure¬ 
ments  (range)  of  specimens  from  Veracruz,  Tlaxcala,  Jalisco,  and  El  Salvador;  Goodwin 
(1934),  external  measurements  of  one  specimen  from  Guatemala;  Sanborn  (1936),  fore¬ 
arm  and  cranial  measurements  (range)  of  1 1  males  and  two  females  from  Guatemala; 
Cunha  Vieira  (1942),  external  measurements  of  a  male  and  three  females  and  cranial 
measurements  of  a  male  from  Brazil;  Goodwin  (1942 a),  external  and  cranial  measurements 
of  one  specimen;  Goodwin  (1953),  external  and  cranial  measurements  of  the  female  holotype 
of  A.  g.  antricola  and  the  holotype  of  Glossophaga  apolinari  from  Colombia;  Sanborn 
(1954),  forearm  measurements  of  one  male  and  one  female  from  Venezuela;  Felten 
(1956a),  external  measurements  of  five  males  and  eight  females  (mean  and  range),  and 
cranial  measurements  of  two  males  and  one  female  from  El  Salvador;  Anderson  (1957), 
external  and  cranial  measurements  (mean,  sd,  range)  of  58  males  and  42  females  from 


30 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Chiapas  and  of  one  specimen  from  Costa  Rica;  Baker  (1960),  external  and  cranial  measure¬ 
ments  of  one  male  from  Durango;  Burt  and  Stirton  (1961),  forearm  and  cranial  measure¬ 
ments  of  a  specimen  from  El  Salvador;  Goodwin  and  Greenhall  (1961),  forearm  measure¬ 
ments  (range)  of  15  males  and  cranial  measurements  of  one  male  from  Trinidad;  Husson 
(1962),  external  and  cranial  measurements  of  six  males  from  Surinam  and  one  male  from 
Cayenne;  Valdivieso  (1964),  external  measurements  of  a  specimen  from  Colombia; 
Tamsitt  and  Valdivieso  (1966a),  forearm  and  cranial  measurements  of  one  female  from 
Colombia;  Villa-R.  (1967),  external  measurements  of  29  males  and  10  females  and  cranial 
measurements  of  28  males  and  10  females  (mean,  sd,  range)  from  Mexico;  Goodwin 
(1969),  forearm  and  cranial  measurements  of  three  males  and  four  females  from  Oaxaca; 
Spenrath  and  LaVal  (1970),  cranial  measurements  of  two  males  from  San  Luis  Potosf 
and  of  seven  males  (mean,  range)  from  Chiapas;  Matson  and  Patten  (1975),  forearm 
measurements  of  seven  males  (mean,  range)  and  two  females,  and  cranial  measurements 
of  five  males  (mean,  range)  and  two  females  from  Zacatecas. 

Secondary  sexual  variation. — Anderson  (1957)  found  no  significant  differences  in  both 
external  and  cranial  measurements  between  58  males  and  42  females  from  Chiapas. 

Geographic  variation. — Anderson  (1957)  found  a  significant  difference  in  forearm 
length  and  length  of  skull  between  specimens  from  South  America  and  Chiapas. 

Anoura  werckleae  Starrett,  1969 

Starrett  (1969)  recorded  external  and  cranial  measurements  of  the  male  holotype  and 
one  female  paratype  from  Costa  Rica. 

Choeroniscus  godmani  (Thomas,  1903) 

Measurements  of  Choeroniscus  godmani  have  been  recorded  as  follows:  Thomas 
(1903a),  external  and  cranial  measurements  of  the  male  holotype  from  Guatemala;  Elliot 
(1904),  external  and  cranial  measurements  of  one  specimen;  Goodwin  (1942a),  external 
and  cranial  measurements  of  the  holotype  from  Guatemala  and  a  male  from  Honduras; 
Goodwin  (1946),  external  and  cranial  measurements  of  one  male  and  female  from  Costa 
Rica;  Sanborn  (1954),  forearm  and  cranial  measurements  (range)  of  three  males  from 
Honduras,  and  two  males,  two  females,  and  one  unsexed  specimen  from  Costa  Rica  combined; 
Hall  and  Kelson  (1959),  external  and  cranial  measurements  of  one  male  and  two  females 
from  Costa  Rica;  Burt  and  Stirton  (1961),  forearm  and  cranial  measurements  of  one  male 
and  female  from  El  Salvador;  Gardner  (1962  ft),  external  and  cranial  measurements  of 
a  female  from  Nayarit;  Carter  et  al.  (1966),  external  and  cranial  measurements  of  one  female 
from  Veracruz  and  one  from  Guatemala;  Villa-R.  (1967),  external  and  cranial  measure¬ 
ments  of  one  female  from  Oaxaca;  Goodwin  (1969),  forearm  and  cranial  measurements 
of  two  males  (subadult)  and  one  female  from  Oaxaca;  LaVal  (1969),  forearm  and  cranial 
measurements  (mean,  range)  of  six  males  and  six  females  from  scattered  localities  in 
Mexico  and  Central  America;  Gardner  et  al.  (1970),  forearm  and  cranial  measurements 
of  one  male  and  three  females  from  Costa  Rica. 

Secondary  sexual  variation. — LaVal  (1969),  in  a  comparison  of  six  males  and  six 
females  from  scattered  localities  in  Mexico  and  Central  America,  found  females  to  be 
generally  larger  than  males.  He  found  no  overlap  in  greatest  skull  length  between  the  sexes. 
The  rostrum  was  larger  relative  to  the  braincase  in  skulls  from  females. 

Gardner  et  al.  (1970)  also  noted  in  a  collection  of  four  specimens  from  Costa  Rica, 
that  the  skull  of  the  one  male  was  considerably  shorter  than  those  of  the  three  females 
from  Costa  Rica. 

Sanborn  (1954)  stated,  contrary  to  the  above,  that  there  is  no  great  difference  in  size 
between  the  sexes. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


31 


Choeroniscus  inca  (Thomas,  1912) 

Measurements  of  Choeroniscus  inca  have  been  recorded  as  follows:  Thomas  (19126), 
external  and  cranial  measurements  of  the  male  holotype  from  Peru;  Sanborn  (1954), 
forearm  and  cranial  measurements  of  the  holotype  (after  Thomas),  external  measurements 
of  one  male  and  two  females,  and  cranial  measurements  of  one  male  and  three  females 
from  Venezuela. 


Choeroniscus  interinedius(J.  A.  Allen  and  Chapman,  1893) 

Measurements  of  Choeroniscus  intermedins  have  been  recorded  as  follows:  J.  A.  Allen 
and  Chapman  (1893),  external  measurements  of  the  female  holotype  and  two  males  from 
Trinidad;  Goodwin  (1953),  forearm  and  cranial  measurements  of  the  female  holotype  from 
Trinidad;  Sanborn  (1954),  forearm  and  cranial  measurements  of  the  holotype  as  given 
by  Goodwin  (1953),  forearm  measurement  of  the  holotype  as  in  the  original  description, 
and  forearm  length  of  an  additional  male  from  Trinidad;  Goodwin  and  Greenhall  (1961), 
external  and  cranial  measurements  of  the  female  holotype,  a  male,  and  a  female  from 
Trinidad;  Genoways  et  al.  (1973),  external  and  cranial  measurements  (mean,  se,  range) 
of  10  males  and  26  females  from  Trinidad. 

Individual  variation. — Coefficients  of  variation  in  external  measurements  ranged  from 
2.5  (total  length  for  males)  to  25.4  (length  of  tail  vertebrae  of  females).  CV  values  in  cranial 
measurements  ranged  from  1.9  (mastoid  breadth  for  females)  to  6.9  (postorbital  constriction 
for  males).  Females  showed  higher  coefficients  of  variation  than  males  in  external  measure¬ 
ments  and  lower  values  than  males  in  cranial  measurements  (Genoways  et  al.,  1973). 

Secondary  sexual  variation. — Females  proved  to  be  significantly  larger  than  males 
in  five  (greatest  length  of  skull,  condylobasal  length,  mastoid  breadth,  breadth  of  brain- 
case,  length  of  maxillary  toothrow)  of  12  measurements  tested.  In  two  of  the  other  seven 
measurements,  males  averaged  larger  than  females  and  in  one  they  were  equal  (Genoways 
etal. ,  1973). 


Choeroniscus  minor  (Peters,  1868) 

Measurements  of  Choeroniscus  minor  have  been  recorded  as  follows:  Peters  (1868), 
external  measurements  of  the  male  holotype  from  Surinam;  Dobson  (1878a),  external 
measurements  of  one  specimen  from  Surinam;  J.  A.  Allen  and  Chapman  (1893),  external 
measurements  as  given  by  Dobson  (1878a);  Elliot  (1904),  external  measurements  of  one 
specimen;  Lima  (1926),  external  measurements  of  a  male  from  Brazil;  Cunha  Vieira  (1942), 
external  and  cranial  measurements  of  a  female  from  Brazil;  Sanborn  (1954),  forearm 
measurements  of  three  specimens  from  Peru;  Husson  (1962),  external  and  cranial  measure¬ 
ments  of  the  male  holotype  from  Surinam;  Valdivieso  (1964),  external  and  cranial  measure¬ 
ments  of  one  female  from  Colombia. 

Choeroniscus  periosus  Handley,  1966 

Handley  (1966a)  recorded  external  and  cranial  measurements  of  the  female  holotype 
from  Colombia. 


Choeronycteris  mexicana  Tschudi,  1844 

Measurements  of  Choeronycteris  mexicana  have  been  recorded  as  follows:  Peters 
(1868),  external  measurements  of  one  specimen  from  Mexico;  Dobson  (1878a),  external 
measurements  of  a  single  specimen;  J.  A.  Allen  and  Chapman  (1893),  external  measure¬ 
ments  as  given  by  Dobson  (1878a);  Elliot  (1904),  external  measurements  of  one  specimen; 
Goodwin  (1934,  1942a,  1946),  external  measurements  of  a  specimen  from  Guatemala; 


32 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Dalquest  (1953a),  external  and  cranial  measurements  (mean)  of  four  males  from  San  Luis 
Potosi;  Baker  (1956),  external  and  cranial  measurements  (mean,  range)  of  three  males 
and  10  females  from  Coahuila;  Hall  and  Kelson  (1959),  external  and  cranial  measurements 
of  a  male  and  female  from  Morelos;  Schaldach  and  McLaughlin  (1960),  external  and 
cranial  measurements  of  two  males  and  six  females  from  Arizona,  one  female  from 
Sonora,  and  four  males  and  a  female  from  Oaxaca  (mean,  range);  Axtell  (1962),  external 
measurements  of  a  male,  female,  and  juvenile,  and  cranial  measurements  of  the  two  adults 
from  Coahuila;  Baker  and  Greer  (1962),  external  and  cranial  measurements  (mean,  range) 
of  six  males  from  Durango;  Davis  et  al.  (1964),  external  and  cranial  measurements  of  one 
female  from  Honduras;  Villa-R.  (1967),  external  measurements  (mean,  range)  of  seven 
males  and  females  combined  and  cranial  measurements  (mean,  range)  of  six  males  and 
females  combined  from  Mexico;  Barbour  and  Davis  (1969),  range  of  forearm  length 
of  the  species;  Goodwin  (1969),  forearm  and  cranial  measurements  of  three  males  from 
Oaxaca;  Anderson  (1972),  external  measurements  of  a  specimen  from  Arizona  and  cranial 
measurements  of  one  from  Sinaloa;  Findley  et  al.  (1975),  external  measurements  (mean, 
range)  of  12  females  from  New  Mexico. 

Glossophaga  alticola  Davis,  1944 

Measurements  of  Glossophaga  alticola  have  been  recorded  as  follows:  Davis  (1944), 
external  and  cranial  measurements  of  the  male  holotype  and  a  female  from  Tlaxcala; 
Davis  and  Russell  (1952),  external  and  cranial  measurements  (mean,  range)  of  seven  males 
and  six  females  from  Morelos;  Gardner  (1962a),  a  graphic  representation  (mean,  se,  range) 
of  variation  in  forearm  and  cranial  measurements  in  the  species;  Villa-R.  (1963),  com¬ 
parison  of  externa!  and  cranial  measurements  as  in  the  original  description  of  Glossophaga 
morenoi,  G.  alticola ,  and  G.  commissarisi  and  external  measurements  of  19  males  and 
18  females  and  cranial  measurements  of  19  males  and  19  females  of  G.  morenoi  (mixed 
sample  of  G.  alticola  and  G.  commissarisi)  from  Mexico;  Villa-R.  (1967),  external 
measurements  (19  males,  18  females)  and  cranial  measurements  (19  males,  19  females) 
of  G.  morenoi  (mixed  sample  of  G.  alticola  and  G.  commissarisi  from  Mexico);  Goodwin 
(1969),  forearm  and  cranial  measurements  of  five  females  and  one  subadult  male  from 
Oaxaca. 


Glossophaga  commissarisi  Gardner,  1962 

Measurements  of  Glossophaga  commissarisi  have  been  recorded  as  follows:  Gardner 
(1962a),  external  and  cranial  measurements  of  the  male  holotype  from  Chiapas  and  a 
graphic  representation  (mean,  se,  range)  of  variation  in  forearm  and  cranial  measure¬ 
ments  in  the  species;  Villa-R.  (1963),  comparison  of  external  and  cranial  measurements 
as  in  the  original  description  of  Glossophaga  morenoi,  G.  alticola,  and  G.  commissarisi, 
external  measurements  of  19  males  and  18  females  and  cranial  measurements  of  19  males 
and  19  females  of  G.  morenoi  (mixed  sample  of  G.  alticola  and  G.  commissarisi )  from 
Mexico;  Villa-R.  (1967),  external  measurements  (18  males,  19  females)  and  cranial  measure¬ 
ments  (19  males,  19  females)  of  G.  morenoi  (mixed  sample  of  G.  alticola  and  G.  com¬ 
missarisi)-,  Goodwin  (1969),  forearm  and  cranial  measurements  of  a  male,  female,  and  three 
unsexed  specimens  from  Oaxaca;  Jones  et  al.  (1972),  forearm  and  cranial  measurements  of 
three  females  from  Sinaloa. 


Glossophaga  longirostris  Miller,  1898 

Measurements  of  Glossophaga  longirostris  have  been  recorded  as  follows:  Miller 
(1898),  external  and  cranial  measurements  of  the  female  holotype  from  Colombia;  Robinson 
and  Lyon  (1901),  external  measurements  and  greatest  length  of  skull  for  nine  males  and 
four  females  from  Venezuela;  G.  M.  Allen  (1908),  external  measurements  (range)  of  ten 
specimens  from  Carriacou,  Lesser  Antilles;  Miller  (1913a),  external  and  cranial  measure- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


33 


ments  of  the  male  holotype  of  G.  I.  rostrata  from  Grenada,  Lesser  Antilles;  Miller  (1913 /?), 
external  and  cranial  measurements  of  nine  males  and  one  female  from  Venezuela,  one 
male  and  one  unsexed  specimen  from  Colombia,  nine  males  from  Grenada,  three  males, 
two  females,  and  three  unsexed  specimens  from  Carriacou,  and  ten  males  and  ten  females 
from  Curasao;  Elliot  (1917),  external  and  cranial  measurements  of  the  holotype  of  G. 
I.  rostrum ;  Hershkovitz  (1949),  external  and  cranial  measurements  (range)  of  five  males 
and  two  females  combined  from  Colombia;  Husson  (1960),  forearm  measurements  (range) 
of  21  males  and  42  females  and  cranial  measurements  (range)  in  12  specimens  from 
Aruba,  Curasao,  and  Bonaire  islands;  Goodwin  and  Greenhall  (1961),  forearm  measure¬ 
ments  of  10  females  and  cranial  measurements  of  four  females  from  Tobago,  forearm 
measurements  of  14  females  and  cranial  measurements  of  10  females  from  Trinidad; 
Tamsitt  and  Valdivieso  (1963«)  and  Valdivieso  (1964),  external  and  cranial  measurements 
of  a  male  and  two  females  from  Girardot,  Colombia;  Smith  and  Genoways  (1974),  fore¬ 
arm  and  cranial  measurements  of  specimens  from  Curasao  (20  from  Miller,  19136), 
Margarita  Island  (9),  Venezuela  (22),  Trinidad  (5),  Grenada  (9),  and  St.  Vincent  (10). 

Geographic  variation. — Smith  and  Genoways  (1974)  stated  that  a  comparison  of 
measurements  obtained  from  specimens  from  Margarita  Island  with  those  of  the  main¬ 
land  and  Antillean  islands  showed  that  the  material  from  Margarita  Island  is  well  within 
the  range  of  variation  of  the  mainland  specimens  and  overlap  those  obtained  from  Antillean 
material. 


Glossophaga  soricina (Pallas,  1766) 

Measurements  of  Glossophaga  soricina  have  been  recorded  as  follows:  Dobson  (1878<v), 
external  measurements  of  a  female;  H.  Allen  (1895),  external  measurements  of  the  holotype 
of  Glossophaga  truer,  Robinson  and  Lyon  (1901),  external  measurements  and  greatest 
length  of  skull  of  one  male  and  three  females  from  Venezuela;  Rehn  (1902rtX  external  and 
cranial  measurements  of  the  female  holotype  of  G.  s.  antillarum  from  Jamaica  and  one 
specimen  each  from  Guyana,  Trinidad,  and  the  Bahamas;  Cabrera  (1903),  external 
measurements  for  the  species  in  Chile;  Elliot  (1904),  external  measurements  of  one 
specimen  from  Tres  Marias  Islands  and  external  and  cranial  measurements  of  two  additional 
specimens;  G.  M.  Allen  (1908),  forearm  measurements  of  three  specimens  from  Peru; 
G.  M.  Allen  (1911),  forearm  and  cranial  measurements  of  a  specimen  from  Jamaica; 
Miller  (1913  6),  external  and  cranial  measurements  of  nine  individuals  (eight  females, 
one  male)  from  Brazil,  one  female  from  Guyana,  seven  (five  females,  one  male,  one  un¬ 
sexed)  from  Venezuela,  10  (five  females,  five  males)  from  Trinidad,  five  (two  females, 
two  males,  one  unsexed)  from  Colombia,  eight  (three  females,  five  males)  from  Moyobamba, 
Peru,  1 1  (seven  females,  four  males)  from  Paraguay,  20  specimens  (nine  females,  1 1  males) 
from  the  mainland  of  Mexico,  two  (one  female,  one  male)  from  Nicaragua,  one  male  from 
Costa  Rica,  five  (three  females,  two  males)  from  Chiriqui,  Panama,  10  (five  males,  five 
females)  from  Panama,  12  (six  females,  six  males)  from  Tres  Marias  Islands,  14  (five  females, 
nine  males)  from  Balsas,  Peru,  three  (two  females,  one  male)  from  Charapex,  Peru,  and 
two  females  from  Jamaica;  Elliot  (1917),  external  and  cranial  measurements  (range)  of 
specimens  from  Nayarit  to  Panama;  Lima  (1926),  external  measurements  of  a  male  from 
Brazil;  Goodwin  (1934),  external  measurements  (mean)  of  five  specimens  from  Guatemala; 
Martinez  and  Villa-R.  (1938),  external  measurements  of  one  specimen  and  cranial 
measurements  of  four  specimens  of  G.  morenoi  (  — G.  soricina )  from  Morelos;  Martinez 
and  Villa-R.  (1941),  external  and  cranial  measurements  (mean,  variance,  and  correlation 
between  measurements)  of  52  males  and  25  females  from  Guerrero;  Cunha  Vieira  (1942), 
external  measurements  of  nine  males  and  one  of  unknown  sex  and  cranial  measurements 
of  three  males  from  Brazil;  Goodwin  (1942 a),  external  and  cranial  measurements  of  two 
specimens  from  Honduras;  Goodwin  (1946),  external  and  cranial  measurements  of  two 
males  from  Costa  Rica;  Hershkovitz  (1949),  external  and  cranial  measurements  of  three 
females  from  Colombia;  Dalquest  (1951),  forearm  and  cranial  measurements  of  one 


34 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


specimen  (sex  unknown)  from  Trinidad;  Davis  and  Russell  (1952),  external  and  cranial 
measurements  (mean,  range)  of  seven  males  and  12  females  from  Morelos  (G.  s.  leachi)', 
Dalquest  (1953a),  external  measurements  (means)  of  seven  males  and  15  females  and 
cranial  measurements  (means)  of  nine  males  and  seven  females  from  San  Luis  Potosi; 
Villa-R.  (1953),  external  and  cranial  measurements  (mean,  range)  of  specimens  from  Tlaxcala 
(1),  Districto  Federal  (15),  Morelos  (12),  and  Guerrero  (5);  de  la  Torre  (1954),  external 
and  cranial  measurements  of  one  female  from  Tamaulipas;  de  la  Torre  (1955),  forearm 
measurements  (mean,  range)  of  nine  specimens  (six  males,  three  females  combined)  from 
Guerrero;  Felten  (1956a),  external  measurements  (mean,  range)  of  286  males  and  200 
females  and  cranial  measurements  of  27  males  and  38  females  from  El  Salvador;  Ryan 
(1960),  external  measurements  of  two  females  from  Guatemala;  Burt  and  Stirton  (1961), 
forearm  and  cranial  measurements  (range)  of  43  males  and  32  females  from  El  Salvador; 
Goodwin  and  Greenhall  (1961),  forearm  measurements  (range)  of  20  specimens  and 
cranial  measurements  of  three  females  from  Trinidad;  Husson  (1962),  external  and  cranial 
measurements  of  five  males  and  five  females  from  Surinam;  Gardner  (1962a),  graphic 
representation  (mean,  range,  se)  of  variation  in  forearm  and  cranial  measurements  of  the 
species;  Tamsitt  and  Valdivieso  (1963a),  external  measurements  (mean,  range)  of  51  specimens 
from  Colombia;  Tamsitt  and  Valdivieso  (19636),  external  measurements  of  one  male  and 
one  female  from  Colombia;  Villa-R.  (1963),  comparison  of  external  and  cranial  measure¬ 
ments  as  in  the  original  description  of  Glossophaga  morenoi,  G.  alticola,  and  G.  commissar  is  i; 
Starrett  and  de  la  Torre  (1964),  forearm  measurements  of  two  males  and  14  females  (mean, 
range)  from  El  Salvador,  Honduras,  Nicaragua,  and  Costa  Rica;  Valdivieso  (1964), 
external  measurements  (mean,  range)  of  77  specimens  from  Colombia;  Aellen  (1965), 
external  and  cranial  measurements  of  one  male  and  one  female  from  Peru;  Villa-R.  (1967), 
external  measurements  (mean,  se,  range)  of  70  males  and  37  females  and  cranial  measure¬ 
ments  of  56  males  and  25  females  from  Mexico;  Pirlot  (1968),  forearm  measurements  of 
a  female  from  Peru;  Goodwin  (1969),  forearm  and  cranial  measurements  of  a  female  from 
Peru;  Goodwin  (1969),  forearm  and  cranial  measurements  of  two  females,  one  subadult 
male,  and  three  unsexed  individuals  from  Oaxaca;  Anderson  (1972),  external  measurements 
of  two  specimens  and  cranial  measurements  of  one  from  Chihuahua;  Jones  et  al.  (1972), 
forearm  and  cranial  measurements  (mean,  range)  of  nine  males  and  one  female  combined 
from  Sinaloa;  Taddei  (19756),  external  measurements  (mean,  se,  range)  of  59  males  and  47 
females  and  cranial  measurements  of  20  males  and  20  females  from  Brazil. 

Individual  variation. — In  specimens  from  Brazil,  coefficients  of  variation  for  external 
measurements  varied  in  59  males  from  2.00  to  5.60  and  in  47  females  from  2.10  to  5.26; 
and  for  cranial  measurements  in  20  males,  CVs  ranged  from  1.75  to  3.44  and  in  20  females 
from  1.65  to  3.37  (Taddei,  19756). 

Secondary  sexual  variation. — Taddei  (19756)  found  females  to  be  significantly  larger 
than  males  in  four  (head  and  body  length,  forearm  length,  fourth  and  fifth  metacarpal)  of 
17  external  measurements.  In  the  case  of  cranial  measurements,  females  were  significantly 
larger  in  two  measurements  (length  of  molar,  mandibular  toothrow)  of  15  but  significantly 
smaller  in  five  (breadth  across  canines,  zygomatic  breadth,  braincase  breadth,  mastoid 
breadth,  cranial  depth). 


Hylonycteris  underwood i  Thomas,  1903 

Measurements  of  Hylonycteris  underwoodi  have  been  recorded  as  follows:  Thomas 
(1903a),  forearm  and  cranial  measurements  of  the  holotype  and  external  measurements 
of  a  second  specimen  from  Costa  Rica;  Elliot  (1904),  external  and  cranial  measurements  of 
one  specimen;  Goodwin  (1942a,  1946),  forearm  and  cranial  measurements  of  the  holotype 
from  Costa  Rica;  Hall  and  Kelson  (1959),  forearm  and  cranial  measurements  of  the 
holotype;  Davis  and  Carter  (1962a),  external  and  cranial  measurements  of  the  holotype 
and  two  additional  specimens  (sex  unknown)  from  Costa  Rica,  one  male  and  four  females 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


35 


from  Veracruz,  and  one  male  and  one  female  from  Oaxaca;  Jones  (1964),  forearm  and 
cranial  measurements  of  one  male  and  one  female  from  Oaxaca;  Villa-R.  (1967),  external 
and  cranial  measurements  of  one  specimen  from  Tabasco;  Goodwin  (1969),  forearm  and 
cranial  measurements  of  two  males  and  two  females  from  Oaxaca;  Gardner  et  al.  (1970), 
forearm  and  cranial  measurements  (mean,  range)  of  14  males  and  seven  females  from 
Costa  Rica;  Phillips  and  Jones  (1971),  forearm  and  cranial  measurements  (mean,  range) 
of  four  males  and  six  females  combined,  additional  measurements  of  the  female  holotype  of 
H.  u.  minor  from  Jalisco,  comparative  measurements  of  a  Veracruz  male,  and  a  male  and 
female  from  Oaxaca;  Jones  and  Homan  (1974),  external  and  cranial  measurements  as 
given  by  Gardner  et  al.  (1970)  and  Phillips  and  Jones  (1971). 

Secondary  sexual  variation. — Females  averaged  larger  than  males  throughout  the  range 
of  the  species  according  to  Phillips  and  Jones  (1971). 

Geographic  variation. — Davis  and  Carter  (1962a)  noted  that  specimens  from  Oaxaca 
appeared  to  be  smaller  than  those  from  Veracruz  and  Costa  Rica.  However,  Jones  (1964) 
found  his  Oaxacan  male  specimen  to  be  larger  than  those  previously  reported  and  the 
measurements  of  his  female  specimens  fell  among  the  largest  known  individuals  of  the 
species. 

Specimens  from  Jalisco  and  southern  Oaxaca  (Davis  and  Carter,  1962a)  were  included 
in  a  subspecies  by  Phillips  and  Jones  (1971).  They  concluded  that  these  specimens  were 
smaller  externally  and  cranially  than  H.  it.  underwoodi  from  northern  Oaxaca,  Veracruz, 
and  Guatemala. 


Leptonycteris  curasoae  Miller,  1900 

Measurements  of  Leptonycteris  curasoae  have  been  recorded  as  follows:  Miller  (19006), 
external  and  cranial  measurements  of  the  male  holotype  from  Curasao;  Hoffmeister 
(1957),  external  measurements  of  the  holotype  and  three  male  topotypes;  Husson  (1960), 
forearm  measurements  (range)  of  21  specimens  and  cranial  measurements  (range)  of  13 
specimens  from  Aruba,  Curasao,  and  Bonaire  islands;  Davis  and  Carter  (19626),  external 
and  cranial  measurements  of  four  males  and  two  females  combined  (mean,  range);  Pirlot 
(1965a),  external  and  cranial  measurements  of  the  male  holotype  of  L.  c.  tarlosti,  a  male, 
and  three  females  from  Margarita  Island;  Marinkelle  and  Cadena  (1972),  external  and 
cranial  measurements  of  two  females  from  Colombia;  Smith  and  Genoways  (1974), 
forearm  and  cranial  measurements  of  12  specimens  from  Margarita  Island,  two  from 
Aruba,  five  from  Curasao,  and  one  from  Bonaire. 

Geographic  variation. — In  his  study  of  this  genus,  Hoffmeister  (1957)  considered  L. 
curasoae  to  be  a  subspecies  of  L.  nivalis.  However,  Davis  and  Carter  (19626)  in  their 
review  of  the  genus  and  subsequent  authors  have  consider  L.  curasoae  a  distinct  species. 
Pirlot  (1965a)  recognized  specimens  from  Margarita  Island  as  a  distinct  subspecies,  however, 
Smith  and  Genoways  (1974),  after  examining  specimens  from  throughout  the  range  of  the 
species,  considered  the  subspecific  status  of  the  island  forms  unwarranted. 

Leptonycteris  nivalis  (Saussure,  1860) 

Measurements  of  Leptonycteris  nivalis  have  been  recorded  as  follows:  Saussure 
(1860c),  external  measurements  of  one  specimen;  Dobson  (1878a),  external  measure¬ 
ments  of  one  specimen;  Miller  (19006),  external  and  cranial  measurements  of  a  male  from 
Colima;  Elliot  (1904),  external  and  cranial  measurements  of  one  specimen;  Martinez 
and  Villa-R.  (1938),  external  measurements  of  one  specimen;  Martinez  and  Villa-R. 
(1940),  external  and  cranial  measurements  (mean,  sd)  of  samples  of  males  and  females  from 
Guerrero;  Goodwin  (1942a,  1946),  external  and  cranial  measurements  of  a  male  from 
Colima;  Dalquest  (1953a),  external  and  cranial  measurements  of  four  males  and  one 
female  from  San  Luis  Potosi;  de  la  Torre  (1955),  forearm  measurements  of  one  male  from 


36 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Guerrero;  Baker  (1956),  external  and  cranial  measurements  of  two  males  and  mean  and 
range  of  five  females  from  Coahuila;  Hoffmeister  (1957),  cranial  measurements  of  the 
holotype  of  L.  n.  nivalis  (Veracruz),  external  and  cranial  measurements  (mean)  of  six 
males  and  eight  females  combined  from  Texas,  and  external  measurements  (mean)  of  11 
males  and  29  females  combined  from  Nuevo  Leon;  Stains  (1957),  external  and  cranial 
measurements  of  the  holotype  and  mean  and  range  of  the  holotype  and  22  topotypes  of 
L.  n.  longala  from  Coahuila  (see  also  Jones,  1958);  Hall  and  Kelson  (1959),  external  and 
cranial  measurements  (range)  of  a  large  series  of  specimens  from  Jalisco;  Davis  and 
Carter  (19626),  external  and  cranial  measurements  of  three  males  and  seven  females  (mean, 
range);  Alvarez  (1963),  external  and  cranial  measurements  of  five  males  and  five  females 
combined  (mean,  range)  from  Tamaulipas;  Baker  and  Cockrum  (1966),  external  and 
cranial  measurements  of  two  females  from  Sinaloa;  Villa-R.  (1967),  external  measure¬ 
ments  of  50  specimens  (mean,  sd,  range)  and  cranial  measurements  of  37  (mean,  sd,  range) 
from  Mexico;  Goodwin  (1969),  external  and  cranial  measurements  of  two  males  and 
two  females  from  Morelos,  and  one  female  from  Veracruz;  Barbour  and  Davis  (1969), 
range  of  forearm  length  in  the  species;  Anderson  (1972),  external  and  cranial  measurements 
of  one  specimen;  Matson  and  Patten  (1975),  forearm  measurements  (mean,  range)  of 
seven  males  from  Zacatecas. 

Individual  variation. — In  specimens  from  Guerrero,  coefficients  of  variation  for 
external  measurements  varied  in  males  from  3.03  to  16.25  and  in  females  from  1.04  to 
16.58;  CV  values  for  cranial  measurements  in  males  ranged  from  1.68  to  7.44  and  in  females 
from  1.23  to  5.58  (Martinez  and  Villa-R.,  1940). 

Geographic  variation. — Hoffmeister  (1957)  and  Davis  and  Carter  (19626)  have  recently 
reviewed  this  genus.  Davis  and  Carter  (19626)  gave  characteristics  by  which  the  currently 
recognized  species  can  be  distinguished. 

Leptonycteris  sanborni  Hoffmeister,  1957 

Measurements  of  Leptonycteris  sanborni  have  been  recorded  as  follows:  Hoffmeister 
(1957),  external  measurements  of  22  females  and  cranial  measurements  of  21  females 
from  Arizona,  external  measurements  (mean)  of  10  males  from  Chihuahua,  and  the  mean 
of  eight  males  from  Colima;  Davis  and  Carter  (19626),  external  and  cranial  measure¬ 
ments  (mean,  range)  of  five  males  and  five  females;  Baker  and  Cockrum  (1966),  external 
and  cranial  measurements  of  one  specimen  from  Sinaloa;  Villa-R.  (1967),  external  measure¬ 
ments  (jV=51)  and  cranial  measurements  (N=  39)  (mean,  sd,  range)  of  L.  yerbabuenae 
(  =  L.  sanborni)  from  Mexico;  Genoways  and  Jones  (1968),  forearm  measurements 
(mean)  of  28  males  from  Zacatecas;  Barbour  and  Davis  (1969),  range  of  forearm  length 
of  the  species;  Anderson  (1972),  external  measurements  (mean,  sd,  range)  of  24  specimens 
from  Chihuahua  and  external  and  cranial  measurements  of  one  specimen  from  Sonora; 
Ramirez-Pulido  and  Alvarez  (1972),  external  and  cranial  measurements  of  a  lectotype  and 
external  measurements  of  a  male  and  female  paralectotype  of  L.  yerbabuenae ;  Jones  and 
Bleier  (1974),  forearm  and  cranial  measurements  of  one  male  from  El  Salvador;  Matson 
and  Patten  (1975),  forearm  and  cranial  measurements  of  five  males  (mean,  range)  and  one 
female  from  Zacatecas. 

Geographic  variation. — The  species  was  originally  described  as  a  subspecies  of  L.  nivalis 
by  Hoffmeister  (1957).  Davis  and  Carter  (19626)  demonstrated  characteristics  by  which  this 
taxon  could  be  distinguished  from  L.  nivalis.  Considerable  controversy  exists  in  the  litera¬ 
ture  over  the  relationships  of  this  taxon  and  L.  yerbabuenae.  Because  the  holotype  of 
yerbabuenae  has  been  lost  and  because  the  original  series  was  a  composite,  Watkins  et  al. 
(1972)  considered  yerbabuenae  to  be  a  nomen  dubium.  However,  as  recently  as  Ramirez- 
Pulido  and  Alvarez  (1972),  authors  have  believed  that  the  name  yerbabuenae  superceded 
sanborni.  The  reader  is  warned  to  take  great  care  in  using  measurements  recorded  in  the 
earlier  literature  concerning  this  genus  because  considerable  confusion  has  existed  in  the 
proper  identification  of  the  species. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


37 


Lichonycteris  degener  Miller,  1931 

Miller  (1931)  gave  external  and  cranial  measurements  of  the  female  holotype  from  Brazil. 

Lichonycteris  obscura  Thomas,  1895 

Measurements  of  Lichonycteris  obscura  have  been  recorded  as  follows:  Thomas  (1895), 
external  and  cranial  measurements  of  the  female  holotype  from  Nicaragua;  Elliot  (1904), 
external  and  cranial  measurements  of  one  specimen  from  Nicaragua;  Sanborn  (1936), 
external  and  cranial  measurements  of  a  female  from  Costa  Rica  and  the  holotype  from 
Nicaragua;  Goodwin  (1942,  1946),  external  and  cranial  measurements  of  two  females  from 
Costa  Rica;  Hall  and  Kelson  (1959),  external  and  cranial  measurements  of  two  females 
from  Costa  Rica;  Husson  (1962),  external  and  cranial  measurements  of  a  male  from  Costa 
Rica;  Davis  et  at.  (1964),  external  and  cranial  measurements  of  a  female  from  Nicaragua; 
Carter  et  al.  (1966),  external  and  cranial  measurements  of  three  females  from  Guatemala; 
Gardner  et  al.  (1970),  external  and  cranial  measurements  (mean,  range)  of  one  male  and 
three  females  from  Costa  Rica;  Jones  et  al.  (19716),  external  and  cranial  measurements  of 
three  males  from  Nicaragua;  Marinkelle  and  Cadena  (1972),  forearm  measurements  (range) 
of  three  females  and  one  unsexed  specimen  from  Colombia;  Gardner  (1976),  external 
and  cranial  measurements  of  two  females  from  Peru. 

Lionycteris  spurrelli  Thomas,  1913 

Measurements  of  Lionycteris  spurrelli  have  been  recorded  as  follows:  Thomas  (1913), 
external  and  cranial  measurements  of  the  immature  male  holotype  from  Colombia;  Gold¬ 
man  (19146),  greatest  length  of  skull  of  a  specimen  from  Colombia;  Sanborn  (1941), 
external  measurements  of  one  male  and  one  female  and  cranial  measurements  of  one 
specimen  from  Guyana,  and  the  measurements  for  the  holotype  from  Colombia. 

Lonchophylla  concava  Goldman,  1914 

Measurements  of  Lonchophylla  concava  have  been  recorded  as  follows:  Goldman 
(1914a),  external  and  cranial  measurements  of  the  male  holotype  from  Panama;  Elliot 
(1917),  external  and  cranial  measurements  of  the  holotype;  Goodwin  (1946),  external 
and  cranial  measurements  of  the  holotype  from  Panama;  Hall  and  Kelson  (1959),  external 
and  cranial  measurements  of  the  holotype  of  L.  concava',  Davis  et  al.  (1964),  external  and 
cranial  measurements  of  one  male  and  two  females  from  Costa  Rica;  Pirlot  (1968),  forearm 
measurements  of  one  male  from  Peru;  Gardner  et  al.  (1970),  external  and  cranial  measure¬ 
ments  (mean,  range)  of  five  specimens  from  Costa  Rica;  Marinkelle  and  Cadena  (1972), 
forearm  measurements  of  two  females  from  Colombia. 

Lonchophylla  hesperiaG.  M.  Allen,  1908 

Measurements  of  Lonchophylla  hesperia  have  been  recorded  as  follows:  G.  M.  Allen 
(1908),  external  and  cranial  measurements  of  the  male  holotype  and  two  additional 
specimens  from  Peru;  Gardner  (1976),  external  and  cranial  measurements  of  one  male 
and  one  female  from  Peru. 


Lonchophylla  mordax  Thomas,  1903 

Measurements  of  Lonchophylla  mordax  have  been  recorded  as  follows:  Thomas 
(1903c),  external  and  cranial  measurements  of  the  male  holotype  from  Brazil;  G.  M.  Allen 
(1908),  external  and  cranial  measurements  of  the  holotype  from  Brazil;  Lima  (1926), 
external  measurements  of  a  specimen  from  Brazil;  Sanborn  (1941),  forearm  measurements 
(range)  of  18  males  from  Brazil;  Cunha  Vieira  (1942),  external  measurements  of  a  male 
and  a  female  and  cranial  measurements  of  a  male  from  Brazil;  Baker  (1974),  external  and 
cranial  measurements  of  one  female  from  Ecuador. 


38 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Lonchophylla  robusta  Miller,  1912 

Measurements  of  Lonchyphylla  robusta  have  been  recorded  as  follows:  Miller  (1912) 
and  Goodwin  (1946),  external  and  cranial  measurements  of  the  male  holotype  and  a  female 
from  Panama;  Hall  and  Kelson  (1959),  external  and  cranial  measurements  of  the  holotype 
and  a  female  topotype;  Walton  (1963),  external  and  cranial  measurements  (mean,  sd,  se, 
range)  of  specimens  from  Panama  (N=21)  and  Costa  Rica  (A=10);  Valdivieso  (1964), 
external  and  cranial  measurements  of  one  female  from  Colombia;  Tuttle  (1970),  external 
measurements  of  one  male  and  one  female  and  cranial  measurements  of  the  female  from 
Peru;  Gardner  (1976),  external  and  cranial  measurements  of  a  male  from  Peru. 

Secondary  sexual  variation. — According  to  Walton  (1963),  no  sexual  dimorphism  in  size 
was  evident  in  specimens  from  Panama  and  Costa  Rica. 

Geographic  variation. — Walton  (1963)  found  specimens  from  Panama  to  be  larger  than 
those  from  Costa  Rica.  Of  seven  external  measurements,  three  (total  length,  length  of  hind 
foot,  ear  length)  proved  to  be  significantly  different,  whereas  in  nine  cranial  measurements 
there  were  four  (skull  length,  skull  width,  interorbital  width,  width  of  rostrum  at  canines) 
that  showed  significant  differences. 

Lonchophylla  thomasiJ.  A.  Allen,  1904 

Measurements  for  Lonchophylla  thomasi  have  been  recorded  as  follows:  J.  A.  Allen 
(1904),  external  and  cranial  measurements  of  the  male  holotype  from  Venezuela;  Goodwin 
(1953),  forearm  and  cranial  measurements  of  the  holotype;  Husson  (1962),  external  and 
cranial  measurements  of  two  males  and  one  female  from  Surinam;  Hill  (1964),  forearm 
and  cranial  measurements  of  a  male  from  Guyana;  Gardner  (1976),  external  and  cranial 
measurements  (mean,  range)  of  six  males  and  six  females  combined  from  Peru. 

Monophyllus  plethodon  Miller,  1900 

Measurements  of  Monophyllus  plethodon  have  been  recorded  as  follows:  Miller  (1900a), 
external  and  cranial  measurements  of  the  male  holotype  of  M.  plethodon  from  Barbados, 
Lesser  Antilles;  Miller  (1902a),  external  and  cranial  measurements  of  the  male  holotype 
of  M.  p.  luciae  from  St.  Lucia,  Lesser  Antilles,  and  of  the  holotype  of  M.  p.  plethodon ;  Elliot 
(1904),  external  and  cranial  measurements  of  a  specimen  from  Barbados  and  one  from  St. 
Lucia;  Anthony  (1917),  cranial  measurements  of  the  holotype  and  two  additional  specimens 
(sub-Recent  fossils)  of  M.  frater  from  Puerto  Rico;  Anthony  (1918,  1925),  cranial  measure¬ 
ments  of  three  specimens  of  sub-Recent  fossils  from  Puerto  Rico;  Goodwin  (1953),  a  cranial 
measurement  of  the  holotype  of  M.  p.  frater  (subfossil)  from  Puerto  Rico;  Hall  and  Kelson 
(1959),  external  and  cranial  measurements  of  the  holotype  of  M.  plethodon,  M.  luciae, 
and  cranial  measurements  of  the  holotype  and  two  topotypes  of  M.  frater,  Schwartz  and 
Jones  (1967),  external  and  cranial  measurements  of  specimens  from  Angulla,  Barbuda, 
Antigua,  Dominica,  St.  Lucia,  and  Barbados;  Choate  and  Birney  (1968),  cranial  measure¬ 
ments  of  six  males  and  nine  females  from  Dominica  and  of  one  sub-Recent  fossil  from  Puerto 
Rico;  Koopman  (1968),  forearm  and  cranial  measurements  of  one  male  from  Dominica, 
a  specimen  from  Antigua,  and  one  female  from  Anguilla;  Homan  and  Jones  (19756), 
external  and  cranial  measurements  (range)  of  Lesser  Antillean  representatives  of  the  species 
(after  Schwartz  and  Jones,  1967). 

Geographic  variation. — Schwartz  and  Jones  (1967)  have  recently  reviewed  geographic 
variation  in  Monophyllus  plethodon.  They  recognized  three  subspecies  occurring  on  Puerto 
Rico  and  the  Lesser  Antilles.  One  subspecies  was  known  only  as  a  fossil  from  Puerto  Rico. 
Specimens  of  M.  plethodon  on  Barbados  were  distinguished  from  all  other  Lesser  Antillean 
populations  by  overall  small  size. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


39 


Monophyllus  redmani  Leach,  1821 

Measurements  of  Monophyllus  redmani  have  been  recorded  as  follows:  Gundlach  (1872, 
1877),  external  measurements  of  a  Cuban  specimen;  Dobson  ( 1 878a),  external  measure¬ 
ments  of  one  male;  Miller  (1900a),  external  and  cranial  measurements  of  the  male  holotype 
of  M.  portoricensis  from  Puerto  Rico,  the  male  holotype  of  M.  clinedaphus  from  an  unknown 
locality,  and  a  male  from  Jamaica,  as  well  as  external  measurements  of  one  male  and  three 
females  from  Puerto  Rico;  Miller  (1902a),  external  and  cranial  measurements  of  the  male 
holotype  of  M.  cubanus  from  Cuba  and  cranial  measurements  of  one  male  from  Jamaica; 
Elliot  (1904),  external  and  cranial  measurements  of  one  specimen  each  from  Puerto  Rico, 
Cuba,  and  Jamaica;  Miller  (1904),  external  measurements  of  eight  males  and  seven  females 
from  Cuba;  Anthony  (1917),  cranial  measurements  of  a  specimen  from  Puerto  Rico; 
Anthony  (1918,  1925),  external  (18  specimens)  and  cranial  (five  specimens)  measurements 
(mean,  range)  of  individuals  from  Puerto  Rico;  Hall  and  Kelson  (1959),  external  and 
cranial  measurements  of  a  male  from  Jamaica,  the  holotype  of  M.  cubanus,  and  of  the 
holotype  of  M.  clinedaphus,  as  well  as  cranial  measurements  of  the  holotype  of  M.  portori¬ 
censis  and  the  range  in  external  measurements  of  five  specimens  from  Puerto  Rico; 
Schwartz  and  Jones  (1967),  external  and  cranial  measurements  of  the  three  recognized 
subspecies  from  Jamaica,  Cuba,  Hispaniola,  and  Puerto  Rico;  Choate  and  Birney  (1968), 
cranial  measurements  of  one  fossil  specimen  from  Puerto  Rico;  Silva-Taboada  (1974), 
measurements  of  fossil  humeri,  crania,  and  mandibles  from  Cuba;  Buden  (1975a),  external 
and  cranial  measurements  (mean,  range)  of  specimens  from  Jamaica,  Cuba,  Hispaniola, 
Bahamas,  and  Puerto  Rico;  Homan  and  Jones  (1975a),  external  and  cranial  measurements 
(range)  of  specimens  of  the  three  recognized  subspecies  (after  Schwartz  and  Jones,  1967; 
Buden,  1975  a). 

Geographic  variation. — Schwartz  and  Jones  (1967)  have  recently  reviewed  geographic 
variation  in  Monophyllus  redmani.  They  recognized  three  subspecies,  all  occurring  in  the 
Greater  Antilles.  Specimens  from  Jamaica  were  characterized  by  large  body  and  cranial 
size  but  a  relatively  short  forearm.  On  Cuba  and  Hispaniola,  bats  were  characterized  by 
small  body,  moderate  skull  size,  and  relatively  long  forearms.  Specimens  of  M.  redmani 
from  Puerto  Rico  are  of  generally  small  size. 

Musonycteris  Harrison i  Schaldach  and  McLaughlin,  1960 

Measurements  of  Musonycteris  harrisoni  have  been  recorded  as  follows:  Schaldach  and 
McLaughlin  (1960),  external  and  cranial  measurements  of  the  male  holotype,  10  male 
paratypes,  and  two  female  paratypes  from  Colima;  Villa-R.  (1967),  external  measurements 
of  nine  specimens  (mean,  range),  and  cranial  measurements  (mean,  range)  of  six  specimens 
from  Colima;  Goodwin  (1969),  forearm  and  cranial  measurements  of  one  male  from 
Guerrero  and  a  male  and  female  from  Colima. 

Platalina  genovensium  Thomas,  1928 

Measurements  of  Platalina  genovensium  have  been  recorded  as  follows:  Thomas 
(1928a),  external  and  cranial  measurements  of  the  male  holotype  from  Peru;  Sanborn 
(1936),  external  and  cranial  measurements  of  the  male  holotype  and  a  second  male  from 
Peru;  Sanborn  (1943),  forearm  measurements  (range)  for  the  species  from  Peru;  Aellen 
(1965),  external  and  cranial  measurements  of  a  male  in  addition  to  the  holotype  (Thomas, 
1928a),  and  one  male  (Sanborn,  1936)  from  Peru. 

Scleronyeteris  ega  Thomas,  1912 

Thomas  (19126)  gave  external  and  cranial  measurements  of  the  female  holotype  from 
Ega,  Brazil. 


40 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Subfamily  Carolliinae 
Carollia  brevicauda  (Schinz,  1821) 

Measurements  of  Carollia  brevicauda  have  been  recorded  as  follows:  Peters  ( 1 865 z/), 
external  measurements  of  one  specimen;  Dobson  (1878a),  external  measurements  of  one 
specimen;  H.  Allen  ( 18906),  external  measurements  of  three  males  and  six  females;  Robin¬ 
son  and  Lyon  (1901),  external  measurements  of  two  males  from  Venezuela;  Goodwin 
(1942a),  external  and  cranial  measurements  of  one  male  and  one  female  from  Honduras 
(originally  reported  as  C.  castanea)',  Dalquest  (1953a),  external  measurements  of  one  male 
and  six  females  (mean)  and  cranial  measurements  (mean)  of  six  males  and  five  females  from 
San  Luis  Potosl  (originally  reported  as  perspicillata)',  Jones  (1966),  forearm  and  cranial 
measurements  (range)  of  12  specimens  from  Guatemala  (another  specimen  representing 
C.  subrufa  is  included  in  ranges);  Goodwin  (1969),  external  and  cranial  measurements 
of  a  male  and  a  female  from  Chiltepec,  Oaxaca  (these  were  originally  listed  as  C.  subrufa , 
but  according  to  Pine,  1972,  these  two  specimens  are  probably  C.  brevicauda)'.  Pine  (1972), 
external  measurements  (mean,  range)  of  four  males  and  10  females  from  San  Luis  Potosl, 
11  males  and  17  females  from  Veracruz,  seven  males  and  23  females  from  Chiapas,  15 
males  and  10  females  from  Guatemala,  one  male  and  one  female  from  Honduras,  20 
males  and  26  females  from  Panama,  nine  males  and  seven  females  from  Ecuador,  eight 
females  from  Brazil,  four  females  from  Peru,  one  male  from  Bolivia,  and  cranial  measure¬ 
ments  (mean,  range)  of  five  males  and  nine  females  from  San  Luis  Potosl,  11  males  and  15 
females  from  Veracruz,  seven  males  and  23  females  from  Chiapas,  15  males  and  10  females 
from  Guatemala,  one  male  and  one  female  from  Honduras,  20  males  and  26  females  from 
Panama,  nine  males  and  seven  females  from  Ecuador,  eight  females  from  Brazil,  five  males 
and  eight  females  from  Peru,  and  one  male  from  Bolivia;  Jones  et  al.  (1973),  external  and 
cranial  measurements  (mean,  range)  of  20  specimens  from  the  Yucatan  Peninsula. 

Geographic  variation. — According  to  Pine  (1972),  specimens  from  the  northernmost 
part  of  the  geographic  range  of  the  species  in  Mexico  are  the  largest. 

Carollia  castanea  H.  Allen,  1890 

Measurements  of  Carollia  castanea  have  been  recorded  as  follows:  H.  Allen  (18906), 
external  measurements  of  the  young  male  holotype  from  Costa  Rica;  Elliot  (1904),  external 
measurements  of  the  holotype  as  given  by  H.  Allen  (18906)  from  Costa  Rica;  Hahn  (1907), 
external  and  cranial  measurements  of  the  holotype  from  Costa  Rica;  Goodwin  (1946),  fore¬ 
arm  and  cranial  measurements  of  the  holotype  and  a  second  male  from  Costa  Rica; 
Hershkovitz  (1949),  external  and  cranial  measurements  of  one  male  and  one  female  from 
Colombia;  Husson  (1962),  external  and  cranial  measurements  of  two  females  from  Surinam; 
Pirlot  (1968),  external  and  cranial  measurements  discussed  in  conjunction  with  C.  perspicillata'. 
Pine  (1972),  external  measurements  of  10  males  and  four  females  from  Honduras,  five 
males  from  Nicaragua,  seven  males  and  eight  females  from  Costa  Rica,  31  males  and  20 
females  from  Panama,  five  males  from  Colombia,  three  males  and  two  females  from  Peru, 
one  female  from  Bolivia,  and  cranial  measurements  of  10  males  and  four  females  from 
Honduras,  five  males  from  Nicaragua,  seven  males  and  eight  females  from  Costa  Rica, 
31  males  and  19  females  from  Panama,  four  females  from  Colombia,  seven  males  and  four 
females  from  Peru,  and  one  female  from  Bolivia. 

Geographic  variation. — Pine  (1972)  could  detect  no  geographic  trends  in  variation  in  this 
species;  therefore,  he  considered  C.  castanea  to  be  monotypic. 

Carollia  perspicillata  (Linnaeus,  1758) 

Measurements  of  Carollia  perspicillata  have  been  recorded  as  follows:  Saussure  (1860c), 
external  measurements  of  one  specimen;  Peters  (1866a),  external  measurements  of  one 
specimen;  Miller  (1902a),  external  and  cranial  measurements  of  the  female  holotype  of  C. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


41 


tricolor  from  Paraguay;  Elliot  (1904),  external  and  cranial  measurements  of  a  single 
specimen;  Hahn  (1907),  external  measurements  (mean)  of  nine  specimens  from  Paraguay, 

10  from  Brazil,  10  from  Trinidad,  two  from  Guyana,  10  from  northern  Ecuador,  nine  from 
Colon,  Panama,  six  from  Panama,  Panama,  nine  from  Nicaragua,  13  from  Veracruz, 

1 1  from  Oaxaca,  two  from  Campeche,  and  13  from  Veracruz  (Jaltipan),  and  cranial  measure¬ 
ments  (mean)  of  eight  specimens  from  Paraguay,  two  from  Sao  Paulo,  Brazil,  five  from 
Naranhoa,  Brazil,  five  from  Trinidad,  two  from  Guyana,  four  from  Venezuela,  nine  from 
Colombia,  10  from  Ecuador,  eight  from  Oaxaca,  six  from  Veracruz,  two  from  Costa  Rica, 
two  from  Campeche,  three  from  Colon,  Panama,  six  from  Boqueron,  Panama,  and  six  from 
Panama,  Panama;  Lima  (1926),  external  measurements  of  a  specimen  from  Brazil;  Sanborn 
(1932),  forearm  measurement  of  one  specimen  from  Bolivia;  Goodwin  (1934),  external 
measurements  of  one  specimen  from  Guatemala;  Cunha  Vieira  (1942),  external  measure¬ 
ments  of  nine  males  and  four  females  and  cranial  measurements  of  three  males  and  one 
female  from  Brazil;  Goodwin  (1942 a),  external  and  cranial  measurements  of  two  males  from 
Honduras;  Goodwin  (1946),  external  and  cranial  measurements  of  a  male  and  female 
from  Costa  Rica;  Hershkovitz  (1949),  forearm  measurements  (range)  in  79  specimens  from 
northern  Colombia  and  the  mean  of  the  greatest  length  of  skull  in  this  sample  (some  specimens 
in  this  sample  are  brevicauda,  see  Pine,  1972);  Dalquest  (1951),  forearm  and  cranial 
measurements  (mean)  of  27  specimens  of  both  sexes  combined  from  Trinidad;  Felten 
(1956a),  external  measurements  (mean,  range)  of  15  males  and  28  females  and  cranial 
measurements  of  10  males  and  19  females  from  El  Salvador;  Felten  (1956*/),  external 
measurements  (mean,  range)  of  specimens  from  El  Salvador;  Ryan  (1960),  external  measure¬ 
ments  of  one  male  from  Guatemala;  Goodwin  and  Greenhall  (1961),  forearm  measurements 
(range)  of  30  specimens  and  cranial  measurements  of  one  male  and  one  female  from  Trinidad; 
Husson  (1962),  external  and  cranial  measurements  of  five  males  and  five  females  from 
Surinam;  Burt  and  Stirton  (1961),  forearm  and  cranial  measurements  (range)  of  22  males 
and  14  females  combined  from  El  Salvador;  Pirlot  (1963),  external  measurements  of  specimens 
from  Venezuela;  Butterworth  and  Starrett  (1964),  cranial  measurements  of  a  male  and 
female  from  Venezuela;  Starrett  and  de  la  Torre  (1964),  external  and  cranial  measure¬ 
ments  of  one  male  from  Nicaragua  and  two  males  and  a  female  from  Costa  Rica;  Tamsitt 
and  Valdivieso  (1963a),  external  measurements  (mean,  range)  of  28  specimens  and  cranial 
measurements  of  11  from  Colombia;  Tamsitt  and  Valdivieso  (19636),  external  measure¬ 
ments  (mean,  range)  of  four  males  from  Colombia;  Valdivieso  (1964),  external  and  cranial 
measurements  (mean,  range)  of  19  specimens  from  Colombia;  Brosset  (1965),  external  and 
cranial  measurements  of  three  males  from  Ecuador;  Jones  (1966),  forearm  and  cranial 
measurements  (range)  of  specimens  from  Guatemala;  Pirlot  (19656),  external  measure¬ 
ments  of  14  males  and  10  females  from  Est  du  Venezuela  and  19  males  and  15  females  from 
Zulia,  Venezuela;  Pirlot  (1968),  external  and  cranial  measurements  discussed  in  conjunction 
with  C.  castanea;  Goodwin  (1969),  forearm  and  cranial  measurements  of  nine  males 
and  three  females  from  Oaxaca;  Pine  (1972),  external  and  cranial  measurements  (mean,  range) 
of  males  and  females  throughout  the  range  of  the  species;  Pirlot  (1972),  external  measure¬ 
ments  of  a  specimen  from  Brazil;  Jones  et  al.  (1973),  external  and  cranial  measurements 
(mean,  range)  of  10  specimens  from  the  Yucatan  Peninsula;  Smith  and  Genoways  (1974), 
forearm  measurements  of  two  specimens  from  Margarita  Island,  Venezuela;  Taddei 
(19756),  external  measurements  (mean,  se,  range)  of  30  males  and  30  females,  and  cranial 
measurements  of  15  males  and  15  females  from  Brazil. 

Individual  variation. — In  specimens  from  Brazil,  coefficients  of  variation  for  external 
measurements  varied  in  30  males  from  2.70  to  6.15  and  in  30  females  from  2.70  to  5.94; 
CV  values  for  cranial  measurements  in  15  males  ranged  from  1.78  to  4.01  and  in  15  females 
from  1.85  to  4. 11  (Taddei,  19756).  According  to  Tamsitt  and  Valdivieso  (1963a),  specimens 
from  central  Colombia  were  homogeneous  in  size. 

Secondary  sexual  variation. — In  specimens  from  Brazil,  females  generally  averaged  larger 
than  males  in  external  measurements  and  in  four  (head  and  body,  ear,  forearm,  metacarpal 


42 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


II)  of  17  measurements  they  proved  to  be  significantly  so.  Cranial  measurements  showed  the 
opposite  in  one  of  15  measurements  (mastoid  breadth),  males  proved  to  be  significantly 
larger  than  females  (Taddei,  1 975 />).  Pine  (1972)  also  found  cranial  measurements  of  males 
to  average  slightly  larger  than  those  of  females.  However,  Tamsitt  and  Valdivieso  ( 1 963a) 
reported  that  their  males  and  females  were  of  the  same  size  in  a  sample  of  16  males  and  12 
females  from  Colombia. 

Geographic  variation. — Tamsitt  and  Valdivieso  (1963a)  found  individuals  from  localities 
on  each  side  of  the  East  Andes  not  to  differ  in  any  way.  Their  specimens,  although  slightly 
smaller,  did  not  differ  significantly  from  the  range  of  measurements  given  by  Hershkovitz 
(1949)  for  northern  Colombian  specimens.  According  to  Pine  (1972),  specimens  from  in 
and  around  the  Panama  drainage  are  characteristically  small.  Dalquest  (1951),  comparing 
forearm  length  and  cranial  measurements  of  his  specimens  from  Trinidad  with  examples 
from  San  Luis  Potosi,  concluded  that  they  are  alike  (however,  the  material  from  San  Luis 
Potosi  was  probably  C.  hrevicauda). 

Carollia  subrufa  (Hahn,  1905) 

Measurements  of  Carollia  subrufa  have  been  recorded  as  follows:  Hahn  (1905),  external 
and  cranial  measurements  of  the  male  holotype  from  Oaxaca;  Hahn  (1907),  external  measure¬ 
ments  of  eight  specimens  from  Oaxaca,  seven  from  Colima,  four  from  Campeche,  and  one 
from  Honduras,  and  cranial  measurements  of  nine  specimens  from  Oaxaca,  four  from 
Colima,  two  from  Campeche,  and  one  from  Honduras;  Elliot  (1917),  external  and  cranial 
measurements  of  the  holotype;  Goodwin  (1934),  external  measurements  of  one  specimen 
from  Guatemala;  Goodwin  (1942a),  external  and  cranial  measurements  of  two  males  from 
Honduras;  Felten  (1956a),  external  measurements  (mean,  range)  of  99  males  and  99  females 
and  cranial  measurements  of  27  males  and  33  females  from  El  Salvador  (as  a  subspecies  of 
C.  castanea)',  Felten  ( 1 956z/),  external  measurements  (mean,  range)  of  specimens  from  El 
Salvador;  Hall  and  Kelson  (1959),  external  measurements  (range)  of  198  (99  males,  99 
females)  specimens  from  El  Salvador  listed  as  C.  castanea ;  Ryan  (1960),  external  measure¬ 
ments  of  one  female  from  Guatemala;  Burt  and  Stirton  (1961),  external  and  cranial  measure¬ 
ments  of  four  males  from  El  Salvador  (as  a  subspecies  of  castanea );  Starrett  and  de  la  Torre 
(1964),  external  and  cranial  measurements  of  two  males  from  El  Salvador;  Jones  (1966), 
forearm  and  cranial  measurements  of  one  male  from  Jocotan,  Guatemala  (others  listed  are 
C.  brevicauda);  Villa-R.  (1967),  external  measurements  (mean,  se,  range)  of  51  males  and 
females  combined  and  cranial  measurements  (mean,  se,  range)  of  38  males  and  females 
combined;  Goodwin  (1969),  forearm  and  cranial  measurements  of  six  males  and  one  female 
from  Oaxaca,  (also  lists  a  male  and  a  female  from  Chiltepec,  but,  according  to  Pine,  1972, 
these  are  probably  C.  brevicauda)',  Pine  (1972),  external  measurements  of  one  male  from 
Colima,  two  males  and  eight  females  from  Oaxaca,  16  males  from  Chiapas,  one  male  and 
one  female  from  Honduras,  two  males  and  seven  females  from  Nicaragua,  and  cranial 
measurements  of  two  males  and  five  females  from  Colima,  two  males  and  eight  females 
from  Oaxaca,  16  males  and  24  females  from  Chiapas,  one  male  and  one  female  from 
Honduras,  and  two  males  and  seven  females  from  Nicaragua;  Watkins  et  al.  (1972), 
external  and  cranial  measurements  of  one  female  from  Jalisco. 

Geographic  variation. — Pine  (1972)  found  specimens  from  the  northern  part  of  the 
geographic  range  of  the  species  to  be  larger  than  those  of  the  southernmost  part  of  the 
geographic  range. 

Rhinophylla  alethina  Handley,  1966 

Handley  (1966a)  gave  external  measurements  (mean,  range)  of  six  males  and  four  females, 
and  cranial  measurements  of  the  male  holotype  from  Colombia. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


43 


Rhinophylla  fischerae  Carter,  1966 

Measurements  of  Rhinophylla  fischerae  have  been  recorded  as  follows:  Carter  (1966), 
external  and  cranial  measurements  of  the  female  holotype  from  Peru,  six  additional  females 
and  two  males,  all  from  the  type  locality  except  one  female  from  Pucallpa,  Peru;  Marinkelle 
and  Cadena  (1972),  external  measurements  of  a  male  and  female  from  Colombia;  Mumford 
(1975),  external  and  cranial  measurements  of  an  unsexed  specimen  from  Ecuador. 

Rhinophylla  pumilio  Peters,  1865 

Measurements  of  Rhinophylla  pumilio  have  been  recorded  as  follows:  Peters  (1865  a), 
external  measurements  of  the  holotype  from  Brazil;  Dobson  ( 1 878«),  external  measurements 
of  one  specimen  from  Brazil;  Sanborn  (1936),  external  and  cranial  measurements  of  a  male 
and  female  from  Ecuador;  Husson  (1962),  external  and  cranial  measurements  of  two  females 
from  Surinam  and  two  from  Guyana;  Hill  (1964),  forearm  measurements  of  two  males  and 
cranial  measurements  of  one  of  these  from  Guyana;  Carter  (1966),  external  measurements 
of  15  males  and  10  females  combined,  and  cranial  measurements  (mean,  range)  of  15  males 
and  13  females  combined  from  Venezuela,  Brazil,  Ecuador,  and  Peru;  Marinkelle  and 
Cadena  (1972),  forearm  and  cranial  measurements  of  a  male  (juvenile)  and  the  range  of  three 
females  from  Colombia. 


Subfamily  Stenoderminae 
Ametrida  centurio  Gray,  1847 

Measurements  of  Ametrida  centurio  have  been  recorded  as  follows:  Peters  ( 1 866c/), 
external  measurements  of  one  specimen;  Dobson  (1878a),  external  measurements  of  the 
female  holotype  from  Brazil;  H.  Allen  (18946),  external  and  cranial  measurements  of  the 
male  holotype  of  A.  minor  from  Surinam  (type  locality  according  to  Peterson,  1965)  and 
external  measurements  of  a  specimen  of  A.  centurio ;  Sanborn  (1938),  external  and  cranial 
measurements  of  a  male  (female  according  to  Peterson,  19656)  from  Brazil;  Husson  (1960), 
cranial  measurements  of  one  specimen  from  Bonaire;  Goodwin  and  Greenhall  (1961), 
forearm  and  cranial  measurements  of  a  male  from  Guyana,  a  female  from  Venezuela,  and 
a  subadult  from  Trinidad;  Husson  (1962),  external  and  cranial  measurements  of  two  males 
and  two  females  (see  Peterson,  19656:3-4,  on  the  question  of  the  sex  of  one  of  these  specimens) 
from  Surinam  and  one  male  from  Bonaire;  Peterson  (1965),  forearm  and  cranial  measure¬ 
ments  of  12  males  from  Brazil,  Guyana,  Surinam,  Venezuela,  Trinidad,  and  Bonaire 
(including  the  holotype  of  A.  minor  from  Surinam),  13  females  from  Brazil,  Guyana, 
Venezuela,  Trinidad,  and  Surinam  (including  the  holotype  of  A.  centurio  from  Brazil),  and 
external  measurements  (mean,  range)  of  males  and  females. 

Secondary  sexual  variation. — Peterson  (19656)  described  distinct  differences  in  size 
between  the  sexes  with  no  overlap  in  forearm  length  or  the  following  cranial  measure¬ 
ments:  condylobasal  length;  least  interorbital  width;  breadth  of  palate  (Ml-Ml);  toothrow 
length  (C-M3). 


Ardops  nichollsi  (Thomas,  1891) 

Measurements  of  Ardops  nichollsi  have  been  recorded  as  follows:  Thomas  (1891a), 
external  and  cranial  measurements  of  the  female  holotype  of  A.  n.  nichollsi  from  Dominica; 
Thomas  (1894),  external  and  cranial  measurements  of  the  male  holotype  of  A.  n.  montser- 
ratensis  from  Montserrat;  Elliot  (1904),  external  and  cranial  measurements  of  one  specimen 
from  Monsterrat,  one  from  Dominica,  and  one  from  St.  Lucia;  Miller  (1902a),  external 
and  cranial  measurements  of  the  female  holotype  of  A.  n.  luciae  from  St.  Lucia  and  of  a 
male  from  Dominica;  Miller  (1913a),  external  and  cranial  measurements  of  the  female 


44 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


holotype  of  A.  n.  annectens  and  a  male  from  Guadeloupe;  Elliot  (1917),  external  and 
cranial  measurements  of  the  holotype  of  A.  n.  annectens ,  G.  M.  Allen  (1942),  forearm  length 
of  taxa  described  at  that  time;  Hall  and  Kelson  (1959),  externa]  and  cranial  measurements  of 
the  holotypes  of  A.  n.  monsterratensis,  A.  n.  annectens ,  and  A.  n.  luciae;  Jones  and  Schwartz 
(1967),  forearm  and  cranial  measurements  of  the  female  holotype  of  A.  n.  nichollsi,  external 
measurements  (mean,  range)  of  six  males  and  seven  females,  cranial  measurements  (mean, 
range)  of  eight  males  and  seven  females  from  Dominica,  external  and  cranial  measurements 
of  a  male  and  a  female  from  St.  Eustatius,  and  the  male  holotype  of  A.  n.  monserratensis 
from  Montserrat,  and  the  female  holotype  of  A.  n.  luciae ,  cranial  measurements  of  a  female, 
forearm  measurements  of  one  male  and  four  females  from  St.  Lucia,  external  measurements 
of  an  adult  male  and  the  female  holotype  of  A.  n.  annectens ,  cranial  measurements  of  the 
holotype,  two  males,  and  two  females,  forearm  measurements  of  four  females  from 
Guadeloupe,  external  measurements  of  the  female  holotype  (A.  n.  koopmani),  another 
female,  and  two  males,  and  cranial  measurements  of  the  female  holotype  and  a  male  from 
Martinique;  Jones  and  Genoways  (1973),  some  measurements  as  given  by  Jones  and 
Schwartz  (1967). 

Secondary  sexual  variation. — In  individuals  from  Dominica,  females  were  clearly  larger 
than  males.  This  was  also  found  to  be  true  in  one  male  and  one  female  from  Martinique 
(Jones  and  Schwartz,  1967). 

Geographic  variation. — According  to  Jones  and  Schwartz  (1967),  specimens  from 
Dominica  were  the  smallest  of  the  species,  whereas  those  from  St.  Eustatius  and  Montserrat 
were  the  largest.  Specimens  from  Martinique  differed  from  those  on  adjacent  islands, 
Dominica  to  the  north  and  St.  Lucia  to  the  south,  in  being  considerably  larger. 

Ariteus  flavescens(Gray,  1831) 

Measurements  of  Ariteus  flavescens  have  been  recorded  as  follows:  Peters  (1876), 
external  measurements  of  a  specimen  of  Peltorhinus  achradophilus  (=  A.  flavescens ); 
Dobson  (1878c/),  external  measurements  of  the  female  holotype  of  Ariteus  achradophilus 
from  Jamaica;  Elliot  (1904),  external  and  cranial  measurements  of  one  specimen  from 
Jamaica;  G.  M.  Allen  (1942),  external  measurements  for  the  species;  Howe  (1974),  external 
measurements  of  two  males  and  two  females  from  Jamaica. 

Artibeus  aztecus  Andersen,  1906 

Measurements  of  Artibeus  aztecus  have  been  recorded  as  follows:  Andersen  (19066), 
external  measurements  of  the  male  holotype  of  A.  aztecus  from  Morelos;  Andersen  (1908), 
external  and  cranial  measurements  (range)  of  four  specimens  from  Morelos;  Elliot  (1917), 
cranial  measurements  of  the  holotype;  Dalquest  (1953c/),  external  measurements  of  a  male 
and  two  females  and  cranial  measurements  of  the  male  and  one  female  from  San  Luis  Potosf; 
Lukins  and  Davis  (1957),  forearm  and  cranial  measurement  (range)  for  the  species;  Villa-R. 
(1967),  external  and  cranial  measurements  of  one  female  from  the  state  of  Mexico;  Koopman 
(1961),  forearm  and  cranial  measurements  (range)  of  four  specimens  (one  male,  three  females) 
from  Sinaloa;  Baker  and  Greer  (1962),  external  and  cranial  measurements  of  a  female  from 
Durango;  Alvarez  (1963),  external  and  cranial  measurements  of  three  males  and  one  female 
from  Tamaulipas;  Jones  (1964),  forearm  and  cranial  measurements  (mean,  range)  of  15 
specimens  (10  males  and  five  females)  from  Sinaloa;  Davis  (1969),  external  and  cranial 
measurements  (mean,  range)  of  33  specimens  from  the  Mexican  highlands,  41  from  the 
Guatemalan  highlands,  and  18  from  the  Costa  Rican  highlands,  and  external  and  cranial 
measurements  of  the  male  holotype  of  A.  aztecus  aztecus  from  Morelos,  the  male  holotype 
of  A.  a.  minor  from  Guatemala,  and  the  male  holotype  of  A.  a.  major  from  Costa  Rica; 
Goodwin  (1969),  forearm  and  cranial  measurements  of  four  males  and  five  females  from 
Oaxaca;  Alvarez  and  Ramirez-Pulido  (1972),  external  and  cranial  measurements  of  two 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


45 


males  from  Michoacan,  and  a  female  from  Oaxaca;  Jones  et  al.  (1972),  forearm  and  cranial 
measurements  as  given  by  Jones  (1964). 

Geographic  variation. — Artiheus  aztecus,  which  occurs  in  the  Middle  American  high¬ 
lands,  was  segregated  into  three  recognizable  populations — aztecus  in  the  Mexican  highlands, 
minor  from  the  Guatemalan  highlands,  and  major  of  the  Costa  Rican  highlands.  With  regard 
to  size,  A.  a.  major  is  the  largest,  and  minor  is  the  smallest  (Davis,  1969). 

Artibeus  cinereus  (Gervais,  1855) 

Measurements  of  Artibeus  cinereus  have  been  recorded  as  follows:  Peters  ( 1 865 r/), 
external  measurements  of  the  holotype  of  A.  quadrivittatum  from  Surinam;  Dobson 
( 1 878«),  external  measurements  of  a  male  and  a  female;  Robinson  and  Lyon  (1901),  external 
measurements  of  three  males  and  six  females  from  Venezuela;  Andersen  (19066),  cranial 
measurements  (range)  of  eight  specimens  including  the  male  holotype  (Colombia)  of  A. 
cinereus  bogotensis  from  Colombia  and  Venezuela  and  seven  additional  specimens  of  A.  c. 
cinereusr,  Andersen  (1908),  external  measurements  (mean,  range)  of  10  specimens  and 
cranial  measurements  (mean,  range)  of  eight  from  Guyana,  Trinidad,  and  Venezuela,  external 
and  cranial  measurements  (mean,  range)  of  eight  specimens  from  Colombia  and  Venezuela 
and  the  range  of  these  measurements  in  three  specimens  of  A.  quadrivittatus  from  Surinam; 
Lima  (1926),  external  measurements  of  a  male  from  Brazil;  Sanborn  (1932),  forearm  measure¬ 
ments  of  a  female  and  a  specimen  of  unknown  sex  and  cranial  measurements  of  the  female 
from  Bolivia;  Cunha  Vieira  (1942),  external  measurements  of  two  females  from  Venezuela 
and  external  measurements  of  a  male  from  Ecuador;  Hershkovitz  (1949),  external  and  cranial 
measurements  of  a  female  from  Colombia;  Goodwin  and  Greenhall  (1961),  forearm  and 
cranial  measurements  of  three  males  and  one  female  from  Trinidad;  Burt  and  Stirton  (1961), 
forearm  and  cranial  measurements  (range)  of  four  males  and  14  females  from  El  Salvador; 
Husson  (1962),  external  and  cranial  measurements  of  three  males,  four  females,  and  the 
unsexed  holotype  of  A.  quadrivittatus  from  Surinam;  Tamsitt  and  Valdivieso  (1963 a), 
external  measurements  of  four  females  from  Colombia;  Brosset  (1965),  external  and  cranial 
measurements  of  a  male  from  Ecuador;  Tamsitt  and  Valdivieso  (1966a),  forearm  and  cranial 
measurements  of  a  male  and  female  from  Colombia  (values  for  the  female  as  given  by 
Hershkovitz,  1949);  Davis  (19706),  external  and  cranial  measurements  (mean,  range)  of 
36  specimens  from  Trinidad;  Tuttle  (1970),  forearm  measurements  (range)  of  specimens 
from  east  of  the  Andes  in  Peru;  Pirlot  (1972),  external  measurements  of  two  males  and  one 
female  from  Brazil  (type  description  of  A.  c.  solimoesi ). 

Artibeus  eoncolor  Peters,  1865 

Measurements  of  Artibeus  eoncolor  have  been  recorded  as  follows:  Peters  (1865a), 
external  measurements  of  the  holotype  from  Surinam;  Thomas  (1892),  forearm  and  cranial 
measurements  of  the  holotype;  Andersen  (1908),  external  and  cranial  measurements  of  a 
female  from  Surinam  and  cranial  measurements  of  the  holotype  from  Surinam;  Cabrera 
(1917),  external  and  cranial  measurements  of  a  female  possibly  from  Brazil;  Cunha  Vieira 
(1942),  external  measurements  based  on  Andersen  (1908);  Husson  (1962),  external  and 
cranial  measurements  of  a  female  as  given  by  Andersen  (1908)  and  measurements  of  the 
holotype  as  given  by  Peters  and  Thomas;  Hill  (1964),  forearm  and  cranial  measurements 
of  one  male  from  Guyana;  Linares  (1969),  external  measurements  of  a  male  and  two  females 
from  Venezuela;  Gardner  (1976),  external  and  cranial  measurements  of  a  male  from  Peru. 

Artibeus  glaucus  Thomas,  1893 

Measurements  of  Artibeus  glaucus  have  been  recorded  as  follows:  Thomas  (1893),  external 
and  cranial  measurements  of  the  female  holotype  from  Peru;  Andersen  (1908),  external  and 
cranial  measurements  of  the  holotype  from  Peru;  Davis  (1970a),  cranial  measurements 
(mean,  range)  of  nine  specimens  from  Peru  and  Ecuador. 


46 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Artibeus  hirsutus  Andersen,  1906 

Measurements  of  Artibeus  hirsutus  have  been  recorded  as  follows:  Andersen  (19066), 
forearm  and  cranial  measurements  (range)  of  eight  specimens  from  Michoacan,  Colima, 
and  Jalisco;  Andersen  (1908),  external  and  cranial  measurements  (mean,  range)  of  eight 
specimens  from  Michoacan,  Colima,  and  Jalisco;  Elliot  (1917),  cranial  measurements  of  the 
holotype;  Davis  and  Russell  (1952),  external  and  cranial  measurements  of  one  male  and 
five  females  (mean,  range)  from  Morelos;  Anderson  (1960),  external  and  cranial  measure¬ 
ments  (mean,  range)  of  28  specimens  from  Guerrero;  Davis  and  Carter  (1964),  external  and 
cranial  measurements  (mean,  range)  of  six  females;  Villa-R.  (1967),  external  measurements 
(mean,  sd,  range)  of  55  specimens  and  cranial  measurements  of  46  specimens  from  Sonora, 
Sinaloa,  Nayarit,  Jalisco,  Morelos,  and  Guerrero;  Genoways  and  Jones  (1968),  forearm 
measurements  (mean,  range)  of  four  young  males  and  four  females  from  Zacatecas;  Goodwin 
(1969),  forearm  and  cranial  measurements  of  two  males  from  Guerrero  and  two  from 
Sonora;  Anderson  (1972),  external  and  cranial  measurements  of  three  specimens  from 
Chihuahua;  Jones  et  al.  (1972),  forearm  and  cranial  measurements  (mean,  range)  of  10 
specimens  (five  males  and  five  females)  from  Sinaloa. 

Secondary  sexual  variation. — Anderson  (1960)  found  no  significant  size  differences 
between  sexes  in  four  external  and  four  cranial  measurements  in  a  sample  of  28  specimens 
from  Guerrero. 


Artibeus  inopinatus  Davis  and  Carter,  1964 

Davis  and  Carter  (1964)  reported  external  and  cranial  measurements  (mean,  range)  of 
eight  females  from  Honduras  and  forearm  measurements  of  one  male  from  Honduras  and 
one  from  Nicaragua.  Although  Davis  and  Carter  did  not  examine  the  specimens  reported 
from  El  Salvador  by  Burt  and  Stirton  (1961),  under  the  name  Artibeus  hirsutus,  they  judged, 
and  we  agree,  from  the  published  measurements  that  the  specimens  are  referable  to  A. 
inopinatus. 


Artibeus  jamaicensis  Leach,  1821 

Measurements  of  Artibeus  jamaicensis  have  been  recorded  as  follows:  Saussure  (18606), 
external  measurements  of  one  specimen;  Gundlach  (1872,  1877),  external  measurements  of 
a  specimen  from  Cuba;  Dobson  (1878a),  external  measurements  for  a  male  of  A.  perspicillatus 
from  Guatemala  and  a  female;  Cope  (1889),  external  measurements  of  one  male  cotype 
of  Dermanura  eva  from  St.  Martin,  Lesser  Antilles;  H.  Allen  (1894a),  external  measurements 
from  three  specimens  (two  from  Mexico,  one  locality  unknown)  and  cranial  measurements 
(mean)  of  three  specimens  from  an  unspecified  locality;  J.  A.  Allen  and  Chapman  (1897a), 
forearm  measurements  of  four  specimens  from  Yucatan,  10  from  Jamaica,  31  females  and 
20  males  from  Cuba;  Rehn  (1900),  cranial  measurements  of  the  two  male  cotypes  of 
Dermanura  eva  Cope  from  St.  Martin,  Lesser  Antilles,  a  specimen  from  Jamaica,  and  one 
from  Brazil;  Robinson  and  Lyon  (1901),  external  measurements  of  a  male  and  two  females 
from  Venezuela;  Rehn  (19026),  external  measurements  of  the  unsexed  holotype  of  A.  hercules 
(  =  A.  jamaicensis)  and  the  mean  of  external  measurements  for  two  additional  specimens, 
cranial  measurements  of  a  specimen  from  Peru,  external  measurements  of  the  male  holotype, 
the  mean  for  six  specimens  of  A.  parvipes  (  —  A.  jamaicensis)  from  Cuba,  and  one  specimen 
of  A.  jamaicensis  from  Jamaica,  the  mean  of  six  specimens  and  external  measurements 
(mean)  of  two  specimens  of  A.  planirostris  and  cranial  measurements  of  one  from  Brazil; 
J.  A.  Allen  (1904),  external  and  cranial  measurements  of  the  male  holotype  of  A.  insularis 
from  St.  Kitts,  Lesser  Antilles,  and  the  male  holotype  of  A.  j.  yucatanicus  from  Yucatan; 
Elliot  (1904),  external  and  cranial  measurements  of  one  specimen  each  of  A.  coryi,  A. 
jamaicensis,  A.j.  parvipes,  and  A.  j.  planirostris;  Miller  (1904),  external  measurements  of  12 
males  and  13  females  from  Cuba;  Elliot  (1905a),  external  and  cranial  measurements  of  a 
specimen  from  St.  Kitts  Island,  Lesser  Antilles;  Andersen  (1906),  cranial  measurements 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


47 


(mean)  of  65  specimens  of  A.  j.  jamaicensis  and  external  measurements  (range)  of  three 
specimens  of  A.  j.  praeceps  from  Guadeloupe;  G.  M.  Allen  (1908),  external  measurements 
of  three  specimens  and  cranial  measurements  of  one  male  from  Brazil,  and  external  measure¬ 
ments  of  one  specimen  from  Jamaica;  J.  A.  Allen  (1908u),  forearm  measurements  (range) 
of  four  specimens  from  the  Dominican  Republic;  J.  A.  Allen  (19086),  external  and  cranial 
measurements  of  the  male  holotype  of  A.  j.  richardsoni  from  Nicaragua;  Andersen  (1908), 
external  and  cranial  measurements  (range)  of  16  specimens  (11  cranial)  from  Brazil,  three 
from  Venezuela,  and  three  from  Chiapas  and  Guerrero,  median  and  range  of  the  above 
combined,  13  specimens  (nine  cranial)  from  Trinidad  and  Tobago,  nine  (eight  cranial)  from 
Grenada,  41  (33  cranial)  from  Surinam,  Cayenne,  Guyana,  and  Lower  Orinoco,  25  speci¬ 
mens  (12  cranial)  from  Cuba,  14  (12  cranial)  from  Yucatan  and  Cozumel  Island,  12  (nine 
cranial)  from  Central  America,  27  (23  cranial)  from  southern  Mexico,  21  (11  cranial)  from 
Puerto  Rico,  three  from  Dominican  Republic,  one  from  St.  Kitts  Island,  eight  (five  cranial) 
from  St.  Andrews  and  Old  Providence  Island,  and  95  (65  cranial)  (median,  range)  of  A.  j. 
jamaicensis  (including  much  of  the  above  data);  Elliot  (1917),  external  and  cranial  measure¬ 
ments  of  the  holotype;  Anthony  (1919),  cranial  measurements  of  fossil  material  from  Cuba; 
Anthony  (1924«),  external  and  cranial  measurements  of  the  female  holotype  of  A.  j.  frater- 
culus  from  Ecuador,  forearm  measurements  (mean)  of  18  specimens  and  cranial  measure¬ 
ments  (mean,  range)  of  13  others;  Anthony  (1918,  1925),  external  measurements  (mean, 
range)  of  24  specimens  and  cranial  measurements  (mean,  range)  of  10  specimens  (five 
males,  five  females)  from  Puerto  Rico;  Goodwin  (1934),  external  measurements  of  one 
specimen  from  Guatemala;  Sanborn  (1936),  forearm  measurements  (range)  of  three  males 
and  four  females  and  cranial  measurements  (range)  of  three  specimens  (one  male,  two  fe¬ 
males)  from  Barbados;  Martinez  and  Villa-R.  (1938),  external  measurements  of  five  males 
and  nine  females  from  Morelos;  Cunha  Vieira  (1942),  external  and  cranial  measurements 
of  a  male  from  Brazil;  Goodwin  (1942),  forearm  and  cranial  measurements  of  two  males 
from  Honduras,  and  these  measurements  of  another  specimen;  Goodwin  (1946),  external 
and  cranial  measurements  (range)  for  the  species;  Hall  and  Villa-R.  (1949),  external  and 
cranial  measurements  of  one  female  from  Michoacan;  Hershkovitz  (1949),  external  and 
cranial  measurements  of  a  male  and  female  (two  males  and  a  female  for  forearm)  from 
Colombia;  Dalquest  (1951),  forearm  and  cranial  measurements  (mean)  of  four  males  and 
eight  females  from  Trinidad;  Dalquest  ( 1 95 3  zz),  external  measurements  (mean)  of  eight  males 
and  eight  females  and  cranial  measurements  (mean)  of  two  males  and  1 1  females  from  San 
Luis  Potosf;  Goodwin  (1953),  forearm  and  cranial  measurements  of  the  male  holotype  of 
A.  coryi  from  St.  Andrews  Island,  the  male  holotype  of  A.  insular  is  from  St.  Kitts,  the  male 
holotype  of  A.  j.  richardsoni  from  Nicaragua,  the  male  holotype  of  A.  j.  yucatanicus 
from  Yucatan,  and  the  female  holotype  of  A.  j.  fraterculus  from  Ecuador;  de  la  Torre 
(1955),  forearm  measurements  (mean,  range)  of  five  specimens  (three  males,  two  fe¬ 
males)  from  Jalisco;  de  la  Torre  (1954),  external  and  cranial  measurements  (mean, 
range)  of  23  specimens  from  Tamaulipas;  Felten  (1956u),  external  measurements  (mean, 
range)  of  16  males  and  five  females  and  cranial  measurements  of  nine  males  (mean, 
range)  and  one  female  from  El  Salvador;  Felten  (1956 d),  external  measurements  (mean, 
range)  of  specimens  from  El  Salvador;  Anderson  (1960),  external  and  cranial  measure¬ 
ments  (range)  of  three  specimens  from  Sinaloa,  and  four  from  Jalisco;  Husson  (1960), 
cranial  measurements  (mean,  range)  of  specimens  from  Curasao  and  St.  Martin;  Burt 
and  Stirton  (1961),  forearm  and  cranial  measurements  (range)  of  44  specimens  (18 
males,  26  females)  from  El  Salvador;  Goodwin  and  Greenhall  (1961),  forearm  measure¬ 
ments  (range)  of  12  males  and  18  females,  and  cranial  measurements  of  one  male  and  one 
female  from  Trinidad;  Baker  and  Greer  (1962),  external  and  cranial  measurements  of  a  male 
and  female  from  Durango;  Pirlot  (1963),  forearm  measurements  (range)  of  35  males  and 
20  females  from  Venezuela;  Tamsitt  and  Valdivieso  (1963«),  external  measurements  of  one 
male  and  three  females  and  cranial  measurements  of  one  female  from  Colombia;  Davis  and 
Carter  (1964),  external  and  cranial  measurements  (mean,  range)  of  eight  females  from  Central 
America;  Hill  (1964),  forearm  measurements  of  two  males  and  three  females  and  cranial 


48 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


measurements  of  two  males  and  three  females  and  cranial  measurements  of  two  males 
and  two  females  from  Guyana;  Valdivieso  (1964),  external  measurements  of  one  male  and 
two  females  and  cranial  measurements  of  one  female  from  Colombia;  Starrett  and  de  la 
Torre  (1964),  external  and  cranial  measurements  of  one  male  from  Nicaragua  and  one  from 
Costa  Rica;  Handley  (1965),  external  and  cranial  measurements  of  the  female  holotype  of 
A.  j.  triomylus  from  Guerrero  and  mean  and  range  of  external  measurements  of  10  males 
and  nine  females  and  cranial  measurements  of  12  females  and  10  males  from  Guerrero; 
Pirlot  (19656),  external  measurements  of  15  males  and  33  females  from  Est  du  Venezuela 
and  of  35  males  and  20  females  from  Zulia,  Venezuela;  Villa-R.  (1967),  external  measure¬ 
ments  of  46  specimens  and  cranial  measurements  of  43  specimens  of  A.  j.  triomylus  from 
Mexico,  and  external  measurements  of  76  specimens  and  cranial  measurements  of  71 
specimens  of  A.  j.  yucatanicus  from  Mexico;  Genoways  and  Jones  (1968),  mean  and  range 
of  forearm  measurements  of  six  young  specimens  (two  males,  four  females)  and  individual 
forearm  measurements  of  two  young  males  and  one  young  female  from  Zacatecas;  Koopman 
(1968),  forearm  and  cranial  measurements  of  the  holotype  of  A.  praeceps  (Guadeloupe) 
and  specimens  (range)  from  Guadeloupe  and  Dominica;  Pirlot  (1968),  forearm  measure¬ 
ments  of  a  female  from  Peru;  Goodwin  (1969),  forearm  and  cranial  measurements  of  four 
males  and  three  females  of  A.  j.  yucatanicus  from  Oaxaca  and  three  males  and  three  females 
of  A.  j.  triomylus  from  Oaxaca;  Jones  and  Phillips  (1970),  forearm  measurements  (mean, 
range)  of  seven  specimens  from  Barbados,  11  from  St.  Lucia,  20  from  St.  Vincent,  23  from 
Grenada,  and  16  from  Trinidad,  and  cranial  measurements  for  7,  15,  32,  15,  and  1 1  specimens, 
respectively;  Davis  (19706),  external  and  cranial  measurements  of  the  male  holotype  of  A. 
j.  richardsoni  from  Nicaragua,  mean  and  range  of  13  topotypes,  means  of  14  from  Chiapas, 
12  from  Guatemala  (Alta  Verapaz),  20  from  Guatemala  (Puerto  Barrios),  20  from  Nicaragua 
(Castillo),  20  from  Honduras  (coastal),  16  from  Costa  Rica  (coastal),  20  from  Panama 
(Veraguas),  21  from  Panama  (Chepo)  of  A.  j.  richardsoni ,  external  and  cranial  measure¬ 
ments  of  the  male  holotype  of  A.  j.  yucatanicus  from  Yucatan,  mean  and  range  of  eight 
topotypes,  mean  of  18  from  Tamaulipas,  25  from  San  Luis  Potosf,  19  from  Veracruz,  14 
from  Campeche  and  Yucatan,  four  from  British  Honduras,  20  from  Honduras  (Bay  Islands) 
of  A.  j.  yucatanicus,  forearm  and  cranial  measurements  of  the  female  holotype  of  A.  j.  trio¬ 
mylus  from  Guerrero,  mean  and  range  of  20  from  near  the  type  locality,  external  and  cranial 
measurements  of  the  female  holotype  of  A.  j.  paulus  from  El  Salvador,  means  of  15  from 
Chiapas  (below  1000  feet),  20  from  Guatemala,  20  from  El  Salvador,  20  from  Honduras 
(Nueva  Ocotepeque),  six  from  Honduras  (Pacific  lowlands),  11  from  Nicaragua  (San 
Antonio),  and  four  from  Costa  Rica  (Guanacaste  Lowlands)  of  A.  j.  paulus ;  Tuttle  (1970), 
cranial  measurements  of  a  female  from  Peru,  and  range  in  forearm  length  of  specimens  east 
of  the  Andes;  Jones  et  al.  (1972),  forearm  and  cranial  measurements  (mean,  range)  of  10 
specimens  (five  males,  five  females)  from  Sinaloa;  Smith  and  Genoways  (1974),  forearm  and 
cranial  measurements  (mean,  range)  from  four  localities  in  Venezuela  (sample  sizes  five, 
22,  17,  22)  and  eight  specimens  from  Trinidad. 

Age  variation. — According  to  Davis  (19706),  young  individuals  in  which  the  cartilaginous 
epiphyses  of  finger  joints  were  readily  discernable  were  consistantly  smaller  than  adults  in 
all  measurements.  However,  individuals  in  which  the  joint  of  the  finger  was  only  swollen  and 
in  which  the  epiphyses  and  diaphyses  appeared  to  be  united  were  as  large  as  adults  in  all 
measurements. 

Individual  variation. — Within  sample  variation  of  cranial  measurements  was  shown  by 
Davis  (19706)  to  be  usually  less  than  10  per  cent  of  the  minimum  value  of  each  variate  tested. 
Of  six  cranial  measurements  tested,  length  of  skull  was  the  least  variable  and  breadth  across 
upper  molars  the  most.  Wing  measurements  varied  more  than  cranial.  Of  four  wing 
measurements  examined,  length  of  forearm  was  the  least  variable  and  length  of  phalanx  1, 
digit  III  the  most. 

Secondary  sexual  variation. — Davis  (19706)  found  no  significant  secondary  sexual 
variation  in  four  wing  and  eight  cranial  measurements. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


49 


Geographic  variation. — Both  Koopman  (1968)  and  Jones  and  Phillips  (1970)  noted  a  trend 
toward  slightly  larger  size  in  specimens  from  the  southern  part  of  the  Lesser  Antilles.  Jones 
and  Phillips  (1970)  found  A.  jamaicensis  from  Grenada  to  approach  those  from  Trinidad 
and  Tobago  in  size.  They  also  found  that  specimens  from  St.  Vincent  averaged  considerably 
larger  than  specimens  from  any  other  Antillean  population. 

Davis  (19706),  studying  geographic  variation  in  Middle  American  populations  of  Artibeus 
jamaicensis,  recognized  four  areas  of  differentiation.  The  largest  individuals  occurred  along 
the  Atlantic  versant  of  Middle  America  (northern  Chiapas  to  eastern  Panama).  Greatest 
length  of  skull  in  this  area  averaged  near  29  and  forearm  near  61.  The  population  along  the 
Atlantic  versant  of  Mexico  (Tamaulipas  to  the  Yucatan  Peninsula  and  into  British  Honduras 
and  on  the  Bay  Islands  of  Honduras)  was  characterized  by  small  size.  More  than  90 
per  cent  of  the  individuals  had  a  skull  length  of  less  than  28.45  combined  with  a  zygomatic 
breadth  of  less  than  17.05.  Populations  from  the  Pacific  versant  were  also  characterized 
by  small  size — those  from  Oaxaca  and  Morelos  northward  into  Sinaloa  and  Durango  normally 
possessed  three  upper  molars  and  had  a  zygomatic  breadth  seldom  less  than  17.0.  Populations 
from  Chiapas  southward  to  Guanacaste,  Costa  Rica,  lacked  the  upper  third  molar. 

Smith  and  Genoways  (1974)  found  their  material  from  Margarita  Island,  Venezuela, 
averaged  slightly  smaller  in  external  and  cranial  measurements  than  specimens  from  the 
adjacent  Venezuelan  mainland  and  Trinidad. 

Artibeus  lituratus(01fers,  1818) 

Measurements  of  Artibeus  lituratus  have  been  recorded  as  follows:  J.  A.  Allen  and 
Chapman  (18976),  external  measurements  of  the  male  holotype  of  A.  I.  palmarum  from 
Trinidad  and  a  female,  mean  external  measurements  for  five  females,  and  cranial  measure¬ 
ments  of  one  female  from  Trinidad;  J.  A.  Allen  (1897),  external  and  cranial  measurements 
of  the  male  holotype  of  A.  lituratus  intermedins  from  Costa  Rica;  Bangs  (1899),  external 
and  cranial  measurements  of  the  male  holotype  of  Artibeus  femurvillosum  from  Colombia; 
Robinson  and  Lyon  (1901),  external  measurements  of  five  males  and  15  females  from 
Venezuela;  Rehn  (19026),  external  measurements  of  the  holotype  of  A.  I.  Hercules  from 
Peru,  the  average  of  these  measurements  for  two  additional  specimens  and  cranial  measure¬ 
ments  for  one;  J.  A.  Allen  (1904),  external  and  cranial  measurements  of  the  male  holotype  of 
A.  rusbyi  from  Peru;  Elliot  (1904),  external  and  cranial  measurements  of  a  specimen  of  A. 
lituratus  intermedins ;  G.  M.  Allen  (1908),  external  measurements  of  three  specimens  and 
cranial  measurements  of  one  from  Brazil  and  forearm  measurements  of  the  holotype  of  A.  I. 
intermedins  and  three  additional  specimens  from  Costa  Rica;  Andersen  (1908),  external 
and  cranial  measurements  (mean,  range)  of  12  specimens  (six  cranial)  from  Paraguay,  20 
(19  cranial)  from  Brazil,  and  nine  (eight  cranial)  from  Ecuador  and  Colombia,  means  for 
these  measurements  for  15  specimens  (10  cranial)  from  Venezuela,  four  (three  cranial)  from 
Trinidad  and  St.  Vincent,  20  (15  cranial)  from  Central  America  (Panama,  Costa  Rica, 
Nicaragua,  Guatemala),  four  (three  cranial)  from  Mexico  (Veracruz,  Jalisco,  Oaxaca) 
and  a  mean  for  these  measurements  from  the  latter  localities,  cranial  measurements  of  six 
specimens  of  A.  I.  aequatorialis  from  Ecuador,  and  external  of  seven  and  cranial  measure¬ 
ments  of  six  specimens  (median,  range)  of  A.  1.  aequatorialis  from  Ecuador  and  Colombia; 
Lima  (1926),  external  measurements  of  a  male  and  cranial  measurements  of  an  unsexed 
individual  from  Brazil;  Cunha  Vieira  (1942),  external  measurements  of  one  male  and  four 
females  and  cranial  measurements  of  three  males  from  Brazil;  Goodwin  (1942u),  external 
and  cranial  measurements  of  two  females  from  Honduras;  Hershkovitz  (1949),  external  and 
cranial  measurements  (range)  of  specimens  from  Colombia;  Dalquest  (1950),  cranial  measure¬ 
ments  (mean)  of  three  males  and  two  females  from  San  Luis  Potosi;  Dalquest  (195  1),  forearm 
and  cranial  measurements  (mean)  of  three  males  and  six  females  from  Trinidad;  Dalquest 
(1953u),  external  measurements  of  a  male  and  two  females  (mean)  and  cranial  measurements 
(mean)  of  three  males  and  two  females  from  San  Luis  Potosi;  Goodwin  (1953),  forearm  and 
cranial  measurements  of  the  male  holotype  of  A.  lituratus  palmarum  from  Trinidad,  the 


50 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


male  holotype  of  A.  lituratus  intermedins  from  Costa  Rica,  and  the  male  holotype  of  A.  rushy i 
from  Peru;  de  la  Torre  (1954),  external  and  cranial  measurements  of  three  specimens  from 
Tamaulipas;  Felten  (1956c),  external  measurements  (mean,  range)  of  six  males  and  six 
females  and  cranial  measurements  of  five  males  and  five  females  from  El  Salvador;  Felten 
( 1956c/),  external  measurements  of  specimens  from  El  Salvador;  Russell  (1956),  forearm 
and  cranial  measurements  of  a  female  from  Morelos;  Lukens  and  Davis  (1957),  forearm 
and  cranial  measurements  (mean,  range)  of  adult  specimens,  one  juvenile  female,  and  a 
subadult  female  from  Guerrero;  Anderson  (1960),  external  and  cranial  measurements  (mean, 
range)  of  17  specimens  from  Sinaloa;  Goodwin  and  Greenhall  (1961),  forearm  measurements 
(range)  of  14  males  and  18  females  and  cranial  measurements  of  one  male  from  Trinidad; 
Tamsitt  and  Valdivieso  (1963a),  external  and  cranial  measurements  (mean,  range)  of  46 
males  and  30  females  combined  from  Colombia;  Tamsitt  and  Valdivieso  (19636),  external 
measurements  of  a  female  from  Colombia;  Hill  (1964),  forearm  and  cranial  measurements 
of  a  female  from  Guyana;  Starrett  and  de  la  Torre  (1964),  forearm  measurements  of  a  male 
and  female  from  El  Salvador  and  a  female  from  Costa  Rica,  other  external  and  cranial 
measurements  of  a  male  and  female  from  Costa  Rica;  Valdivieso  (1964),  external  and 
cranial  measurements  (mean,  range)  of  specimens  from  Colombia;  Brosset  (1965),  external 
and  cranial  measurements  of  five  males  (including  the  lectotype  of  A.  fallax)  and  five  females 
from  Surinam;  Pirlot  (19656),  external  measurements  of  eight  males  and  eight  females  from 
Est  du  Venezuela;  Tamsitt  and  Valdivieso  (1965a),  forearm  measurements  (mean,  range) 
of  monthly  samples  of  males  from  Colombia;  Tamsitt  and  Valdivieso  (19656),  external 
measurements  (mean,  sd,  se,  range)  of  80  adult  and  18  young  adult  females  from  Colombia; 
Tamsitt  and  Valdivieso  (19666),  external  measurements  (mean,  range)  of  14  specimens 
(four  males,  10  females)  and  cranial  measurements  of  five  females  from  Colombia;  Villa-R. 
(1967),  external  measurements  of  46  specimens  and  cranial  measurements  of  34  specimens 
from  Mexico;  Koopman  (1968),  forearm  measurements  (range)  of  seven  specimens  from 
St.  Vincent;  Goodwin  (1969),  forearm  and  cranial  measurements  of  four  males  and  four 
females  from  Oaxaca;  Burt  and  Stirton  (1969),  forearm  and  cranial  measurements  (range) 
of  five  specimens  from  El  Salvador;  Villa-R.  and  Villa  Cornejo  (1969),  external  and 
cranial  measurements  (mean,  range)  of  seven  specimens  from  Argentina;  Tuttle  (1970), 
forearm  measurements  (range)  of  specimens  from  east  of  the  Andes  in  Peru;  Jones  et  al. 
(1972),  forearm  and  cranial  measurements  (mean,  range)  of  10  specimens  (five  males,  five 
females)  from  Sinaloa;  Pirlot  (1972),  external  measurements  of  specimens  from  Brazil. 

Age  variation. — Lukens  and  Davis  (1957)  presented  forearm  and  cranial  measurements 
of  a  juvenile  female  and  a  subadult  female  from  Guerrero.  Anderson  (1960)  gave  external 
and  cranial  measurements  of  an  immature  female  from  Sinaloa. 

Secondary  sexual  variation. — Tamsitt  and  Valdivieso  (1963a)  found  that  females  from 
Colombia  averaged  larger  than  males  in  all  body  measurements  and  in  four  of  nine  cranial 
measurements.  Anderson  (1960)  found  no  significant  differences  in  size  between  males 
and  females  from  Sinaloa. 

Geographic  variation. — San  Luis  Potosi  material  was  found  to  be  comparable  in  cranial 
size  to  topotypes  of  A.  1.  palmarum  from  Trinidad  (Dalquest,  1950).  Specimens  from  Girardot, 
Mariquita,  and  Puente  Nacional  in  the  Magdalena  River  Valley,  Colombia,  averaged  slightly 
larger  in  body  size  than  did  those  from  two  other  localities:  Mesitas  del  Colegio,  at  a  higher 
elevation  on  the  western  slope  of  the  East  Andes,  and  Villavicencio,  at  the  base  of  the 
eastern  slope  of  the  East  Andes  (Tamsitt  and  Valdivieso,  1963  a). 

Artibeus  phaeotis (Miller,  1902) 

Measurements  of  Artibeus  phaeotis  have  been  recorded  as  follows:  Miller  (1902a),  external 
and  cranial  measurements  of  the  female  holotype  from  Yucatan;  Elliot  (1904),  external  and 
cranial  measurements  of  a  single  specimen;  Andersen  (19066),  cranial  measurements  of  the 
female  holotype  of  A.  turpis  (=  A.  phaeotis)  from  Tabasco  and  the  female  holotype  of  A.  p. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


51 


nanus  from  Guerrero;  Andersen  (1908),  external  and  cranial  measurements  of  the  female 
holotype  of  A.  phaeotis  from  Yucatan,  the  holotype  of  A.  jucundus  (  —  A.  phaeotis)  from 
Veracruz,  the  female  holotype  of  A.  turpis  (  =  A.  phaeotis)  from  Tabasco,  and  mean  and  range 
of  these  measurements  in  eight  specimens  from  Guerrero,  Sinaloa,  and  Colima;  Goodwin 
(1934),  external  measurements  of  a  specimen  from  Guatemala;  Goodwin  (1942 a),  forearm 
and  cranial  measurements  of  one  specimen;  Dalquest  (1953/?),  forearm  and  cranial  measure¬ 
ments  of  a  male  and  female  from  Veracruz;  Jones  and  Lawlor  (1965),  external  and  cranial 
measurements  of  a  male  and  two  females  from  Cozumel  Island,  Quintana  Roo;  Jones 
(1966),  forearm  and  cranial  measurements  (mean,  range)  of  five  specimens  (three  males,  two 
females)  from  El  Peten,  Guatemala,  and  for  a  male  and  female  from  Santa  Rosa,  Guatemala; 
Villa-R.  (1967),  external  measurements  of  28  specimens  and  cranial  measurements  of  22  of 
A.  turpis  turpis,  which  more  or  less  include  A.  p.  phaeotis  and  A.  p.  palatinus  of  Davis 
(1970a),  external  measurements  of  38  specimens  and  cranial  measurements  of  35  specimens 
of  A.  p.  nanus  and  two  males  and  three  females  of  A.  cinerus  phaeotis  from  Veracruz, 
Oaxaca,  and  Tabasco;  Rick  (1968),  external  measurements  of  three  females  and  one  male, 
and  cranial  measurements  of  three  females,  one  male,  and  an  unsexed  specimen  from 
Guatemala;  Goodwin  (1969),  forearm  and  cranial  measurements  of  four  males  and  nine 
females  from  Oaxaca;  Davis  (1970a),  cranial  measurements  (mean,  range)  of  135  specimens 
from  the  Pacific  versant  of  Sinaloa  to  Guerrero,  19  from  Oaxaca  to  Chiapas,  37  from 
Guatemala,  El  Salvador,  and  Nicaragua,  34  from  the  Pacific  versant  of  Costa  Rica  and  seven 
from  the  Caribbean  versant,  124  from  the  Caribbean  versant  of  Guatemala  and  British 
Honduras,  67  from  Honduras  and  Nicaragua,  and  cranial  measurements  of  the  female 
holotype  of  A.  phaeotis  phaeotis  from  Yucatan,  the  female  holotype  of  A.  p.  nanus  from 
Guerrero,  and  the  male  holotype  of  A.  p.  palatinus  from  Guatemala;  Jones  et  al.  (1972), 
forearm  and  cranial  measurements  (mean,  range)  of  five  males  and  five  females  combined 
from  Sinaloa. 

Age  variation. — Juveniles  (cartilaginous  epiphyses  and  unworn  dental  cusps)  could  not 
be  distinguished  from  adults  on  the  basis  of  seven  cranial  measurements  (Davis,  1970a). 

Secondary  sexual  variation. — Davis  (1970a)  found  no  significant  secondary  sexual 
dimorphism  in  four  external  and  seven  cranial  measurements. 

Geographic  variation. — Davis  (1970a)  noted  the  following  size  variation  throughout  the 
geographic  range  of  this  species.  Members  of  the  population  in  western  Mexico  (Sinaloa  to 
Guerrero)  were  generally  the  smallest  for  the  species.  The  rostrum  in  this  population  was 
short,  which  was  reflected  in  the  shortness  of  the  palate.  In  the  Pacific  lowlands  (Oaxaca 
to  Costa  Rica),  specimens  had  a  longer  palate,  skull,  and  forearm;  they  were,  however, 
smaller  than  those  from  the  Caribbean-Gulf  versant.  The  population  occupying  the  Caribbean- 
Gulf  versant  (Veracruz  to  South  America)  was  the  largest  in  the  species. 

Artibeus  toltecus(Saussure,  1860) 

Measurements  of  Artibeus  toltecus  have  been  recorded  as  follows:  Saussure  (18606), 
external  measurements  of  a  single  specimen;  Miller  (1902a),  external  and  cranial  measure¬ 
ments  of  the  male  holotype  of  A.  t.  ravus  from  Ecuador  and  a  specimen  from  Morelos; 
Andersen  (1908),  external  and  cranial  measurements  (range)  of  three  specimens  from  Costa 
Rica,  Nicaragua,  and  Guatemala,  two  (one  cranial)  from  Oaxaca,  nine  (five  cranial)  from 
Jalisco  and  Durango,  and  three  from  Veracruz,  external  measurements  (mean,  range)  of 
18  specimens  (cranial  of  13)  from  Costa  Rica,  Nicaragua,  Guatemala,  Jalisco,  Durango, 
Oaxaca,  and  Veracruz,  and  11  specimens  (mean,  range)  from  Ecuador;  Goodwin  (1934), 
external  measurements  of  a  specimen  from  Guatemala;  Goodwin  (1942a),  external  and 
cranial  measurements  of  two  males  from  Honduras;  Goodwin  (1946),  external  and  cranial 
measurements  (range)  for  the  species;  Dalquest  (1953a),  external  measurements  (mean)  of 
two  males  and  cranial  measurements  (mean)  of  two  males  and  five  females  from  San  Luis 
Potosi;  de  la  Torre  (1954),  external  and  cranial  measurements  (mean,  range)  of  six  specimens 


52 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


from  Tamaulipas;  de  la  Torre  (1955),  forearm  measurements  (mean,  range)  of  five  males 
and  three  females  combined  from  Jalisco;  Felten  (1956 d),  external  measurements  of  a 
specimen  from  El  Salvador;  Jones  et  al.  (1962),  forearm  and  total  length  of  skull  (range)  of 
12  specimens  from  Mexico  (Oaxaca  6,  Tamaulipas  3,  Jalisco  2,  Sinaloa  1);  Alvarez  (1963), 
external  and  cranial  measurements  of  a  male  and  two  females  from  Tamaulipas;  Jones  and 
Alvarez  (1964),  forearm  measurements  of  a  female  and  cranial  measurements  of  this  female 
and  a  specimen  of  unknown  sex  from  San  Luis  Potosf;  Jones  (1964),  forearm  and  cranial 
measurements  of  a  specimen  from  Sinaloa;  Jones  (1966),  forearm  and  cranial  measurements 
(mean,  range)  of  six  specimens  (five  males,  one  female)  from  Guatemala;  Villa-R.  (1967), 
external  measurements  of  20  specimens  and  cranial  measurements  of  18  from  Mexico; 
Genoways  and  Jones  (1968),  forearm  measurements  of  two  males  and  four  females  from 
Zacatecas;  Davis  (1969),  forearm  and  cranial  measurements  (mean,  range)  of  samples  from 
the  Pacific  versant  including  14  from  Sinaloa  and  Nayarit,  12  from  Guerrero,  18  from 
Chiapas,  18  from  Guatemala,  and  17  from  the  Honduran  highlands,  from  the  Atlantic 
versant  including  nine  from  Tamaulipas  and  San  Luis  Potosi,  eight  from  Veracruz,  16 
from  Chiapas,  14  from  Guatemala,  and  29  from  the  Costa  Rican  highlands,  external  and 
cranial  measurements  of  the  male  holotype  of  A.  t.  Hesperus  from  Guerrero  and  the  male 
neotype  of  A.  t.  toltecus  from  Veracruz;  Goodwin  (1969),  forearm  and  cranial  measurements 
of  four  males  and  four  females  from  Oaxaca;  Jones  et  al.  (19716),  forearm  and  cranial 
measurements  (mean,  range)  of  six  specimens  (three  males,  three  females)  from  Departa- 
mento  de  Matagalpa,  Nicaragua,  and  external  and  cranial  measurements  of  10  specimens 
(four  males,  six  females)  from  Isla  de  Ometepe,  Rivas,  Nicaragua;  Alvarez  and  Ramirez- 
Pulido  (1972),  external  and  cranial  measurements  of  two  males  and  two  females  from 
Morelos;  Jones  et  al.  (1972),  forearm  and  cranial  measurements  (mean,  range)  of  10  specimens 
(five  males,  five  females)  from  Sinaloa. 

Geographic  variation. — According  to  Jones  (1966),  specimens  from  Guatemala  averaged 
larger  than  specimens  from  western  Mexico.  Davis  (1969)  showed  that  specimens  from  the 
Pacific  versant  (El  Salvador  to  Sinaloa)  averaged  smaller  for  almost  all  measurements 
compared  to  those  occupying  the  remainder  of  the  species  geographic  range.  Jones  et  al. 
(19716)  reported  two  size  groups  (subspecies)  occurring  in  Nicaragua.  Those  of  smaller  size 
from  Isla  de  Ometepe,  Rivas,  and  the  others  from  Departamento  de  Matagalpa. 

Artibeus  watson i  Thomas,  1901 

Measurements  of  Artibeus  watsoni  have  been  recorded  as  follows:  Thomas  (1901  a), 
forearm  and  cranial  measurements  of  the  male  holotype  and  external  measurements  of 
another  male  from  Panama;  Elliot  (1904),  external  and  cranial  measurements  of  the  holotype 
(after  Thomas,  1901);  Elliot  (1906),  external  and  cranial  measurements  of  the  holotype  of 
Dermanura  jucundum  from  Veracruz;  Andersen  (1908),  external  and  cranial 
measurements  (mean,  range)  of  nine  specimens  from  Panama  and  Nicaragua;  Sanborn 
(1936),  external  measurements  of  two  males  and  cranial  measurements  of  one  male  from 
Guatemala;  Goodwin  (1942u),  external  and  cranial  measurements  of  a  single  specimen; 
Goodwin  (19426),  external  and  cranial  measurements  of  the  male  holotype  from  Panama 
and  the  range  for  these  measurements  in  the  species;  Jones  (1966),  forearm  and  cranial 
measurements  of  a  male  and  female  from  Guatemala;  Davis  (1970a),  cranial  measurements 
of  the  holotype,  external  and  cranial  measurements  (mean,  range)  of  62  males  and  46  females 
from  the  Pacific  versant  of  Costa  Rica,  and  from  the  Atlantic  versant  25  males  and  19  fe¬ 
males  from  Costa  Rica,  22  males  and  17  females  from  Nicaragua,  11  males  and  four  fe¬ 
males  from  Honduras,  and  eight  males  and  four  females  from  Guatemala,  and  cranial  mea¬ 
surements  (mean,  range)  of  120  specimens  from  southwestern  Costa  Rica  (near  type 
locality). 

Geographic  variation. — Davis  (1970a)  considered  Artibeus  watsoni  to  be  monotypic. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


53 


Centurio  senex  Gray,  1 842 

Measurements  of  Centurio  senex  have  been  recorded  as  follows:  Lichtenstein  and  Peters 
(1855),  external  measurements  of  the  holotype  of  Centurio  flavogularis’,  Saussure  (1860a), 
external  measurements  of  the  female  holotype  of  Centurio  mexicanus  from  Mexico;  H. 
Allen  (1861),  external  measurements  of  the  holotype  of  Centurio  mcmurtrii  from  Veracruz; 
Dobson  (1878a),  external  measurements  of  the  female  holotype;  Ward  (1891),  external 
measurements  of  the  female  holotype  of  Centurio  minor  from  Veracruz  and  measure¬ 
ments  given  by  Dobson  ( 1 878c/);  Rehn  (1901),  external  measurements  from  the  litera¬ 
ture  including  Dobson’s  for  C.  senex ,  Lichtenstein's  and  Peters’  for  C.  flavogularis, 
Saussure’s  for  C.  mexicanus  and  Ward’s  for  C.  minor,  external  measurements  of  five  and 
cranial  of  two  specimens  from  Veracruz  and  external  and  cranial  measurements  of  one 
specimen  from  Costa  Rica;  Elliot  (1904),  external  and  cranial  measurements  of  a  specimen; 
Sanborn  (1936),  external  measurements  (range)  of  12  specimens  and  forearm  and  cranial 
measurements  (range)  of  24  specimens  from  Guatemala;  Goodwin  (1942a),  external  and 
cranial  measurements  (range)  in  the  species;  Goodwin  (1946),  forearm  and  cranial  measure¬ 
ments  (range)  of  24  specimens  from  Guatemala  (as  given  by  Sanborn,  1936)  and  the  holotype; 
Felten  (1956c),  external  and  cranial  measurements  of  a  female  from  El  Salvador;  Felten 
( 1 956z/),  external  measurements  of  a  specimen  from  El  Salvador;  Hall  and  Kelson  (1959), 
forearm  and  cranial  measurements  (range)  of  specimens  from  Guatemala;  Burt  and  Stirton 
(1961),  forearm  and  cranial  measurements  of  a  male  from  El  Salvador;  Goodwin  and 
Greenhall  (1961),  forearm  measurements  of  four  males  and  one  female  and  cranial  measure¬ 
ments  of  three  males  and  one  female  from  Trinidad;  Alvarez  (1963),  external  and  cranial 
measurements  of  a  female  from  Tamaulipas;  Villa-R.  (1967),  external  and  cranial  measure¬ 
ments  (mean,  sd,  range)  of  10  specimens  from  Mexico;  Paradiso  (1967),  forearm  and  cranial 
measurements  of  the  female  holotype  of  C.  s.  greenhalli  from  Trinidad,  forearm  measurements 
(mean,  range)  of  28  topotypes,  cranial  measurements  of  1 1  topotypes,  and  forearm  and 
cranial  measurements  (mean,  range)  of  20  specimens  of  C.  5.  senex  from  Panama,  1 1  from 
Guatemala,  and  two  from  Oaxaca;  Goodwin  (1969),  forearm  and  cranial  measurements  of 
a  male  and  female  from  Oaxaca;  Jones  et  al.  (19716),  forearm  and  cranial  measurements 
(mean,  range)  of  11  specimens  (seven  males,  four  females)  from  Nicaragua;  Jones  et  al. 
(1972),  external  and  cranial  measurements  of  two  males  and  one  female  from  Sinaloa; 
Watkins  et  al.  (1972),  forearm  and  cranial  measurements  of  a  male  and  five  females  (mean, 
range)  from  Jalisco,  and  seven  males  and  four  females  from  Nicaragua. 

Secondary  sexual  variation. — Females  from  Nicaragua  averaged  slightly  larger  than  males 
in  both  external  and  cranial  measurements  (Jones  et  al.,  1971  6). 

Geographic  variation. — Specimens  from  Trinidad  were  clearly  larger  than  those  from 
Panama,  Guatemala,  and  Oaxaca  in  most  measurements.  No  overlap  in  forearm  measure¬ 
ments  were  found  (Paradiso,  1967).  Jones  et  al.  (19716)  reported  that  measurements  of  their 
specimens  from  Nicaragua  agreed  in  general  with  those  given  by  Paradiso  (1967)  for 
material  from  Panama.  Specimens  from  Jalisco  compare  favorably  in  size  with  those  from 
the  vicinity  of  the  type  locality  (restricted  by  Goodwin,  1946)  and  elsewhere  in  Nicaragua 
(Watkins  et  al.,  1972). 


Chiroderma  doriae  Thomas,  1891 

Measurements  of  Chiroderma  doriae  have  been  recorded  as  follows:  Thomas  (18916), 
forearm  and  cranial  measurements  for  the  species  (material  described  by  Dobson,  1878a, 
as  C.  villosum  is  actually  C.  doriae  and  formed  the  basis  for  Thomas’  description);  Goodwin 
(1958),  forearm  and  cranial  measurements  of  the  holotype  from  Brazil;  Baker  and  Genoways 
(1976),  external  and  cranial  measurements  (mean,  range)  of  15  males  and  21  females  from 
Brazil. 


54 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Chiroderma  iinprovisum  Baker  and  Genoways,  1976 

Baker  and  Genoways  (1976)  recorded  external  and  cranial  measurements  of  the  male 
holotype  from  Guadeloupe,  Lesser  Antilles. 

Chiroderma  salvini  Dobson,  1878 

Measurements  of  Chiroderma  salvini  have  been  recorded  as  follows:  Elliot  (1904),  external 
and  cranial  measurements  of  one  specimen;  Sanborn  (1941),  forearm  measurements  (range) 
of  22  specimens  and  cranial  measurements  of  three  from  Honduras;  Goodwin  (1942«), 
external  and  cranial  measurements  of  two  males  from  Honduras;  Goodwin  (1946),  external 
and  cranial  measurements  of  two  males  from  Honduras  and  one  from  Costa  Rica;  Goodwin 
(1958),  forearm  and  cranial  measurements  of  a  female  from  Costa  Rica;  Hall  and  Kelson 
(1959),  external  and  cranial  measurements  of  a  male  from  Costa  Rica;  Brosset  (1965), 
external  and  cranial  measurements  of  a  female  from  Ecuador;  Handley  (1965),  external 
and  cranial  measurements  of  two  males  and  1 1  females  (mean,  range)  of  C.  s.  scopaeum  from 
Chihuahua,  Sinaloa,  Nayarit,  Jalisco,  Colima,  and  Guerrero;  Carter  el  al.  (1966),  external 
and  cranial  measurements  of  a  female  from  Guerrero  and  one  from  Honduras;  Villa-R. 
(1967),  external  and  cranial  measurements  of  a  male  from  Costa  Rica;  Genoways  and 
Jones  (1968),  forearm  measurements  of  five  males  from  Zacatecas;  Alvarez  and  Ramirez- 
Pulido  (1972),  external  and  cranial  measurements  of  one  female  from  Puebla;  Anderson 
(1972),  external  and  cranial  measurements  of  two  females  from  Chihuahua;  Baker  (1974), 
forearm  measurements  of  three  specimens  from  Ecuador. 

Geographic  variation.- — Handley  (1965)  distinguished  specimens  from  western  Mexico 
from  typical  members  of  the  species  in  Costa  Rica  and  Panama  by  their  smaller  size  and 
paler  coloration. 


Chiroderma  trinitatum  Goodwin,  1958 

Measurements  of  Chiroderma  trinitatum  have  been  recorded  as  follows:  Goodwin 
(1958),  external  and  cranial  measurements  of  the  female  holotype  from  Trinidad;  Handley 
(1960),  external  and  cranial  measurements  of  the  male  holotype  of  C.  gorgasi  ( =  C.  trinitatum ) 
from  Panama,  a  female  paratype,  and  the  female  holotype  of  C.  trinitatum  from  Trinidad; 
Goodwin  and  Greenhall  (1961),  forearm  and  cranial  measurements  of  the  female  holotype 
from  Trinidad;  Ojasti  and  Linares  (1971),  external  and  cranial  measurements  of  two  females 
from  Venezuela;  Pirlot  (1972),  forearm  measurements  of  a  single  specimen  from  Brazil; 
Gardner  (1976),  external  and  cranial  measurements  (mean,  range)  of  two  males  and  six 
females  from  Peru. 


Chiroderma  villosum  Peters,  1860 

Measurements  of  Chiroderma  villosum  have  been  recorded  as  follows:  Thomas  (18916), 
forearm  and  cranial  measurements  for  the  species;  J.  A.  Allen  (1900),  external  and  cranial 
measurements  of  the  male  holotype  of  C.  villosum  jesupi  from  Colombia;  Miller  (1912), 
external  and  cranial  measurements  of  the  female  holotype  of  C.  isthmicum  (=C.  villosum 
jesupi)  from  Panama;  Elliot  (1917),  external  and  cranial  measurements  of  the  holotype  of 
C.  isthmicunr,  Sanborn  (1936),  forearm  and  cranial  measurements  of  a  male  from  Veracruz; 
Goodwin  (1946),  external  and  cranial  measurements  of  the  female  holotype  of  C.  isthmicum ; 
Goodwin  (1953),  forearm  and  cranial  measurements  of  the  male  holotype  of  C.  villosum 
jesupi  from  Colombia;  Goodwin  (1958),  forearm  and  cranial  measurements  of  the  holotype 
of  C.  v.  jesupi  from  Colombia,  male  holotype  and  female  topotype  of  C.  isthmicum  from 
Panama,  and  a  male  from  Trinidad;  Hall  and  Kelson  (1959),  external  and  cranial  measure¬ 
ments  of  the  holotype  of  C.  isthmicum;  Goodwin  and  Greenhall  (1961),  forearm  and  cranial 
measurements  of  one  male  and  three  females  from  Trinidad;  Husson  (1962),  external  and 
cranial  measurements  of  a  female  from  Surinam;  Villa-R.  (1962),  cranial  measurements  of 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


55 


three  specimens  from  Chiapas,  two  from  Colima,  and  of  the  holotype  of  C.  isthmicum ; 
Davis  et  al.  (1964),  forearm  measurements  (range)  of  12  females  from  Chiapas;  Hill  (1964), 
forearm  and  cranial  measurements  of  a  female  from  Guyana;  Villa-R.  (1967),  external  and 
cranial  measurements  of  three  females  from  Chiapas;  Goodwin  (1969),  forearm  and  cranial 
measurements  of  a  female  from  Oaxaca;  Gardner  et  al.  (1970),  forearm  and  cranial  measure¬ 
ments  of  two  males  from  Costa  Rica;  Birney  et  al.  (1974),  external  and  cranial  measurements 
of  one  male  from  Quintana  Roo. 

Geographic  variation. — Husson  (1962)  found  the  measurements  of  his  female  from  Surinam 
to  correspond  well  with  those  of  the  four  specimens  reported  by  Goodwin  and  Greenhall 
(1961)  from  Trinidad.  According  to  Birney  et  al.  (1974),  their  male  specimen  corresponded 
closely  in  size  to  a  female  reported  by  Goodwin  (1969)  from  Oaxaca. 

Ectophylla  alba  H.  Allen,  1892 

Measurements  of  Ectophylla  alha  have  been  recorded  as  follows:  H.  Allen  (1892),  external 
measurements  of  the  holotype  from  Nicaragua;  H.  Allen  (1898),  external  measurements  of 
the  holotype  and  of  an  Oldfield  Thomas  specimen;  Goodwin  (1946),  external  measurements 
of  the  holotype  from  Nicaragua;  Casebeer  et  al.  (1963),  external  and  cranial  measurements 
of  three  females  from  Costa  Rica;  Starrett  and  Casebeer  (1968),  forearm  measurements  of  a 
male  and  two  females  and  cranial  measurements  of  one  male  from  Costa  Rica;  Gardner 
et  al.  (1970),  forearm  measurements  (eight  males,  two  females)  and  cranial  measurements 
(mean,  range)  of  seven  males  and  two  females  from  Costa  Rica. 

Enchisthenes  hartii  (Thomas,  1892) 

Measurements  of  Enchisthenes  hartii  have  been  recorded  as  follows:  Thomas  (1892), 
external  and  cranial  measurements  of  the  “slightly  immature"  male  holotype  from  Trinidad; 
Andersen  (1908),  external  and  cranial  measurements  of  the  male  holotype  from  Trinidad; 
Sanborn  (1932),  external  and  cranial  measurements  of  a  female  from  Venezuela;  Goodwin 
(1940,  1942,  1946),  external  and  cranial  measurements  of  a  specimen  from  Honduras;  de  la 
Torre  (1955),  forearm  measurements  (mean,  range)  of  12  specimens  (eight  males,  four  females), 
and  cranial  measurements  of  one  male  and  two  females  from  Jalisco;  Hall  and  Kelson  (1959), 
external  and  cranial  measurements  of  a  male  from  Honduras;  Goodwin  and  Greenhall 
(1961),  forearm  and  cranial  measurements  of  the  holotype  from  Trinidad;  Villa-R.  (1967), 
external  measurements  of  a  male  and  female  from  Jalisco;  Baker  and  Lopez  (1968),  forearm 
and  cranial  measurements  of  a  male  from  Tamaulipas  and  a  male  and  female  from  Trinidad; 
Goodwin  (1969),  forearm  and  cranial  measurements  of  a  female  from  Oaxaca;  LaVal  (1969), 
external  and  cranial  measurements  of  one  female  from  Honduras;  Gardner  et  al.  (1970), 
forearm  and  cranial  measurements  (mean,  range)  of  13  specimens  from  Costa  Rica;  Gardner 
(1976),  external  and  cranial  measurements  of  a  female  from  Peru. 

Geographic  variation. — When  comparing  one  male  from  Tamaulipas  with  a  male  and 
female  from  Trinidad,  Baker  and  Lopez  (1968)  concluded  that  no  outstanding  variation 
was  obvious. 

Mesophy  11a  (=  Ectophylla)  macconnelli Thomas,  1901 

Measurements  of  Mesophylla  macconnelli  have  been  recorded  as  follows:  Thomas 
(19016),  external  measurements  of  the  female  holotype  and  one  male  and  cranial  measure¬ 
ments  of  the  holotype  from  Guyana;  Lima  (1926),  external  measurements  of  a  specimen  from 
Brazil;  Cunha  Vieira  (1942),  external  and  cranial  measurements  of  a  female  from  Brazil; 
Sanborn  (1951),  forearm  and  cranial  measurements  of  one  specimen  from  Peru;  Goodwin 
and  Greenhall  (1962),  external  and  cranial  measurements  of  the  female  holotype  of  M.  m. 
flavescens  from  Trinidad,  forearm  and  cranial  measurements  of  one  male  and  two  females 
(including  the  holotype  of  M.  macconnelli)  from  Guyana,  two  males  and  three  females 
from  Peru,  one  male  from  Brazil,  and  one  male  and  two  females  from  Ecuador;  Starrett 
and  Casebeer  (1968),  forearm  and  cranial  measurements  of  a  female  from  Costa  Rica. 


56 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Phyllops  falcatus  (Gray,  1839) 

Measurements  of  Phyllops  falcatus  have  been  recorded  as  follows:  Gundlach  (1872, 
1877),  external  measurements  of  a  specimen  from  Cuba;  Dobson  ( 1 878«),  external  measure¬ 
ments  of  the  male  holotype  from  Cuba;  Elliot  (1904),  external  and  cranial  measurements  of 
one  specimen  from  Cuba;  G.  M.  Allen  (1942),  external  measurements  for  the  species. 

Phyllops  haitiensis(J.  A.  Allen,  1908) 

Measurements  of  Phyllops  haitiensis  have  been  recorded  as  follows:  J.  A.  Allen  (1908  a), 
external  measurements  of  the  holotype  of  P.  haitiensis  from  the  Dominican  Republic; 
Elliot  (1917),  external  and  cranial  measurements  of  the  holotype;  Sanborn  (1941),  external 
measurements  of  two  females  and  cranial  measurements  of  one  from  Haiti;  Goodwin 
(1953),  forearm  and  cranial  measurements  of  the  holotype  from  the  Dominican  Republic. 

Pygoderma  bilabiatum  (Wagner,  1843) 

Measurements  of  Pygoderma  bilabiatum  have  been  recorded  as  follows:  Peters  (1863), 
external  measurements  of  the  holotype  of  Stenoderma  ( Pygoderma )  microdon  from  Surinam; 
Dobson  (1878a),  external  measurements  of  one  specimen;  Elliot  (1904),  external  and 
cranial  measurements  of  a  single  specimen;  Lima  (1926),  external  measurements  of  a  specimen 
from  Brazil;  Cunha  Vieira  (1942),  external  measurements  of  two  females  and  two  of  unknown 
sex  and  cranial  measurements  of  a  female  from  Brazil;  Goodwin  (1942,  1946),  external 
measurements  of  a  specimen  from  Paraguay;  Husson  (1962),  external  and  cranial  measure¬ 
ments  of  two  Brazilian  specimens  and  several  measurements  of  the  male  holotype  of  P. 
microdon  from  Surinam,  as  given  by  Peters  (1863). 

Sphaeronycteris  toxophyllum  Peters,  1 882 

Measurements  of  Sphaeronycteris  toxophyllum  have  been  recorded  as  follows:  Peters 
(1882),  external  measurements  of  the  holotype  from  tropical  America;  Husson  (1958), 
external  and  cranial  measurements  of  four  males,  five  females,  and  one  of  unknown  sex 
from  Venezuela. 


Stenoderma  rufum  Desmarest,  1820 

Measurements  of  Stenoderma  rufum  have  been  recorded  as  follows:  Peters  (1869),  external 
measurements  of  the  holotype  of  S.  r.  rufum,  Anthony  (1918,  1925),  cranial  measurements 
of  fossil  material  from  Puerto  Rico;  G.  M.  Allen  (1942),  cranial  measurements  of  a  single 
specimen;  Hall  and  Bee  (1960),  external  measurements  of  the  holotype  from  an  unknown 
locality  and  external  and  cranial  measurements  of  a  male  and  female  from  St.  John  Island; 
Tamsitt  and  Valdivieso  (1966c),  external  measurements  of  a  female  and  her  one-day-old 
young  (male)  from  Puerto  Rico;  Choate  and  Birney  (1968),  cranial  measurements  of  10 
specimens  of  sub-Recent  material  from  Puerto  Rico  (type  description  of  S.  r.  anthonyi), 
six  specimens  of  Recent  material  from  Puerto  Rico,  and  two  specimens  from  St.  John; 
Hall  and  Tamsitt  (1968),  external  and  cranial  measurements  of  the  female  holotype  of  S.  r. 
darioi  from  Puerto  Rico,  and  the  mean  and  range  of  these  measurements  in  three  males  and 
four  females;  Jones  et  al.  (1971a),  external  and  cranial  measurements  (mean,  sd,  range)  of 
15  males  and  seven  females  from  Puerto  Rico,  and  one  male  and  female  from  St.  John; 
Genoways  and  Baker  (1972),  external  measurements  (mean,  range)  of  14  males  and  six 
females  and  cranial  measurements  of  15  males  and  seven  females  from  Puerto  Rico  (from 
Jones  et  al.,  1971a). 

Individual  variation. — Forearm  and  cranial  measurements  of  specimens  with  a  greyish 
pelage  and  unfused  or  incompletely  fused  phalangeal  epiphyses  (immature)  were  significantly 
smaller  than  adults  (Jones  et  al.,  197 1  a). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


57 


Secondary  sexual  variation. — According  to  Choate  and  Birney  (1968),  females  were 
larger  than  males  in  material  from  Puerto  Rico  and  St.  John  Island.  Indications  also  exist  that 
this  was  true  in  sub- Recent  material.  Jones  et  al.  (1971  a)  found  females  significantly  larger 
than  males  in  all  external  and  cranial  measurements  tested. 

Geographic  variation. — Hall  and  Bee  (1960)  stated  that  cranial  dimensions  of  Puerto 
Rican  specimens  were  larger  than  those  from  St.  John.  Sub-Recent  material  from  Puerto 
Rico  was  larger  throughout  than  the  Recent  material  from  Puerto  Rico  and  St.  John  (Choate 
and  Birney,  1968). 

Hall  and  Tamsitt  (1968)  assigned  specimens  from  St.  John  Island  and  St.  Thomas  Island 
to  S.  r.  rufum  because  they  closely  resembled  the  holotype.  They  named  a  new  subspecies 
from  Puerto  Rico  on  the  basis  of  external  color,  although  they  found  no  differences  between 
the  two  in  overall  size  or  shape  and  size  of  skull. 

Jones  et  al.  (1971a)  confirmed  that  Stenoderma  rufum  was  a  polytypic  species  with  three 
distinct  subspecies.  Recent  Puerto  Rican  specimens  were  characterized  by  marked  secondary 
sexual  dimorphism  and  by  darker  color  than  the  other  Recent  race  from  the  Virgin  Islands; 
subfossil  material  from  Puerto  Rico  was  distinguished  by  larger  size  and  several  details  of 
dentition. 


Sturnira  aratathomasi  Peterson  and  Tamsitt,  1968 

Measurements  of  Sturnira  aratathomasi  have  been  recorded  as  follows:  Peterson  and 
Tamsitt  (1968),  external  and  cranial  measurements  of  the  male  holotype  from  Colombia  and 
a  male  and  female  from  Ecuador;  Thomas  and  McMurry  (1974),  external  and  cranial 
measurements  of  the  holotype  and  three  males  and  three  females  from  Colombia. 

Sturnira  bidens (Thomas,  1915) 

Measurements  of  Sturnira  bidens  have  been  recorded  as  follows:  Thomas  (1915),  external 
and  cranial  measurements  of  the  immature  male  holotype  from  Ecuador;  Gardner  and 
O’Neill  (1969),  forearm  and  cranial  measurements  (mean,  range)  of  six  specimens  from 
Peru  and  the  holotype  from  Ecuador;  Gardner  and  O’Neill  (1971),  forearm  and  cranial 
measurements  (mean,  range)  of  11  specimens  from  Peru;  Marinkelle  and  Cadena  (1972), 
forearm  measurements  (range)  of  two  males  and  seven  females  and  cranial  measurements 
(range)  of  two  males  and  four  females  from  Colombia. 

Geographic  variation. — Marinkelle  and  Cadena  (1972)  found  that  their  specimens  from 
Colombia  averaged  slightly  larger  in  cranial  measurements  than  those  from  Peru  reported 
by  Gardner  and  O'Neill  (1969). 

Sturnira  erythromos(Tschudi,  1844) 

Measurements  of  Sturnira  erythromos  have  been  recorded  as  follows:  Gardner  et  al. 
(1969),  forearm  and  cranial  measurements  (mean,  range)  of  24  specimens  from  Peru;  Tuttle 
(1970),  forearm  measurement  range  in  species. 

Sturnira  lilium  (E.  Geoffroy  St. -Hilaire,  1810) 

Measurements  of  Sturnira  lilium  have  been  recorded  as  follows:  Dobson  (1878a),  external 
measurements  of  one  male;  Cabrera  (1903),  external  measurements  for  the  species  in  Chile; 
Elliot  (1904),  external  and  cranial  measurements  of  a  specimen;  Goldman  (1917),  external 
and  cranial  measurements  of  the  female  holotype  S.  I.  parvidens  from  Guerrero;  Lima 
(1926),  external  measurements  of  a  male  from  Brazil;  Cunha  Vieira  (1942),  external  measure¬ 
ments  of  five  males  and  three  females  and  cranial  measurements  of  four  males  from  Brazil; 
Goodwin  (1942a),  external  and  cranial  measurements  of  the  holotype  of  S.  I.  parvidens  and 
a  male  and  female  from  Honduras;  Goodwin  (1946),  external  and  cranial  measurements  of 


58 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


one  male  from  Honduras,  also  given  by  Goodwin  (1942a);  Hershkovitz  (1949),  external  and 
cranial  measurements  of  a  male  and  female  from  northern  Colombia;  Dalquest  (1953  a), 
external  measurements  (mean)  of  three  males  and  seven  females  combined,  and  cranial 
measurements  (mean)  of  three  males  and  five  females  combined  from  San  Luis  Potosi; 
de  la  Torre  (1954),  external  and  cranial  measurements  of  two  specimens  from  Tamaulipas; 
Felten  (1956c),  external  and  cranial  measurements  of  a  female  from  El  Salvador;  Felten 
(1956 d),  external  measurements  of  a  specimen  from  El  Salvador;  Hall  and  Kelson  (1959), 
forearm  measurements  (mean)  of  12  topotypes  from  Paraguay;  Goodwin  and  Greenhall 
(1961),  forearm  and  cranial  measurements  of  a  male  and  female  from  Trinidad  and  two  males 
from  Paraguay;  Husson  (1962),  external  and  cranial  measurements  of  one  male  and  four 
females  from  Surinam;  Pirlot  (1963),  external  measurements  of  seven  males  and  seven  females 
from  Venezuela  and  cranial  measurements  of  one  female;  Tamsitt  and  Valdivieso  (1963a), 
external  measurements  of  three  males  and  one  female  and  cranial  measurements  of  one 
female  from  Colombia;  Tamsitt  and  Valdivieso  (19636),  external  measurements  of  two 
males  from  Colombia;  Starrett  and  de  la  Torre  (1964),  external  and  cranial  measurements 
of  a  male  and  two  females  from  El  Salvador  and  one  female  from  Nicaragua;  Valdivieso 
(1964),  external  and  cranial  measurements  of  a  specimen  from  Colombia;  de  la  Torre 
(1966),  external  and  cranial  measurements  of  the  male  holotype  and  the  mean  and  range 
of  four  male  and  five  female  paratypes  combined  of  S.  1.  angeli  from  Dominica,  Lesser  Antil¬ 
les;  de  la  Torre  and  Schwartz  (1966),  external  and  cranial  measurements  of  the  female  holo¬ 
type  of  S.  I.  paulsoni  from  St.  Vincent,  Lesser  Antilles;  Villa-R.  (1967),  external  and  cranial 
measurements  (mean,  sd,  range)  of  nine  specimens  from  Mexico;  Pirlot  (1968),  forearm 
measurement  of  a  female  from  Peru;  Goodwin  (1969),  forearm  and  cranial  measurements 
of  four  males  and  five  females  from  Oaxaca;  Villa-R.  and  Villa  Cornejo  (1969),  external  and 
cranial  measurements  (mean,  range)  of  15  specimens  from  Argentina;  Anderson  (1972), 
external  measurements  of  one  adult  specimen  and  cranial  measurements  of  two  from 
Chihuahua;  Jones  et  al.  (1973),  greatest  length  of  skull  (mean,  range)  of  three  males  and  five 
females  combined  from  the  Yucatan  Peninsula;  Taddei  (19756),  external  measurements 
(mean,  se,  range)  of  20  males  and  20  females  and  cranial  measurements  of  15  males  and 
15  females  from  Brazil;  Jones  and  Phillips  (1976),  forearm  and  cranial  measurements  (mean 
and  range  of  sexes  combined)  from  four  Lesser  Antillean  islands — Dominica,  two  males 
and  12  females;  Martinique,  four  males  and  four  females;  St.  Lucia,  four  males  and  three 
females;  and  St.  Vincent,  three  males. 

Individual  variation. — In  specimens  from  Brazil,  coefficients  of  variation  for  external 
measurements  varied  in  20  males  from  2.85  to  5.86  and  in  20  females  from  2.48  to  7.08; 
CV  values  for  cranial  measurements  in  15  males  ranged  from  1.47  to  3.57  and  in  15 
females  from  1.75  to  3.01  (Taddei,  19756). 

Secondary  sexual  variation. — Although  males  generally  averaged  larger  than  females  in 
specimens  from  Brazil,  no  significant  differences  in  external  measurements  were  found.  How¬ 
ever,  in  15  cranial  measurements,  only  two  (braincase  breadth,  cranial  depth)  did  not  differ 
significantly  (Taddei,  19756). 

Geographic  variation. — Comparing  Mexican  material  with  species  from  Paraguay, 
Goldman  (1917)  concluded  that  the  forearm  was  shorter  in  most  of  the  specimens  available 
from  Mexico  and  that  the  skull  was  narrower.  Goodwin  (1942a)  stated  that  size  in  a  Honduran 
series,  including  both  males  and  females,  was  smaller  than  specimens  from  Mexico. 
Jones  et  al.  (1973)  noted  that  the  greatest  length  of  skull  of  a  specimen  from  La  Tuxpena, 
Campeche,  which  Goldman  reported  (1917)  to  be  abnormally  small,  fell  within  the  range 
of  that  observed  for  three  males  and  five  females  combined  from  the  Yucatan  Peninsula — 
their  specimens  averaged  only  slightly  smaller  than  specimens  from  adjacent  Chiapas  and 
Guatemala.  Jones  and  Phillips  (1976)  stated  that  Antillean  S.  I  ilium  generally  fell  within  the 
size  range  of  populations  of  this  species  from  Middle  and  South  America.  They  did  find 
some  variation  between  insular  samples,  although  no  clinal  geographic  trend  could  be 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


59 


demonstrated.  Bats  from  St.  Vincent  tended  to  be  the  largest  cranially  among  Antillean 
populations,  whereas  specimens  from  Martinique  had  proportionally  broader  zygomatic 
arches  and  longer  maxillary  toothrows.  Forearm  length  in  specimens  from  Dominica 
averaged  slightly  larger  than  did  specimens  from  other  islands.  No  other  differences  in 
external  proportions  were  demonstrated. 

Sturnira  ludovici  Anthony,  1924 

Measurements  of  Sturnira  ludovici  have  been  recorded  as  follows:  Anthony  (1924/?), 
external  and  cranial  measurements  of  the  male  holotype  from  Ecuador;  Shamel  (1927), 
external  and  cranial  measurements  of  the  female  holotype  of  S.  I.  bogotensis  ( =  S.  ludovici) 
from  Colombia;  Goodwin  (1940),  external  and  cranial  measurements  of  the  female  holotype 
of  5.  hondurensis  (=5.  ludovici)  from  Honduras;  Goodwin  (1942a),  external  and  cranial 
measurements  of  two  specimens  from  Honduras;  Goodwin  (1946),  forearm  and  cranial 
measurements  of  the  holotype  of  S.  hondurensis,  and  a  male  from  Costa  Rica;  Hershkovitz 
(1949),  external  and  cranial  measurements  of  the  holotype  of  5.  /.  bogotensis  and  the  range 
of  these  measurements  in  two  males  and  two  females  combined  from  Colombia;  de  la  Torre 
(1952),  external  and  cranial  measurements  of  a  male  and  female  from  Michoacan;  Dalquest 
(1953  a),  external  measurements  (mean)  of  three  males  and  cranial  measurements  of  one  of 
unknown  sex,  from  San  Luis  Potosi;  Goodwin  (1953),  external  and  cranial  measurements 
of  the  holotypes  of  S.  ludovici  and  S.  hondurensis ;  Lukins  and  Davis  (1957),  external  and 
cranial  measurements  of  a  female  from  Guerrero;  Baker  and  Greer  (1962),  external  and 
cranial  measurements  of  one  male  and  two  females  from  Durango;  Tamsitt  and  Valdivieso 
(1963a),  external  and  cranial  measurements  (mean,  range)  of  six  males  and  six  females 
combined  from  Colombia;  Jones  and  Phillips  (1964),  external  and  cranial  measurements 
of  the  female  holotype  of  S.  I.  occidentalis  from  Sinaloa,  mean  and  range  of  these  measure¬ 
ments  for  specimens  from  Durango  and  Jalisco  (S.  /.  occidentalis),  Puebla,  Michoacan, 
Oaxaca,  Honduras,  Colombia  (after  Hershkovitz,  1949),  and  Ecuador  (S.  /.  ludovici)-, 
Starrett  and  de  la  Torre  (1964),  external  and  cranial  measurements  of  a  male  and  female  from 
Costa  Rica;  Valdivieso  (1964),  external  and  cranial  measurements  (mean,  range)  of  specimens 
from  Colombia;  Jones  and  Dunnigan  (1965),  forearm  and  cranial  measurements  of  12 
males  and  15  females  (mean,  range)  from  Oaxaca;  Villa-R.  (1967),  external  and  cranial 
measurements  of  five  specimens  from  Mexico;  Goodwin  (1969),  forearm  and  cranial 
measurements  of  eight  males  and  one  female  from  Oaxaca;  Jones  et  al.  (19716),  external 
and  cranial  measurements  of  one  male  from  Nicaragua;  Jones  et  al.  (1972),  forearm  and 
cranial  measurements  of  the  female  holotype  of  S.  1.  occidentalis  and  three  males  from 
Sinaloa. 

Secondary  sexual  variation. — Jones  and  Dunnigan  (1965),  examining  the  mean  and 
extremes  of  forearm  and  six  cranial  measurements,  suggested  that  males  average  slightly 
larger  than  females. 

Geographic  variation. — Lukins  and  Davis  (1957)  concluded  that  their  female  specimens 
from  Guerrero  were  somewhat  smaller  than  those  recorded  by  Hershkovitz  (1949)  from 
Colombia  and  Dalquest  (1953a)  from  San  Luis  Potosi  but  corresponded  closely  to  one 
regarded  as  S.  hondurensis  from  Costa  Rica  (Goodwin,  1946).  Jones  and  Phillips  (1964) 
found  specimens  in  the  northern  part  of  the  range  of  the  species  to  be  smaller  than  speci¬ 
mens  from  Central  America  and  northern  South  America  and  described  them  as  S.  I. 
occidentalis. 


Sturnira  niagna  de  la  Torre,  1966 

Measurements  of  Sturnira  magna  have  been  recorded  as  follows:  de  la  Torre  (1966), 
external  and  cranial  measurements  of  the  male  holotype  and  mean  and  range  of  five  male 
and  three  female  paratypes  from  Peru;  Peterson  and  Tamsitt  (1968),  external  and  cranial 


60 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


measurements  of  the  male  holotype,  mean  and  range  of  five  males  and  three  females  (after 
de  la  Torre,  1966),  and  two  females  from  Peru;  Marinkelle  and  Cadena  (1972),  external 
measurements  of  one  specimen  from  Colombia;  Baker  (1974),  forearm  measurement 
of  a  female  from  Ecuador;  Gardner  (1976),  external  and  cranial  measurements  (mean,  range) 
of  one  male  and  three  females  from  Peru. 

Sturnira  mordax  (Goodwin,  1938) 

Measurements  of  Sturnira  mordax  have  been  recorded  as  follows:  Goodwin  (1938,  1946), 
external  and  cranial  measurements  of  the  male  holotype  from  Costa  Rica;  Hall  and  Kelson 
(1959),  external  and  cranial  measurements  of  the  holotype;  Davis  et  al.  (1964),  external 
and  cranial  measurements  of  six  males  and  two  females  from  Costa  Rica;  Gardner  et  al. 
(1970),  forearm  and  cranial  measurements  (mean,  range)  of  12  specimens  from  Costa  Rica. 

Sturnira  nana  Gardner  and  O’Neill,  1971 

Gardner  and  O’Neill  (1971)  recorded  external  and  cranial  measurements  of  the  female 
holotype  and  forearm  and  cranial  measurements  (mean,  range)  of  five  other  specimens 
from  Peru. 


Sturnira  tildae  de  la  Torre,  1959 

Measurements  of  Sturnira  tildae  have  been  recorded  as  follows:  de  la  Torre  (1959), 
external  and  cranial  measurements  of  the  male  holotype  and  a  female  paratype  from 
Trinidad;  Goodwin  and  Greenhall  (1961),  forearm  and  cranial  measurements  of  two  males 
and  two  females  from  Trinidad;  Hill  (1964),  external  and  cranial  measurements  of  two 
females  from  Guyana;  Marinkelle  and  Cadena  (1971),  external  measurements  of  60  males 
and  60  females  from  Colombia  (mean,  range),  male  holotype  and  female  paratype  from 
Trinidad  (after  de  la  Torre,  1959),  two  females  from  Guyana  (after  Hill,  1964),  and  cranial 
measurements  of  50  males  and  50  females  from  Colombia  (mean,  range),  one  male  and 
five  females  from  Guyana,  holotype,  paratype,  and  three  females  from  Trinidad. 

Geographic  variation. — Marinkelle  and  Cadena  (1971)  found  external  measurements 
of  Colombian  specimens  generally  averaged  larger  than  the  holotype  and  paratype  from 
Trinidad. 


Sturnira  thomaside  la  Torre  and  Schwartz,  1966 

Measurements  of  Sturnira  thomasi  have  been  recorded  as  follows:  de  la  Torre  and 
Schwartz  (1966),  external  and  cranial  measurements  of  the  male  holotype  from  Guadeloupe, 
Lesser  Antilles;  Genoways  and  Jones  (1975),  external  and  cranial  measurements  of  the 
male  holotype  (after  de  la  Torre  and  Schwartz,  1966)  and  four  females  (including  one  juve¬ 
nile)  from  Guadeloupe;  Jones  and  Genoways  (1975),  external  and  cranial  measurements 
(after  Genoways  and  Jones,  1975);  Jones  and  Phillips  (1976),  external  and  cranial  measure¬ 
ments  of  the  same  individuals  as  given  by  Genoways  and  Jones  (1975). 

Uroderma  bilobatum  Peters,  1866 

Measurements  of  Uroderma  bilobatum  have  been  recorded  as  follows:  Peters  (1866a), 
external  measurements  of  a  single  specimen;  Dobson  (1878a),  external  measurements  of 
one  specimen;  Rehn  (1900),  cranial  measurements  of  a  specimen  from  Brazil;  Lyon  (1902a), 
external  and  cranial  measurements  of  the  female  holotype  of  U.  b.  convexum  from  Panama 
and  a  specimen  from  Brazil;  Elliot  (1904),  external  and  cranial  measurements  of  the  holo¬ 
type  of  U.  b.  convexum  (after  Lyon,  1902a)  from  Panama;  Andersen  (19066),  measure¬ 
ments  (range)  of  two  specimens,  including  the  male  holotype  of  U.  b.  thomasi,  from 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


61 


Bolivia;  Andersen  (1908),  external  and  cranial  measurements  (range)  of  one  specimen  from 
Brazil,  one  from  Amazonas,  two  from  Peru,  one  from  Ecuador,  one  from  Cali,  Colombia, 
three  from  Santa  Marta,  Colombia,  and  Valencia,  Venezuela,  two  from  Colon,  Panama, 
two  from  Chiriqui,  Panama,  nine  (eight  cranial)  from  the  islands  off  Panama,  and  one  from 
Costa  Rica;  Lima  (1926),  external  measurements  of  a  male  from  Brazil;  Cunha  Vieira 
(1942),  external  and  cranial  measurements  of  a  male  from  Peru;  Goodwin  (1946),  external 
and  cranial  measurements  of  two  males  from  Costa  Rica;  Hershkovitz  (1949),  external 
and  cranial  measurements  (range)  of  specimens  from  Colombia;  Sanborn  (1951),  greatest 
length  of  skull  of  one  female  from  Peru;  Felten  ( 1956c),  external  measurements  of  a  male 
and  four  females  and  cranial  measurements  of  one  male  and  two  females  from  El  Salvador; 
Felten  (1956^0,  external  measurements  (mean,  range)  of  specimens  from  El  Salvador;  Hall 
and  Kelson  (1959),  external  and  cranial  measurements  of  two  males  from  Costa  Rica;  Burt  and 
Stirton  (1961),  forearm  and  cranial  measurements  (range)  of  16  males  and  13  females 
from  El  Salvador;  Goodwin  and  Greenhall  (1961),  external  measurements  of  a  subadult 
male  and  four  females  and  cranial  measurements  of  the  subadult  male  and  two  females 
from  Trinidad;  Husson  (1962),  external  and  cranial  measurements  of  four  females  from 
Surinam;  Tamsitt  and  Valdivieso  (1963  a),  external  measurements  (mean,  range)  of  nine 
males  and  five  females  combined  from  Colombia;  Valdivieso  (1964),  external  and  cranial 
measurements  (mean,  range)  of  one  male  and  nine  females  combined  from  Colombia; 
Brosset  (1965),  external  and  cranial  measurements  of  one  female  from  Ecuador;  Villa-R. 
(1967),  external  measurements  (mean,  sd,  range)  of  22  specimens  and  cranial  measure¬ 
ments  of  20  from  Chiapas;  Davis  (1968),  forearm  and  cranial  measurements  of  the  holotype 
(juvenile,  unsexed)  of  U.  b.  bilobatum  from  Brazil,  18  males  and  30  females  from  Bolivia, 
eastern  Brazil,  Cayenne,  Guyana,  and  Venezuela,  external  and  cranial  measurements  of  the 
male  holotype  of  U.  b.  trinitatum,  mean  and  range  of  eight  males,  and  five  females  from 
Trinidad,  a  male  paratype  of  U.  b.  thomasi  from  Bolivia,  21  males  and  14  females  from 
Ecuador,  Peru,  and  western  Bolivia,  the  female  holotype  (young)  of  U.  b.  convexum  from 
Panama,  77  males,  and  124  females  from  western  Venezuela,  Colombia,  Panama  (exclusive 
of  the  Bocas  del  Toro  region),  the  Pacific  versant  of  Middle  America  as  far  as  Oaxaca, 
the  male  holotype  of  U.  b.  molaris  from  Chiapas,  36  males  and  58  females  from  the 
Atlantic  versant  of  Middle  America  from  the  Bocas  del  Toro  region  of  Panama  north¬ 
ward  to  southern  Veracruz;  Goodwin  (1969),  forearm  and  cranial  measurements  of  one 
male  and  two  females  from  Oaxaca  and  one  subadult  male  and  two  females  of  Uroderma  sp. 
from  Oaxaca;  Baker  and  McDaniel  (1972),  forearm  and  cranial  measurements  of  the  female 
holotype  of  U.  b.  davisi  from  El  Salvador,  forearm  and  cranial  measurements  (mean, 
sd)  of  16  males  and  10  females  from  Chiapas  and  El  Salvador  (U.  b.  davisi),  33  males 
and  29  females  from  Nicaragua,  Costa  Rica,  and  Colombia  ( U.  b.  convexum),  and  25 
males  and  26  females  from  Tabasco,  Honduras,  Nicaragua,  and  Costa  Rica  (  U.  b.  molaris). 

Secondary  sexual  variation. — Baker  et  al.  (1972c;)  described  sexual  dimorphism  in  this 
species  with  males  larger  than  females. 

Geographic  variation. — According  to  Davis  (1968),  specimens  from  Trinidad  ( U .  b. 
trinitatum)  were  noticeably  larger  than  those  from  the  adjacent  mainland  ( U.  b.  bilobatum) 
but  were  difficult  to  separate  from  specimens  from  Ecuador,  Peru,  and  western  Bolivia 
( U.  b.  thomasi).  Specimens  from  western  Bolivia  were  larger  than  specimens  from  Colombia 
and  the  Pacific  versant  of  Central  America  ( U.  b.  convexum).  U.  b.  convexum,  again,  was 
smaller  in  most  measurements  than  specimens  from  Bolivia,  eastern  Brazil,  the  Guianas, 
and  Venezuela  ( U.  b.  bilobatum).  Specimens  from  the  Atlantic  versant  of  Middle  America 
(U.  b.  molaris)  from  Bocas  de  Toro,  Panama,  northwest  to  Veracruz,  Mexico,  were  of 
moderate  size  for  the  species.  Uroderma  b.  davisi  from  the  Pacific  versant  of  Middle  Ameri¬ 
ca  (Chiapas,  El  Salvador,  Honduras)  averaged  smaller  both  externally  and  cranially  than 
either  convexum  or  molaris  (Baker  and  McDaniel,  1972). 


62 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Uroderma  magnirostrum  Davis,  1968 

Measurements  of  Uroderma  magnirostrum  have  been  recorded  as  follows:  Davis 
(1968),  external  and  cranial  measurements  of  the  male  holotype  from  Honduras  and 
26  males  and  51  females  (mean,  range)  from  Oaxaca,  Chiapas,  El  Salvador,  Honduras, 
Nicaragua,  Panama,  Colombia,  Peru,  Bolivia,  Venezuela,  and  Brazil;  Jones  et  al.  (1971  6), 
external  and  cranial  measurements  of  one  male  and  two  females  from  Nicaragua. 

Geographic  variation. — Davis  (1968)  found  little  evidence  of  geographic  variation  but 
his  findings  were  based  on  relatively  small  sample  sizes  of  U.  magnirostrum. 

Vampyressa  bidens (Dobson,  1878) 

Measurements  of  Vampyressa  bidens  have  been  recorded  as  follows:  Dobson  (1878 z/), 
external  measurements  of  the  female  holotype  from  Peru;  Sanborn  (1936),  forearm 
measurements  (range)  of  two  males  and  one  female,  wing  measurements  of  one  male  and 
one  female  from  Ecuador,  cranial  measurements  of  a  male  and  female  from  Ecuador  and 
the  range  of  these  measurements  in  three  males  and  one  female  from  Peru;  Cunha  Vieira 
(1942),  external  measurements  of  a  male  and  female  and  cranial  measurements  of  a  male 
from  Brazil;  Hill  (1964),  forearm  and  cranial  measurements  of  four  males  and  one  female 
from  Guyana;  Marinkelle  and  Cadena  (1972),  external  and  cranial  measurements  of  one 
female  from  Colombia;  Davis  (1975),  external  and  cranial  measurements  of  13  males 
and  10  females  (mean,  sd,  range)  from  Peru. 

Individual  variation. — Coefficients  of  variation,  as  given  by  Davis  (1975),  varied  from 
1.28  in  greatest  length  of  skull  in  females  to  3.27  in  postorbital  constriction  of  females. 
The  two  external  measurements,  which  were  tested,  fell  within  this  range. 

Secondary  sexual  variation. — Comparing  two  external  and  eight  cranial  measurements 
of  13  males  with  those  of  10  females  showed  no  significant  differences.  Females  generally 
averaged  larger  than  males  (Davis,  1975). 

Vampyressa  brocki  Peterson,  1968 

Measurements  of  Vampyressa  brocki  have  been  recorded  as  follows:  Peterson  (1968), 
external  and  cranial  measurements  of  the  female  holotype  from  Guyana;  Baker  et  al. 
(19726),  external  and  cranial  measurements  of  three  females  from  Colombia;  Peterson 
(1972),  external  and  cranial  measurements  of  the  holotype  and  a  male  from  Guyana; 
Davis  (1975),  forearm  and  cranial  measurements  (range)  of  published  data. 

Vampyressa  melissa  Thomas,  1926 

Measurements  of  Vampyressa  melissa  have  been  recorded  as  follows:  Thomas  (1926), 
external  and  cranial  measurements  of  the  female  holotype  from  Peru;  Goodwin  (1963), 
forearm  and  cranial  measurements  of  the  female  holotype;  Peterson  (1968),  forearm  and 
cranial  measurements  of  one  specimen;  Gardner  (1976),  external  and  cranial  measurements 
of  four  specimens  (one  male,  three  females)  from  Peru. 

Vampyressa  nymphaea  Thomas,  1909 

Measurements  of  Vampyressa  nymphaea  have  been  recorded  as  follows:  Thomas  (1909), 
forearm  and  cranial  measurements  of  the  male  holotype  from  Colombia;  Hall  and  Kelson 
(1959),  forearm  and  cranial  measurements  of  the  holotype  and  external  measurements  of  a 
specimen  from  Panama;  Goodwin  (1963),  forearm  and  cranial  measurements  of  two  males 
from  Colombia  and  two  females  from  Panama;  Peterson  (1968),  forearm  and  cranial 
measurements  (range)  in  specimens  of  the  species;  Gardner  et  al.  (1970),  forearm  and 
cranial  measurements  (mean,  range)  of  five  specimens  (three  males,  two  females)  from 
Costa  Rica;  Jones  et  at.  (19716),  external  and  cranial  measurements  of  one  female  from 
Nicaragua. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


63 


Vampyressa  pusilla  (Wagner,  1843) 

Measurements  of  Vampyressa  pusilla  have  been  recorded  as  follows:  Peters  ( 1 866«), 
external  measurements  of  a  specimen  from  Brazil;  Dobson  (1878a),  external  measurements 
of  one  specimen  from  Brazil;  Thomas  (1909),  forearm  and  cranial  measurements  of  the  male 
holotype  of  V.  p.  thy  one  from  Colombia;  Miller  (1912),  external  and  cranial  measurements 
of  the  immature  female  holotype  of  V.  minuta  (=  V.  pusilla)  from  Panama;  Elliot  (1917), 
external  and  cranial  measurements  of  the  holotype  of  V.  minuta-,  Cunha  Vieira  (1942), 
external  measurements  of  a  specimen  from  Brazil;  Goodwin  (1946),  external  and  cranial 
measurements  of  the  female  holotype  of  V.  minuta  from  Panama  and  those  of  a  male  from 
Costa  Rica;  Hershkovitz  (1949),  external  and  cranial  measurements  of  one  female  from 
Colombia;  Sanborn  (1953),  forearm  and  cranial  measurements  (range)  of  two  males  and 
one  female  from  Peru;  Hall  and  Kelson  (1959),  cranial  measurements  of  the  holotype  of 
V.  p.  thyone;  Davis  et  al.  (1964),  external  and  cranial  measurements  of  a  female  from  Chiapas; 
Goodwin  (1963),  external  and  cranial  measurements  of  the  male  holotype  of  V.  pusilla  from 
Brazil,  the  male  holotype  of  V.  nattereri  (—  V.  pusilla )  from  Brazil,  and  forearm  and  cranial 
measurements  of  the  female  holotype  of  V.  p.  venilla  from  Peru,  three  females  from  Panama, 
two  males  from  Costa  Rica,  one  male  and  three  females  from  Colombia,  two  males  and 
one  female  from  Ecuador,  five  males  and  five  females  from  Peru,  and  one  female  from 
Venezuela;  Starrett  and  de  la  Torre  (1964),  external  and  cranial  measurements  of  one  female 
from  Nicaragua;  Peterson  (1965a),  external  and  cranial  measurements  of  a  female  from 
British  Honduras;  Tamsitt  and  Valdivieso  (1966a),  forearm  and  cranial  measurements  of  a 
male  and  female  from  Colombia  (the  latter  as  given  by  Hershkovitz,  1949);  Rick  (1968), 
external  and  cranial  measurements  of  one  male  and  female  from  Guatemala;  Gardner 
et  al.  (1970),  forearm  and  cranial  measurements  (mean,  range)  of  five  specimens  (one  male, 
four  females)  from  Costa  Rica;  Jones  et  al.  (19716),  forearm  and  cranial  measurements  of 
two  males  and  mean  and  range  of  six  females  from  Nicaragua;  Baker  et  al.  (1973),  external 
and  cranial  measurements  of  36  specimens  from  Colombia,  Ecuador,  and  Venezuela,  four 
specimens  from  the  Darien  of  Panama,  14  from  the  remainder  of  Panama,  and  seven  from 
Nicaragua;  Jones  et  al.  (1973),  external  and  cranial  measurements  of  one  female  from 
Campeche. 

Individual  variation. — Baker  et  al.  (1973)  found  coefficients  of  variation  for  forearm 
and  cranial  measurements  in  four  samples  from  Central  and  South  America  ranged 
between  1.5  and  7.2.  Lowest  values  were  for  breadth  across  upper  molars  in  the  sample 
from  the  Darien  of  Panama  and  postorbital  breadth  in  the  sample  from  Nicaragua;  the 
highest  CV  value  was  for  postorbital  breadth  in  the  sample  from  the  Darien  of  Panama. 
All  samples  had  coefficients  of  variation  exceeding  4.0  for  palatal  length. 

Geographic  variation. — Goodwin  (1963),  in  his  review  of  the  genus,  recognized  three 
subspecies  of  V.  pusilla.  These  were  based  primarily  on  minor  details  of  coloration  and 
slight  size  differences.  Handley  (19666)  believed  that  the  subspecific  variations  noted  by 
Goodwin  could  be  attributed  to  variation  with  age  and  chose  to  consider  V.  pusilla  as 
being  monotypic.  Two  years  later,  Peterson  (1968)  recognized  two  subspecies — one  from 
southeastern  Brazil  and  the  other  occupying  the  remainder  of  the  geographic  range  of 
the  species  in  South  and  Central  America.  He  did  not  give,  however,  the  characteristics 
used  to  distinguish  them. 

Starrett  and  de  la  Torre  (1964)  concluded  that  their  female  specimen  from  Nicaragua 
was  similar  in  size  to  measurements  given  by  Goodwin  (1946)  for  the  holotype  of  V.  minuta 
(=  V.  pusilla)  from  Panama  and  for  a  specimen  from  Costa  Rica.  They  also  found  their 
specimen  from  Nicaragua  indistinguishable  from  three  specimens  from  Peru. 

Baker  et  al.  (1973)  found  no  significant  differences  in  forearm  and  cranial  measurements 
of  specimens  from  four  geographic  areas  including  Colombia,  Ecuador,  Venezuela,  the 
Darien  and  remainder  of  Panama,  and  Nicaragua. 

Jones  et  al.  (1973)  followed  Handley  (19666)  in  considering  V.  pusilla  monotypic  when 
assigning  their  specimen  from  Campeche. 


64 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Vampyrodes  caraccioli  (Thomas,  1889) 

Measurements  of  Vampyrodes  caraccioli  have  been  recorded  as  follows:  Thomas  (1889), 
external  and  cranial  measurements  of  the  holotype  from  Trinidad;  G.  M.  Allen  (1908), 
external  and  cranial  measurements  of  the  female  holotype  of  V.  major  from  Panama; 
Sanborn  (1936),  forearm  and  cranial  measurements  (range)  of  two  males,  one  female, 
and  one  unsexed  specimen,  and  wing  measurements  of  one  male  from  Guatemala;  Sanborn 
(1941),  external  and  cranial  measurements  of  a  male  from  Trinidad;  Goodwin  (1942«), 
external  and  cranial  measurements  of  the  female  holotype  of  V.  major  from  Panama; 
Goodwin  (1946),  external  and  cranial  measurements  of  the  holotype  of  V.  major  (as  in 
Goodwin,  1942)  and  of  one  specimen  from  Nicaragua;  Husson  (1954),  external  and  cranial 
measurements  of  four  males  from  Tobago;  Hall  and  Kelson  (1959),  cranial  measurements 
of  a  male  from  Guatemala;  Goodwin  and  Greenhall  (1961),  forearm  and  cranial  measure¬ 
ments  of  the  unsexed  holotype  from  Trinidad  and  a  female  from  Tobago;  Villa-R.  (1967), 
external  and  cranial  measurements  of  two  males  and  one  female  from  Veracruz;  Starrett 
and  Casebeer  (1968),  forearm  measurements  of  three  males  and  nine  females,  and  cranial 
measurements  of  three  males  and  two  females  from  Costa  Rica;  Goodwin  (1969),  forearm 
and  cranial  measurements  of  one  male  from  Oaxaca;  Linares  (1969),  external  and  cranial 
measurements  of  one  specimen  from  Venezuela;  Gardner  et  al.  (1970),  forearm  measure¬ 
ment  of  a  female  from  Costa  Rica. 

Geographic  variation. — According  to  Sanborn  (1936),  his  series  of  specimens  from 
Guatemala  agreed  closely  in  measurements  with  the  original  description  of  V.  major 
from  Danama.  Gardner  et  al.  (1970)  noted  that  the  forearm  length  of  their  female  from 
Costa  Rica  greatly  exceeded  the  range  for  three  males  and  nine  females  recorded  by  Starrett 
and  Casebeer  (1968)  from  Costa  Rica. 

Vampyrops  aurarius  Handley  and  Ferris,  1972 

Measurements  of  Vampyrops  aurarius  have  been  recorded  as  follows:  Handley  and  Ferris 
(1972),  external  and  cranial  measurements  of  the  male  holotype  from  Venezuela;  Carter 
and  Rouk  (1973),  forearm  and  cranial  measurements  of  the  male  holotype  from  Venezuela 
and  the  mean  and  range  for  Peruvian  specimens. 

Vampyrops  brachycephalus  Rouk  and  Carter,  1972 

Measurements  of  Vampyrops  brachycephalus  have  been  recorded  as  follows:  Rouk  and 
Carter  (1972),  external  and  cranial  measurements  of  the  male  holotype  from  Huanuco, 
Peru  and  mean  and  range  for  13  specimens  from  Loreto,  Peru,  six  from  Huanuco,  Peru, 
three  from  Colombia,  and  13  from  Venezuela;  Gardner  and  Carter  (19726),  external  and 
cranial  measurements  of  the  male  holotype  and  measurements  (mean,  range)  of  13  specimens 
from  Loreto  and  six  specimens  from  Huanuco,  Peru  (see  also  Rouk  and  Carter,  1972); 
Handley  and  Ferris  (1972),  external  and  cranial  measurements  of  the  male  holotype  of 
V.  latus  1  =  V.  brachycephalus)  from  Peru  and  similar  measurements  for  the  male  holotype  of 
V.  latus  saccharus  from  Venezuela;  Carter  and  Rouk  (1973),  forearm  and  cranial  measure¬ 
ments  of  the  holotype  of  V.  latus  and  V.  latus  saccharus  as  well  as  mean  and  range  of  these 
measurements  for  13  specimens  from  Loreto,  Peru,  and  an  unspecified  number  of  specimens 
from  Tingo  Maria,  Peru. 


Vampyrops  dorsalis  Thomas,  1900 

Measurements  of  Vampyrops  dorsalis  have  been  recorded  as  follows:  Thomas  (1900), 
external  and  cranial  measurements  of  the  holotype  from  Ecuador;  Lyon  (19026),  external 
and  cranial  measurements  of  the  female  holotype  of  V.  umbratus  from  Colombia;  Thomas 
(1914),  external  and  cranial  measurements  of  the  male  holotype  of  V.  oratus  from  Venezuela; 
Sanborn  (1951),  forearm  and  cranial  measurements  of  the  holotype  and  a  male  from  Peru; 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


65 


Sanborn  (1955),  external  measurements  of  two  males  and  cranial  measurements  (range)  of 
10  specimens  (eight  males,  one  female,  one  unsexed)  from  Colombia,  Ecuador,  Peru, 
and  Venezuela;  Tamsitt  and  Valdivieso  (1966a),  forearm  and  cranial  measurements  (range) 
of  four  males  from  Colombia,  and  those  given  by  Sanborn  (1955),  Handley  and  Ferris 
(1972),  external  and  cranial  measurements  of  the  female  holotype  of  V.  aquilus  from 
Panama;  Gardner  and  Carter  (19726),  external  and  cranial  measurements  of  the  immature 
male  holotype  from  Ecuador  and  mean  and  range  for  one  specimen  from  Ecuador  and 
eight  from  Peru;  Carter  and  Rouk  (1973),  forearm  and  cranial  measurements  of  the  holo¬ 
type  of  V.  aquilus  (=  V.  dorsalis)  as  reported  by  Handley  and  Ferris  (1972)  and  mean  and 
range  for  specimens  from  Peru  of  V.  dorsalis  reported  by  Gardner  and  Carter  (19726). 

Vampyrops  helleri  Peters,  1866 

Measurements  of  Vampyrops  helleri  have  been  recorded  as  follows:  Peters  (1866a), 
external  measurements  of  the  holotype  from  Mexico;  Dobson  (1878a),  measurements  of 
one  specimen  from  Mexico;  H.  Allen  (1891),  external  and  cranial  measurements  of  the 
female  holotype  of  Vampyrops  zarhinus  from  Brazil  (holotype  now  considered  to  be  from 
Panama  according  to  Jones  and  Carter,  1976);  Robinson  and  Lyon  (1901),  external  measure¬ 
ments  of  four  females  from  Venezuela;  Elliot  (1904),  external  and  cranial  measurements  of 
one  specimen;  Thomas  (1912a),  external  and  cranial  measurements  of  the  male  holotype 
of  V.  incarum  from  Peru;  Cunha  Vieira  (1942),  external  measurements  of  a  male  and  female 
and  cranial  measurements  of  a  male  of  Vampyrops  zarhinus  (=V.  heller i)  from  Brazil; 
Goodwin  (1942a),  external  and  cranial  measurements  of  a  single  specimen;  Goodwin 
(1946),  forearm  and  cranial  measurements  of  one  female  from  Costa  Rica;  Sanborn 
(19496),  forearm  measurement  of  one  female  and  cranial  measurements  of  two  females 
from  Peru;  Sanborn  (1955),  external  and  cranial  measurements  (range)  of  specimens  from 
Oaxaca,  Honduras,  Costa  Rica,  Panama,  Cayenne,  Trinidad,  Brazil,  Venezuela,  Colombia, 
and  Peru;  Sherman  (1955),  external  measurements  of  a  male  from  Paraguay;  Hall  and 
Kelson  (1959),  forearm  and  cranial  measurements  of  one  female  from  Costa  Rica;  Goodwin 
and  Greenhall  (1961),  external  and  cranial  measurements  of  one  male  and  three  females 
from  Trinidad;  Husson  (1962),  external  and  cranial  measurements  of  eight  males  from 
Surinam;  Tamsitt  and  Valdivieso  (1963a),  external  measurements  of  three  males  and  one 
female  and  cranial  measurements  of  three  males  from  Colombia;  Starrett  and  de  la  Torre 
(1964),  external  and  cranial  measurements  of  a  female  from  Costa  Rica;  Davis  et  al. 
(1964),  external  and  cranial  measurements  (mean,  range)  of  six  specimens  from  Chiapas 
and  Central  America;  Valdivieso  (1964),  external  measurements  of  one  specimen  from 
Colombia;  Villa-R.  (1967),  external  and  cranial  measurements  of  a  male  and  two  females 
from  Oaxaca,  Chiapas,  and  Tabasco;  Rick  (1968),  external  and  cranial  measurements  of 
a  male  and  female  from  Guatemala;  Goodwin  (1969),  forearm  and  cranial  measurements 
of  one  female  from  Oaxaca;  Gardner  and  Carter  (19726),  external  measurements  of  the 
holotype  (sex  unknown)  from  Mexico,  and  external  and  cranial  measurements  (mean, 
range)  of  four  specimens  from  Peru;  Rouk  and  Carter  (1972),  forearm  and  cranial  measure¬ 
ments  (mean,  range)  of  four  specimens  from  Peru,  one  from  Ecuador,  nine  from  Colombia, 
three  from  Venezuela,  one  from  Panama,  two  from  Costa  Rica,  20  from  Nicaragua,  and 
12  from  Honduras. 


Vampyrops  infuscus  Peters,  1880 

Measurements  of  Vampyrops  infuscus  have  been  recorded  as  follows:  Peters  (1880), 
external  measurements  of  the  holotype  from  Peru;  Miller  (1902a),  external  and  cranial 
measurements  of  the  female  holotype  of  V.  fumosus  from  Brazil;  Sanborn  (1936),  forearm 
and  cranial  measurements  (range)  of  three  males  and  one  female  from  Ecuador;  Cunha 
Vieira  (1942),  external  measurements  of  the  holotype  of  V.  fumosus  based  on  Miller 
(1902a);  Sanborn  (1951),  forearm  measurements  of  the  holotype  of  V.  infuscus  from  Brazil 


66 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


and  a  series  of  specimens  from  Peru,  Ecuador,  and  Colombia;  Marinkelle  (1970),  external 
and  cranial  measurements  of  the  female  holotype  of  V.  intermedius  from  Colombia  and  the 
range  of  these  measurements  in  the  paratypes  (five  males,  ten  females);  Gardner  and  Carter 
(19726),  external  and  cranial  measurements  of  the  adult  male  neotype  of  V.  infuscus  and 
the  mean  and  range  of  several  external  and  cranial  measurements  of  six  specimens,  including 
the  neotype  from  Peru. 

Secondary  sexual  variation. — Marinkelle  (1970)  found  no  significant  differences  in  size 
between  five  males  and  10  females  from  Colombia. 

Vampyrops  lineatus(E.  Geoffroy  St.-Hilaire,  1810) 

Measurements  of  Vampyrops  Uneatus  have  been  recorded  as  follows:  Dobson  (1878 z/), 
external  measurements  of  the  holotype;  H.  Allen  (1891),  external  and  cranial  measurements 
of  one  specimen;  Elliot  (1904),  external  measurements  of  a  single  specimen;  Lima  (1926), 
external  measurements  of  a  specimen  from  Brazil;  Cunha  Vieira  (1942),  external  measure¬ 
ments  of  three  males,  three  females,  and  one  unsexed  specimen,  and  cranial  measurements 
of  three  males  and  one  female  from  Brazil;  Goodwin  (1946),  external  and  cranial  measure¬ 
ments  of  a  male  from  Paraguay;  Hershkovitz  (1949),  external  measurements  of  four  males 
and  a  female  and  cranial  measurements  of  one  male  from  Colombia;  Sanborn  (1955), 
external  measurements  of  one  male  and  seven  females  and  cranial  measurements  of  an 
unspecified  number  of  specimens  from  Brazil,  Paraguay,  and  Bolivia. 

Vampyrops  nigellus  Gardner  and  Carter,  1972 

Gardner  and  Carter  ( 1 927 c/,  19726)  gave  external  and  cranial  measurements  of  the  male 
holotype  from  Peru  and  mean  and  range  of  measurements  of  17  specimens  from  Peru. 

Vampyrops  recifinus  Thomas,  1901 

Measurements  of  Vampyrops  recifinus  have  been  recorded  as  follows:  Thomas  (1901c), 
external  and  cranial  measurements  of  the  male  holotype  from  Brazil;  Cunha  Vieira 
(1942),  external  measurements  of  a  male  and  a  female  from  Brazil;  Sanborn  (1955), 
external  and  cranial  measurements  (range)  of  specimens  from  Brazil  and  Guyana. 

Vampyrops  vittatus( Peters,  1859) 

Measurements  of  Vampyrops  vittatus  have  been  recorded  as  follows:  Dobson  (1878u), 
external  measurements  of  one  specimen;  Goodwin  (1946),  external  and  cranial  measure¬ 
ments  of  a  specimen  from  Costa  Rica;  Sanborn  (1955),  forearm  and  cranial  measurements 
(range)  of  specimens  from  Venezuela,  Colombia,  Brazil,  Ecuador,  and  Peru  (he  considered 
V.  vittatus  and  V.  fuscus nonspecific);  Hall  and  Kelson  (1959),  external  and  cranial  measure¬ 
ments  of  a  single  specimen  from  Colombia;  Davis  et  al.  (1964),  external  and  cranial  measure¬ 
ments  of  a  male  and  two  females  from  Costa  Rica;  Gardner  et  al.  (1970),  forearm  and 
cranial  measurements  (mean,  range)  of  six  males  and  nine  females  from  Costa  Rica;  Gardner 
and  Carter  (19726),  external  and  cranial  measurements  of  the  male  holotype  from  Venezuela 
and  several  of  these  measurements  (mean,  range)  for  six  specimens  from  Peru. 

Geographic  variation. — According  to  Gardner  and  Carter  (19726)  measurements 
of  six  specimens  from  Peru  were  much  the  same  as  those  reported  by  Gardner  et  al.  (1970) 
for  19  specimens  from  Costa  Rica. 

Subfamily  Brachyphyllinae 
Brachyphylla  cavernarum  Gray,  1834 

Measurements  of  Brachyphylla  cavernarum  have  been  recorded  as  follows:  Gray  (1834), 
external  measurements  of  the  holotype  from  St.  Vincent;  Dobson  (1878u),  external  measure- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


67 


ments  of  one  specimen;  Miller  (1902//),  cranial  measurements  of  a  male  topotype  from 
St.  Vincent;  Miller  (19026),  external  measurements  of  a  female  specimen;  Elliot  (1904), 
external  and  cranial  measurements  of  one  specimen;  Miller  (1913 c/),  external  and  cranial 
measurements  of  the  female  holotype  of  B.  c.  minor  from  Barbados  and  cranial  measure¬ 
ments  for  an  additional  male;  Elliot  (1917),  external  and  cranial  measurements  of  the  holo¬ 
type  of  B.  c.  minor,  Anthony  (1918,  1925),  external  measurements  (mean,  range)  of  11 
specimens  (2  males,  9  females)  and  cranial  measurements  of  10  specimens  (3  males,  7  females) 
from  Puerto  Rico;  Hall  and  Kelson  (1959),  external  and  cranial  measurements  (range)  of 
10  specimens  and  external  and  cranial  measurements  of  the  holotype  of  B.  6.  minor  from 
Barbados;  Husson  (1960),  forearm  and  cranial  measurements  (range)  of  18  specimens  from 
St.  Martin  and  Saba;  Choate  and  Birney  (1968),  cranial  measurements  of  two  samples  of 
sub-Recent  material  from  Puerto  Rico;  Koopman  (1968),  cranial  measurements  of  a  male 
and  female  from  Barbados  (as  given  by  Miller,  1913 a)  and  the  range  of  a  series  of  males 
from  Anguilla  and  females  from  St.  Martin;  Buden  (1977),  forearm  measurements  (mean, 
range)  of  three  males  and  eight  females,  cranial  measurements  of  four  males  and  eight 
females  from  Puerto  Rico,  forearm  measurements  (mean,  range)  of  seven  males  and  three 
females,  and  cranial  measurements  of  1 1  males  and  four  females  from  St.  John. 

Geographic  variation. — Buden  (1977)  treated  all  members  of  the  genus  as  a  single  species. 
Within  the  species,  he  recognized  several  areas  of  morphological  variation.  Individuals 
from  Puerto  Rico,  Virgin  Islands,  and  most  of  the  Lesser  Antilles  were  the  largest.  Specimens 
from  Barbados  in  the  Lesser  Antilles  were  small  compared  to  populations  on  adjacent 
islands.  Specimens  from  Cuba,  Hispaniola,  and  the  Bahamas  were  also  small,  with  Cuban 
material  being  distinguished  by  deeper  and  more  robust  zygomatic  arches.  However,  Silva- 
Taboada  (1976),  after  examining  this  group,  concluded  that  it  contained  two  species,  each 
with  two  subspecies. 

Initially,  populations  from  Barbados  (minor)  and  the  remainder  of  the  Lesser  Antilles 
(caver narum)  were  considered  two  separate  species.  Koopman  (1968),  however,  showed  that 
there  was  overlap  in  size  among  both  males  and  females  and  concluded  from  this  that  the 
two  were  subspecies  of  B.  cavernarum. 

Brachyphylla  nana  Miller,  1902 

Measurements  of  Brachyphylla  nana  have  been  recorded  as  follows:  Gundlach  (1872, 
1877),  external  measurements  of  a  specimen  from  Cuba;  Miller  (1902c/),  cranial  measure¬ 
ments  of  the  holotype  from  Cuba;  Miller  (19026),  external  measurements  of  one  female  from 
Cuba;  Elliot  (1904),  external  and  cranial  measurements  of  a  single  specimen;  Miller  (1918), 
cranial  measurements  of  the  holotype  and  an  additional  specimen  of  B.  nana  pumila  from 
the  type  locality  on  Haiti;  Miller  (1929),  cranial  measurements  of  one  specimen  from  Haiti; 
Goodwin  (1933),  external  measurements  of  five  males  from  the  Dominican  Republic 
and  one  female  from  Cuba;  Sanborn  (1941),  external  measurements  of  three  females  (range) 
and  cranial  measurements  of  one  female  from  Haiti;  Hall  and  Kelson  (1959),  cranial 
measurements  of  the  holotype  of  Brachyphylla  nana  and  B.  pumila:,  Silva-Taboada  (1974), 
measurements  of  fossil  humeri,  crania,  and  mandibles  from  Cuba;  Buden  (1977),  forearm 
measurements  (mean,  range)  of  eight  males  and  13  females,  cranial  measurements  (mean, 
range)  of  five  males  and  nine  females  from  Cuba,  forearm  measurements  of  seven  males 
and  three  females,  and  cranial  measurements  of  10  males  and  three  females  from  Hispaniola 
and  of  seven  males  and  12  females  from  Middle  Caicos,  Bahamas. 

Geographic  variation. — Buden  (1977),  considering  B.  nana  and  B.  cavernarum 
conspecific,  found  populations  from  Middle  Caicos,  Cuba,  and  Hispaniola  (nana)  to  be 
distinctly  smaller  than  individuals  from  Puerto  Rico,  Virgin  Islands,  and  the  remainder 
of  the  Lesser  Antilles  (cavernarum).  Many  characters  of  specimens  from  Caicos  and 
Hispaniola  overlap  broadly,  but  Buden  distinguished  specimens  from  the  two  areas  by  the 
deeper  and  more  robust  zygomatic  arch  of  specimens  from  Cuba. 


68 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Erophylla  bonibifrons( Miller,  1899) 

Measurements  of  Erophylla  bombifrons  have  been  recorded  as  follows:  Miller  (1899), 
external  and  cranial  measurements  of  the  male  holotype  from  Puerto  Rico;  Elliot  (1904), 
external  and  cranial  measurements  of  the  holotype  from  Puerto  Rico  as  given  by  Miller 
(1899);  Elliot  (19056),  external  and  cranial  measurements  of  the  holotype  of  E.  h.  santacristo- 
balensis  from  the  Dominican  Republic;  Elliot  (1917),  external  and  cranial  measure¬ 
ments  of  the  holotype  of  E.  b.  santacristobalensis ,  Anthony  (1918,  1925),  external  measure¬ 
ments  (mean,  range)  of  six  specimens  and  cranial  measurements  (mean,  range)  of  three 
specimens  from  Puerto  Rico;  Miller  (1929),  cranial  measurements  of  three  specimens  from 
Haiti  and  three  from  Puerto  Rico;  Hall  and  Kelson  (1959),  forearm  and  cranial  measure¬ 
ments  of  the  holotype  of  E.  b.  bombifrons ;  Buden  (1976),  external  and  cranial  measure¬ 
ments  (mean,  sd,  range)  of  49  specimens  (21  cranial)  from  Hispaniola  and  47  (18  cranial) 
from  Puerto  Rico. 

Individual  variation. — Coefficients  of  variation  in  external  measurements  of  specimens 
from  Hispaniola  and  Puerto  Rico  varied  from  1.98  to  4.94  and  in  cranial  measurements 
from  1.84  to  3.45  (Buden,  1976). 

Geographic  variation. — Buden  (1976)  treated  the  two  recognized  species  ( bombifrons 
and  sezekorni)  of  the  genus  as  conspecifics  and  relegated  them  to  subspecific  status.  Differences 
between  many  of  the  currently  recognized  taxa  were  considered  slight.  Skull  shape  was 
considered  the  main  diagnostic  factor  in  distinguishing  bombifrons  and  sezekorni. 

Erophylla  sezekorni  (Gundlach,  1861) 

Measurements  of  Erophylla  sezekorni  have  been  recorded  as  follows:  Gundlach  (1877), 
external  measurements  of  a  specimen  from  Cuba;  Dobson  ( 1 878c/),  external  measurements 
of  a  single  specimen;  Miller  (1899),  external  and  cranial  measurements  of  the  male  holotype 
of  E.  s.  plantifrons  from  the  Bahamas;  Elliot  (1904),  external  and  cranial  measurements  of 
two  specimens;  G.  M.  Allen  (1917),  external  and  cranial  measurements  of  the  male  holotype 
from  Jamaica;  Shamel  (1931),  external  and  cranial  measurements  of  the  male  holotype  of 
E.  s.  mariguanensis  from  Mariguana  Island,  southern  Bahamas,  cranial  measurements 
(range)  of  eight  additional  specimens,  and  eight  from  the  northern  Bahamas;  Buden  (1976), 
external  and  cranial  measurements  (mean,  sd,  range)  of  50  specimens  (19  cranial)  from  New 
Providence,  Bahamas,  35  (six  cranial)  from  Mayaguana,  Bahamas,  88  (44  cranial)  from 
Cuba,  and  66  (29  cranial)  from  Jamaica. 

Individual  variation. — Coefficients  of  variation  in  external  measurements  of  specimens 
from  the  Bahamas,  Cuba,  and  Jamaica  varied  from  2.06  to  4.40  and  in  cranial  measurements 
from  1.58  to  2.93  (Buden,  1976). 

Geographic  variation. — See  geographic  variation  in  E.  bombifrons. 

Phyllonycteris  aphylla  (M  iller,  1 898) 

Measurements  of  Phyllonycteris  aphylla  have  been  recorded  as  follows:  Miller  (1898), 
external  and  cranial  measurements  of  the  male  holotype  from  Jamaica;  Elliot  (1904), 
external  and  cranial  measurements  of  one  specimen;  G.  M.  Allen  (1942),  external  and 
cranial  measurements  for  the  species;  Hall  and  Kelson  (1959),  external  and  cranial  measure¬ 
ments  of  the  holotype;  Henson  and  Novick  (1966),  external  measurements  of  a  female  from 
Jamaica;  Howe  (1974),  external  measurements  of  three  females  from  Jamaica. 

Phyllonycteris  mjyor  Anthony,  1917 

Measurements  of  Phyllonycteris  major  have  been  recorded  as  follows:  Anthony  (1917, 
1918,  1925),  cranial  measurements  of  the  holotype  and  eight  additional  specimens 
(sub-Recent  fossils)  from  Puerto  Rico;  G.  M.  Allen  (1942),  cranial  measurements  for  the 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


69 


species;  Goodwin  (1953),  cranial  measurements  of  the  holotype  from  Puerto  Rico;  Choate 
and  Birney  (1968),  measurements  (mean,  range)  of  partial  crania  and  partial  lower  jaws 
from  Puerto  Rico. 


Phyllonycteris  poeyi  Gundlach,  1861 

Measurements  of  Phyllonycteris  poeyi  have  been  recorded  as  follows:  Gundlach  (1872, 
1877),  external  measurements  of  a  specimen  from  Cuba;  Dobson  (1878c/),  external  measure¬ 
ments  of  one  specimen  from  Cuba;  Elliot  (1904),  external  and  cranial  measurements  of  a 
single  specimen  from  Cuba;  Miller  (1904),  external  measurements  of  a  single  specimen  from 
Cuba;  Miller  (1904),  external  measurements  of  12  males  and  13  females  from  Cuba;  Anthony 
(1917,  1918,  1925),  cranial  measurements  of  two  specimens  from  Cuba;  Miller  (1929), 
cranial  measurements  of  the  holotype  of  P.  p.  ohtusci  and  an  additional  specimen  from 
Haiti;  G.  M.  Allen  (1942),  cranial  measurements  for  P.  p.  obtusa;  Hall  and  Kelson  (1959), 
cranial  measurements  of  the  holotype  of  P.  p.  obtusa  and  two  specimens  of  P.  p.  poeyi; 
Silva-Taboada  (1974),  measurements  of  fossil  humeri,  crania,  and  mandibles  from  Cuba. 

Subfamily  Desmodontinae 
Desmodus  rotundus  (E.  Geoffroy  St.-Hilaire,  1810) 

Measurements  of  Desmodus  rotundus  have  been  recorded  as  follows:  Dobson  (1878c/), 
external  measurements  of  one  specimen;  Flower  and  Lydekker  (1891),  forearm  length  of 
the  species;  Jentink  (1893),  external  measurements  probably  of  a  female  from  Guyana;  H. 
Allen  (1896),  cranial  measurements  of  a  single  specimen;  Cabrera  (1903),  external  measure¬ 
ments  for  the  species  in  Chile;  Elliot  (1904),  external  and  cranial  measurements  of  one 
specimen;  J.  A.  Allen  (1906),  external  measurements  (mean,  range)  of  five  specimens 
from  Jalisco;  Miller  (1912),  external  and  cranial  measurements  of  a  female  from  Taboga 
Island,  Panama;  Lima  (1926),  external  and  cranial  measurements  of  a  specimen  from 
Brazil;  Goodwin  (1934),  external  measurements  of  one  specimen  from  Guatemala;  Martinez 
and  Villa-R.  (1940),  external  and  cranial  measurements  of  males  and  females  combined 
from  Guerrero;  Cunha  Vieira  (1942),  external  measurements  of  four  males  and  four  females 
and  cranial  measurements  of  three  males  and  one  female  from  Brazil;  Goodwin  (1942 a), 
external  and  cranial  measurements  of  two  females  from  Honduras;  Osgood  (1943),  fore¬ 
arm  measurements  of  two  specimens  from  Chile;  Goodwin  (1946),  external  and  cranial 
measurements  of  a  male  and  female  from  Costa  Rica;  Hershkovitz  (1949),  external  and 
cranial  measurements  (range)  of  14  females  and  a  large  male  obtained  in  a  sample  from 
Colombia;  Dalquest  (1953//),  external  measurements  (mean)  of  10  males  and  10  females 
and  cranial  measurements  of  one  male  and  one  female  from  San  Luis  Potosi;  de  la  Torre 
(1954),  external  and  cranial  measurements  of  a  female  from  Tamaulipas;  de  la  Torre 
(1955),  forearm  measurements  of  one  male  and  one  female  from  Guerrero;  Felten  (1956c), 
external  measurements  (mean,  range)  of  33  males  and  23  females  and  cranial  measurements 
(mean,  range)  of  19  females  and  eight  females  from  El  Salvador;  Felten  (1956//),  cranial 
measurements  of  a  single  specimen  from  El  Salvador;  Jones  (1958),  cranial  measurements 
(mean,  range)  of  three  males  and  seven  females  (combined)  from  Tamaulipas;  Koopman 
(1958),  cranial  measurements  of  a  sub-Recent  fossil  from  Cuba  and  the  range  of  these 
measurements  in  seven  specimens  from  Tamaulipas;  Hall  and  Kelson  (1959),  external  and 
cranial  measurements  of  a  male  and  female  from  Costa  Rica;  Burt  and  Stirton  (1961),  fore¬ 
arm  and  cranial  measurements  (range)  of  14  males  and  23  females;  Goodwin  and  Green- 
hall  (1961),  forearm  measurements  (range)  of  15  males  and  16  females  and  cranial  measure¬ 
ments  of  one  male  and  one  female  from  Trinidad;  Husson  (1962),  external  and  cranial 
measurements  of  a  male  and  five  females  from  Surinam;  Tamsitt  and  Valdivieso  (1962), 
external  measurements  of  a  male  from  Colombia  and  a  large  male  reported  from 
Colombia  by  Hershkovitz  (1949);  Tamsitt  and  Valdivieso  (1963//),  external  measurements 
of  one  male  and  one  female  from  Colombia;  Valdivieso  (1964),  external  measurements  of  a 


70 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


specimen  from  Colombia;  Aellen  (1965),  forearm  measurements  of  two  males,  the  range  of 
eight  females,  and  cranial  measurements  of  one  male  from  Peru;  Brosset  (1965),  external 
measurements  of  two  males  and  a  female  and  cranial  measurements  of  a  male  and  female  from 
Ecuador;  Tamsitt  and  Valdivieso  ( 1 966c/),  forearm  and  cranial  measurements  of  one  male 
and  the  range  of  four  females  from  Colombia;  Villa-R.  (1967),  external  measurements 
(mean,  sd,  range)  of  53  specimens  and  cranial  measurements  (mean,  sd,  range)  of  42  specimens 
from  Mexico;  Genoways  and  Jones  (1968),  forearm  measurements  (mean,  range)  of  10 
young  specimens  (seven  males,  three  females)  from  Zacatecas;  Goodwin  (1969),  forearm 
and  cranial  measurements  of  seven  males  and  seven  females  from  Oaxaca;  Anderson  (1972), 
external  measurements  (mean,  sd,  range)  of  21  specimens  and  cranial  measurements 
(mean,  sd,  range)  of  six  specimens  from  Chihuahua;  Smith  and  Genoways  (1974),  external 
and  cranial  measurements  of  a  male  from  Margarita  Island,  Venezuela,  and  mean  and 
range  of  four  males  from  the  adjacent  mainland;  Woloszyn  and  Mayo  (1974),  cranial  measure¬ 
ments  of  the  holotype  of  the  sub-Recent  D.  r.  puntajudensis  from  Cuba,  one  sub-Recent 
specimen  from  Mexico,  10  Recent  specimens  (mean,  range)  from  Mexico,  and  measure¬ 
ments  after  Koopman  (1958)  and  Husson  (1962). 

Individual  variation. — In  specimens  from  Guerrero,  coefficients  of  variation  for  external 
measurements  of  sexes  combined  varied  from  2.51  to  16.80  and  for  cranial  measurements 
from  1.48  to  4.41  (Martinez  and  Villa-R.,  1940). 

Secondary  sexual  variation. — Hershkovitz  (1949)  noted  that  males  were  smaller  than 
females,  and  Husson  (1962)  concluded  from  published  accounts  that  males  were  smaller  than 
females. 

Geographic  variation.— Measurements  of  individuals  from  Surinam  agreed  well, 
according  to  Husson  (1962),  with  those  from  Colombia  (Hershkovitz,  1949)  and  Trinidad 
(Goodwin  and  Greenhall,  1961). 

Diaemus  youngii  (Jentink,  1893) 

Measurements  of  Diaemus  youngii  have  been  recorded  as  follows:  Jentink  (1893), 
external  measurements  of  the  male  holotype  of  D.  y.  youngii  from  Guyana;  Thomas  (19286), 
external  and  cranial  measurements  of  the  female  holotype  of  D.  y.  cypselinus  from  Peru; 
Cunha  Vieira  (1942),  external  and  cranial  measurements  of  a  male  and  female  from  Brazil; 
Sanborn  (1949),  external  and  cranial  measurements  of  one  specimen  from  Venezuela  and 
another  from  Peru;  Goodwin  and  Greenhall  (1961),  forearm  measurements  of  one  male 
and  two  females  and  cranial  measurements  of  one  male  and  female  from  Trinidad;  Husson 
(1962),  external  and  cranial  measurements  of  the  holotype  from  Guyana;  Lay  (1962), 
external  and  cranial  measurements  of  a  male  and  female  from  Tabasco;  Villa-R.  (1965), 
external  and  cranial  measurements  of  a  female  from  Tamaulipas;  Villa-R.  (1967),  external 
and  cranial  measurements  of  a  specimen  from  Mexico;  Gardner  et  al.  (1970),  external  and 
cranial  measurements  of  a  male  from  Costa  Rica;  Smith  and  Genoways  (1974),  external 
and  cranial  measurements  of  one  specimen  from  Margarita  Island,  Venezuela,  three  males 
(mean,  range)  and  one  female  from  the  adjacent  mainland,  and  the  holotype  of  D.  youngii. 

Geographic  variation. — Gardner  et  al.  (1970)  reported  that  measurements  of  their 
Costa  Rican  specimen  were  much  larger  than  the  holotype  of  D.  y.  youngii  from  Guyana  but 
that  it  agreed  closely  with  the  holotype  of  D.  y.  cypselinus  from  Peru  and  with  a  specimen 
from  Tamaulipas  recorded  by  Villa-R.  (1965).  Measurements  of  two  specimens  from 
Tabasco  (Lay,  1962)  were  somewhat  larger  than  those  of  a  specimen  from  Costa  Rica 
(Gardner  et  al.,  1970). 


Diphylla  ecaudata  Spix,  1823 

Measurements  of  Diphylla  ecaudata  are  recorded  as  follows:  Dobson  (1878a),  external 
measurements  of  a  specimen  from  Brazil;  H.  Allen  (1896),  external  measurements  of  two 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


71 


specimens  and  cranial  measurements  of  one  from  Mexico;  Thomas  (1903/)),  external  and 
cranial  measurements  of  the  male  holotype  of  D.  e.  centralis  from  Panama;  Elliot  (1904), 
external  and  cranial  measurements  of  the  male  holotype  of  D.  e.  centralis  from  Panama 
(after  Thomas,  1903  6)  and  another  specimen;  Lima  (1926),  external  measurements  of  a 
specimen  from  Brazil;  Sanborn  (1936),  external  and  cranial  measurements  of  one  female 
from  Ecuador;  Cunha  Vieira  (1942),  external  and  cranial  measurements  of  a  male  from 
Brazil;  Goodwin  (1942a),  external  and  cranial  measurements  of  two  males  from  Honduras; 
Goodwin  (1946),  external  and  cranial  measurements  of  two  males  from  Honduras 
(as  given  by  Goodwin,  1942a)  and  the  holotype  of  D.  e.  centralis  from  Panama;  Dalquest 
(1950),  cranial  measurements  (mean)  of  seven  males  and  three  females  from  San  Luis 
Potosi;  Dalquest  (1953a),  external  measurements  (mean)  for  two  males  and  13  females 
and  cranial  measurements  (mean)  of  seven  males  and  three  females  from  San  Luis 
Potosi;  de  la  Torre  (1954),  external  and  cranial  measurements  of  a  male  from  Tamaulipas; 
Felten  (1956c),  cranial  measurements  of  five  males  from  El  Salvador;  Felten  ( 1 956c/), 
external  measurements  of  one  specimen  from  El  Salvador;  Hall  and  Kelson  (1959),  external 
and  cranial  measurements  of  the  holotype  of  D.  e.  centralis ,  Burt  and  Stirton  (1961), 
forearm  and  cranial  measurements  (range)  of  six  males  and  nine  females  from  El  Salvador; 
Villa-R.  (1967),  external  measurements  of  20  specimens  and  cranial  measurements  of  19 
from  Mexico;  Reddell  (1968),  external  measurements  of  one  female  from  Texas;  Goodwin 
(1969),  forearm  and  cranial  measurements  of  a  male  from  San  Luis  Potosi  and  a  female 
from  Yucatan;  Ojasti  and  Linares  (1971),  forearm  measurements  (mean,  se,  range)  of  16 
males  and  10  females  and  cranial  measurements  of  10  males  and  nine  females  from 
Venezuela;  Starrett  (1976),  forearm  measurement  of  a  single  female  from  Costa  Rica. 

Geographic  variation. — Ojasti  and  Linares  (1971)  compared  length  of  forearm  and 
length  of  skull  of  specimens  of  Diphylla  ecaudata  from  Central  and  South  America.  They 
concluded  that  these  populations  were  sufficiently  distinct  to  warrant  recognition  as 
separate  subspecies. 


Literature  Cited 

Aellen,  V.  1965.  Sur  une  petite  collection  de  Chiropteres  du  nordouest  du  Perou. 
Mammalia,  29:563-571. 

Allen,  G.  M.  1906.  Vertebrata  from  Yucatan.  Mammalia.  Bull.  Mus.  Comp.  Zool., 
50:106-109. 

- .  1908.  Notes  on  Chiroptera.  Bull.  Mus.  Comp.  Zool.,  52:25-62. 

- .  1911.  Mammals  of  the  West  Indies.  Bull.  Mus.  Comp.  Zool.,  54:175-263. 

- .  1917.  Two  undescribed  West  Indian  bats.  Proc.  Biol.  Soc.  Washington,  30: 

165-170. 

- .  1929.  Mammals.  Pp.  129-130,  in  Vertebrates  from  the  Corn  Islands.  Bull. 

Mus.  Comp.  Zool.,  69:127-146. 

- .  1942.  Extinct  and  vanishing  mammals  of  the  Western  Hemisphere.  Spec. 

Publ.  Amer.  Committee  Internat.  Wild  Life  Protection,  ll:xv  +  1-620. 

Allen,  H.  1861.  Description  of  a  new  Mexican  bat.  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
13:359-361. 

- .  1864.  Monograph  of  the  bats  of  North  America.  Smithsonian  Misc.  Coll., 

165:xxiii+  1-85. 

- .  1890a.  Description  of  a  new  species  of  Macrotus.  Proc.  Amer.  Phil.  Soc., 

28:72-74. 

- .  18906.  Description  of  a  new  species  of  bat  of  the  genus  Carollia,  and  remarks 

on  Carollia  brevicauda.  Proc.  U.S.  Nat.  Mus.,  13:291-298. 

- .  1891.  Description  of  a  new  species  of  Vampyrops.  Proc.  Acad.  Nat.  Sci. 

Philadelphia,  43:400-405. 

- .  1892.  Description  of  a  new  genus  of  phyllostome  bats.  Proc.  U.S.  Nat.  Mus., 

15:441-442. 


72 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


- .  1894a.  A  monograph  of  the  bats  of  North  America.  Bull.  U.S.  Nat.  Mus., 

43:ix  +  1-198. 

- .  1894 6.  On  a  new  species  of  Ametrida.  Proc.  Boston  Soc.  Nat.  Hist.,  26:240-246. 

- .  1895.  Description  of  a  new  species  of  bat  of  the  genus  Glossophaga.  Proc. 

U.S.  Nat.  Mus.,  18:779-781. 

- .  1896.  Notes  on  the  vampire  bat  ( Diphylla  ecaudata )  with  special  reference  to 

its  relationships  with  Desmodus  rufus.  Proc.  U.S.  Nat.  Mus.,  18:769-777. 

- .  1898.  The  skull  and  teeth  of  Ectophylla  alba.  Trans.  Amer.  Philos.  Soc., 

19:267-272. 

Allen,  J.  A.  1897.  Additional  notes  on  Costa  Rican  mammals,  with  description  of  new 
species.  Bull.  Amer.  Mus.  Nat.  Hist.,  9:31-44. 

- .  1900.  List  of  bats  collected  by  Mr.  H.  H.  Smith  in  the  Santa  Marta  region  of 

Colombia,  with  descriptions  of  new  species.  Bull.  Amer.  Mus.  Nat.  Hist.,  13:87-94. 

- .  1904.  New  bats  from  tropical  America  with  notes  on  species  of  Otopterus.  Bull. 

Amer.  Nat.  Hist.,  20:227-237. 

- .  1906.  Mammals  from  the  states  of  Sinaloa  and  Jalisco,  Mexico,  collected  by 

J.  H.  Batty  during  1904  and  1905.  Bull.  Amer.  Mus.  Nat.  Hist.,  22:191-262. 

- .  1908a.  Mammalogical  notes.  II.  Bat  from  the  island  San  Domingo.  Bull. 

Amer.  Mus.  Nat.  Hist.,  24:579-589. 

- .  19086.  Mammals  from  Nicaragua.  Bull.  Amer.  Mus.  Nat.  Hist.,  24:647-670. 

- .  1910.  Mammals  from  the  Cauva  District  of  Venezuela,  with  description  of  a 

new  species  of  Chrotopterus.  Bull.  Amer.  Mus.  Nat.  Hist.,  28:45-149. 

Allen,  J.  A.,  and  F.  M.  Chapman.  1893.  On  a  collection  of  mammals  from  the  island 
of  Trinidad,  with  descriptions  of  new  species.  Bull.  Amer.  Mus.  Nat.  Hist., 
5:203-234. 

- .  1897a.  On  mammals  from  Yucatan,  with  descriptions  of  new  species.  Bull. 

Amer.  Mus.  Nat.  Hist.,  9:1-12. 

- .  18976.  On  a  second  collection  of  mammals  from  the  island  of  Trinidad,  with 

descriptions  of  new  species,  and  a  note  on  some  mammals  from  the  island  of 
Dominica,  West  Indies.  Bull.  Amer.  Mus.  Nat.  Hist.,  9:13-30. 

Alvarez,  T.  1963.  The  Recent  mammals  of  Tamaulipas,  Mexico.  Univ.  Kansas  Publ., 
Mus.  Nat.  Hist.,  14:363-473. 

Alvarez,  T.,  and  J.  Ramirez-Pulido.  1972.  Notes  acera  de  murcielagos  Mexicanos. 

An.  Esc.  Nac.  Cienc.  Biol.,  Mexico,  19:167-178. 

Andersen,  K.  1906a.  On  the  bats  of  the  genera  Micronycteris  and  Glyphonycteris.  Ann. 
Mag.  Nat.  Hist.,  ser.  7,  18:50-65. 

- .  19066.  Brief  diagnoses  of  a  new  genus  and  ten  new  forms  of  stenodermatous 

bats.  Ann.  Mag.  Nat.  Hist.,  ser.  7,  18:419-423. 

- .  1908.  A  monograph  of  the  chiropteran  genera  Uroderma,  Enchisthenes,  and 

Artibeus.  Proc.  Zool.  Soc.  London,  pp.  204-319. 

Anderson,  S.  1957.  New  records  of  the  bat,  Anoura  geoffroyi  lasiopyga.  Chicago  Acad. 
Sci.  Nat.  Hist.  Misc.,  159:1-3. 

- .  1960.  Neotropical  bats  from  western  Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat. 

Hist.,  14:1-8. 

- .  1969.  Macrotus  waterhousi.  Mammalian  Species,  1: 1-4. 

- .  1972.  Mammals  of  Chihuahua:  taxonomy  and  distribution.  Bull.  Amer.  Mus. 

Nat.  Hist.,  148:149-410. 

Anderson,  S.,  and  C.  E.  Nelson.  1965.  A  systematic  revision  of  Macrotus  (Chiroptera). 
Amer.  Mus.  Novit.,  2212:1-39. 

Anthony,  H.  E.  1917.  Two  new  fossil  bats  from  Porto  Rico.  Bull.  Amer.  Mus.  Nat. 
Hist.,  37:565-568. 

- .  1918.  The  indigenous  land  mammals  of  Porto  Rico,  living  and  extinct.  Mem. 

Amer.  Mus.  Nat.  Hist.,  new  ser.,  2(2):33 1-435. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


73 


- .  1919.  Mammals  collected  in  eastern  Cuba  in  1917  with  descriptions  of  two  new 

species.  Bull.  Amer.  Mus.  Nat.  Hist.,  41:625-643. 

- .  1920.  New  rodents  and  new  bats  from  neotropical  regions.  J.  Mamm.,  1: 

81-86. 

- .  1921.  Preliminary  report  on  Ecuadorean  mammals.  No.  1.  Amer.  Mus.  Novit., 

20:1-6. 

- .  1923.  Preliminary  report  on  Ecuadorean  mammals.  No.  3.  Amer.  Mus.  Novit., 

55:1-14. 

- .  1924a.  Preliminary  report  on  Ecuadorean  mammals.  No.  4.  Amer.  Mus.  Novit., 

114:1-6. 

- .  19246.  Preliminary  report  on  Ecuadorean  mammals.  No.  6.  Amer.  Mus. 

Novit.,  139:1-9. 

- .  1925.  Mammals  of  Porto  Rico,  living  and  extinct — Chiroptera  and  Insectivora. 

Scientific  Survey  of  Porto  Rico  and  the  Virgin  Islands,  New  York  Acad.  Sci., 
9:1-238. 

Axtell,  R.  W.  1962.  An  easternmost  record  for  the  bat  Choeronycteris  mexicana  from 
Coahuila,  Mexico.  Southwestern  Nat.,  7:76. 

Baird,  S.  F.  1858.  Description  of  a  phyllostome  bat  from  California,  in  the  Museum  of 
the  Smithsonian  Institution.  Proc.  Acad.  Nat.  Sci.  Philadelphia,  10:1 16-1 17. 

Baker,  R.  H.  1956.  Mammals  of  Coahuila,  Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat. 
Hist.,  9:125-335. 

- .  1960.  Geoffroy’s  tailless  bat  in  Durango.  J.  Mamm.,  41:51 1-514. 

- .  1974.  Records  of  mammals  from  Ecuador.  Publ.  Mus.,  Michigan  State  Univ. 

Biol.  Ser.,  5:129-146. 

Baker,  R.  H.,  and  J.  K.  Greer.  1962.  Mammals  of  the  Mexican  state  of  Durango.  Publ. 

Mus.,  Michigan  State  Univ.  Biol.  Ser.,  2:25-154. 

Baker,  R.  J.,  and  E.  L.  Cockrum.  1966.  Geographic  and  ecological  range  of  the  long- 
nosed  bats,  Leptonycteris.  J.  Mamm.,  47:329-331. 

Baker,  R.  J.,  and  H.  H.  Genoways.  1976.  A  new  species  of  Chiroderma  from 
Guadeloupe,  West  Indies  (Chiroptera:  Phyllostomatidae).  Occas.  Papers  Mus., 
Texas  Tech  Univ.,  39:1-9. 

Baker,  R.  J.,  and  J.  K.  Jones,  Jr.  1975.  Additional  records  of  bats  from  Nicaragua,  with  a 
revised  checklist  of  Chiroptera.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  32:1-13. 
Baker,  R.  J.,  and  G.  Lopez.  1968.  Notes  on  some  bats  from  Tamaulipas.  Southwestern 
Nat.,  13:361-362. 

Baker,  R.  J.,  and  V.  R.  McDaniel.  1972.  A  new  subspecies  of  Uroderma  bilobatum 
(Chiroptera:  Phyllostomatidae)  from  Middle  America.  Occas.  Papers  Mus., 
Texas  Tech  Univ.,  7:1-4. 

Baker,  R.  J.,  W.  R.  Atchley,  and  V.  R.  McDaniel.  1972a.  Karyology  and  morphometries 
of  Peters’  tentmaking  bat,  Uroderma  bilobatum  Peters  (Chiroptera,  Phyllostomati¬ 
dae).  Syst.  Zool.,  21:414-429. 

Baker,  R.  J.,  H.  H.  Genoways,  and  A.  Cadena.  19726.  The  phyllostomatid  bat,  Vampyres- 
sa  brock i,  in  Colombia.  Bull.  S.  California  Acad.  Sci.,  71:54. 

Baker,  R.  J.,  H.  H.  Genoways,  W.  J.  Bleier,  and  J.  W.  Warner.  1973.  Cytotypes  and 
morphometries  of  two  phyllostomatid  bats,  Micronycteris  hirsuta  and  Vampyressa 
pusilla.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  17:1-10. 

Bangs,  O.  1899.  A  new  bat  from  Colombia.  Proc.  New  England  Zool.  Club.,  1:73-74. 
Barbour,  R.  W„  and  W.  H.  Davis.  1969.  Bats  of  America.  Univ.  Press  Kentucky, 

286  pp. 

Birney,  E.  C.,  J.  B.  Bowles,  R.  M.  Timm,  and  S.  L.  Williams.  1974.  Mammal  distri¬ 
butional  records  in  Yucatan  and  Quintana  Roo,  with  comments  on  reproduction, 
structure,  and  status  of  peninsular  populations.  Occas.  Papers  Bull.  Mus.  Nat. 
Hist.,  Univ.  Minnesota,  13:1-25. 


74 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Brosset,  A.  1965.  Contribution  a  L’etude  des  Chiropteres  de  L'ouest  de  L'ecuador. 
Mammals,  29:209-227. 

Buden,  D.  W.  1975a.  Monophyllus  redmani  Leach  (Chiroptera)  from  the  Bahamas,  with 
notes  on  variation  in  the  species.  J.  Mamm.,  56:369-377. 

- .  19756.  A  taxonomic  and  zoogeographic  appraisal  of  the  big-eared  bat  ( Macrotus 

waterhousi Gray)  in  the  West  Indies.  J.  Mamm.,  56:758-769. 

- .  1976.  A  review  of  the  bats  of  the  endemic  West  Indian  genus  Erophylla.  Proc. 

Biol.  Soc.  Washington,  89:1-16. 

- .  1977.  First  records  of  bats  of  the  genus  Brachyphyllci  from  the  Caicos  islands, 

with  notes  on  geographic  variation.  J.  Mamm.,  58:221-225. 

Burt,  W.  H.,  and  R.  A.  Stirton.  1961.  The  mammals  of  El  Salvador.  Misc.  Publ.  Mus. 
Zool.,  Univ.  Michigan,  117:1-69. 

Butterworth,  B.  B.,  and  A.  Starrett.  1964.  Mammals  collected  by  the  Los  Angeles 
County  Museum  expedition  to  northeastern  Venezuela.  Los  Angeles  Co.  Mus., 
Contrib.  Sci.,  85:1-8. 

Cabrera,  A.  1903.  Sinopsis  de  los  quiropteros  Chilenos.  Rev.  Chilena  Hist.  Nat., 
7:278-308. 

- .  1917.  Mamlferos  del  Viaje  al  Pacifico.  Trab.  Mus.  Nac.  Cien.  Nat.  Zool.,  Ma¬ 
drid,  31:1-62. 

Carter,  D.  C.  1966.  A  new  species  of  Rhinophylla  (Mammalia,  Phyllostomatidae)  from 
South  America.  Proc.  Biol.  Soc.  Washington,  79:235-238. 

- .  1968.  A  new  species  of  Anoura  (Mammalia:  Chiroptera:  Phyllostomatidae)  from 

South  America.  Proc.  Biol.  Soc.  Washington,  81:427-430. 

Carter,  D.  C.,  and  C.  S.  Rouk.  1973.  Status  of  recently  described  species  of  Vamprops 
(Chiroptera:  Phyllostomatidae).  J.  Mamm.,  54:975-977. 

Carter,  D.  C.,  R.  H.  Pine,  and  W.  B.  Davis.  1966.  Notes  on  Middle  American  bats. 
Southwestern  Nat.,  1 1:488-499. 

Casebeer,  R.  S„  R.  B.  Linsky,  and  G.  E.  Nelson.  1963.  The  phyllostomatid  bats, 
Ectophylla  alba  and  Vampyrum  spectrum,  in  Costa  Rica.  J.  Mamm.,  44: 186-189. 
Choate,  J.  R.,  and  E.  C.  Birney.  1968.  Sub-Recent  Insectivora  and  Chiroptera  from 
Puerto  Rico,  with  description  of  a  new  bat  of  the  genus  Stenoderma.  J.  Mamm., 
49:400-412. 

Cunha  Vieira,  C.  O.  da.  1942.  Ensaio  monografico  sobre  os  Quiropteros  do  Brasil. 

Arquinos  Zool.,  Estado  de  Sao  Paulo,  3:219-471. 

Cope,  E.  D.  1889.  On  the  Mammalia  obtained  by  the  Naturalist  Exploring  Expedition  to 
southern  Brazil.  Amer.  Nat.,  23:128-150. 

Dalquest,  W.  W.  1950.  Records  of  mammals  from  the  Mexican  state  of  San  Luis 
Potosl.  Occas.  Papers  Mus.  Zool.,  Louisiana  State  Univ.,  23:1-15. 

- .  1951.  Bats  from  the  island  of  Trinidad.  Proc.  Louisiana  Acad.  Sci.,  14:26-33. 

- .  1953a.  The  mammals  of  the  Mexican  state  of  San  Luis  Potosl.  Louisiana  State 

Univ.  Studies,  Biol.  Ser.,  1:1-229. 

- .  19536.  Mexican  bats  of  the  genus  Artibeus.  Proc.  Biol.  Soc.  Washington,  66:61- 

66. 

- .  1957.  American  bats  of  the  genus  Minion.  Proc.  Biol.  Soc.  Washington,  70:45- 

47. 

Davis,  B.  L.,  and  R.  J.  Baker.  1974.  Morphometries,  evolution,  and  cytotaxonomy  of 
mainland  bats  of  the  genus  Macrotus  (Chiroptera:  Phyllostomatidae).  Syst. 
Zool.,  23:26-39. 

Davis,  W.  B.  1944.  Notes  on  Mexican  mammals.  J.  Mamm.,  25:370-403. 

- .  1968.  Review  of  the  genus  Urodernm  (Chiroptera).  J.  Mamm.,  49:676-698. 

- .  1969.  A  review  of  the  small  fruit  bats  (genus  Artibeus)  of  Middle  America.  I. 

Southwestern  Nat.,  14:15-29. 

- .  1970a.  A  review  of  the  small  fruit  bats  (genus  Artibeus )  of  Middle  America.  II. 

Southwestern  Nat.,  14:389-402. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


75 


- .  1970/?.  The  large  fruit  bats  (genus  Artibeus)  of  Middle  America,  with  a  review  of 

the  Artibeus  jamaicensis  complex.  J.  Mamm.,  51:105-122. 

- .  1975.  Individual  and  sexual  variation  in  Vampyressa  bidens.  J.  Mamm., 

56:262-265. 

Davis,  W.  B.,  and  D.  C.  Carter.  1962«.  Notes  on  Central  American  bats  with  description 
of  a  new'  subspecies  of  Marmoops.  Southwestern  Nat.,  7:64-74. 

- .  1962 b.  Review  of  the  genus  Leptonycteris  (Mammalia:  Chiroptera).  Proc. 

Biol.  Soc.,  Washington,  75:193-198. 

- .  1964.  A  new  species  of  fruit-eating  bat  (genus  Artibeus)  from  Central  America. 

Proc.  Biol.  Soc.  Washington,  77:119-122. 

Davis,  W.  B.,  and  R.  J.  Russell,  Jr.  1952.  Bats  of  the  Mexican  state  of  Morelos.  J. 
Mamm.,  33:234-239. 

Davis,  W.  B.,  D.  C.  Carter,  and  R.  H.  Pine.  1964.  Noteworthy  records  of  Mexican  and 
Central  American  bats.  J.  Mamm.,  45:375-387. 
de  la  Torre,  L.  1952.  An  additional  record  of  the  bat,  Sturnira  ludovici,  in  Mexico.  Nat. 
Hist.,  Miscellanea,  Chicago  Acad.  Sci.,  105:1-2. 

- .  1954.  Bats  from  southern  Tamaulipas,  Mexico.  J.  Mamm.,  35:1 13-1 16. 

- .  1955.  Bats  from  Guerrero,  Jalisco  and  Oaxaca,  Mexico.  Fieldiana  Zool., 

37:695-701. 

- .  1959.  A  new  species  of  bat  of  the  genus  Sturnira  (Phyllostomatidae)  from  the 

island  of  Trinidad,  West  Indies.  Nat.  Hist.,  Miscellanea,  Chicago  Acad.  Sci., 
166:1-6. 

- .  1966.  New  bats  of  the  genus  Sturnira  (Phyllostomatidae)  from  the  Amazonian 

lowlands  of  Peru  and  the  Windward  islands.  West  Indies.  Proc.  Biol.  Soc.  Wash¬ 
ington,  79:267-272. 

de  la  Torre,  L.,  and  A.  Schwartz.  1966.  New  species  of  Sturnira  (Chiroptera:  Phyllosto¬ 
matidae)  from  the  islands  of  Gualeloupe  and  Saint  Vincent,  Lesser  Antilles.  Proc. 
Biol.  Soc.  Washington,  79:297-304. 

Dobson,  G.  E.  1876.  Description  of  a  new  species  of  Macrotus.  Ann.  Mag.  Nat.  Hist., 
ser.  4,  18:436-437. 

- .  1878a.  Catalogue  of  the  Chiroptera  in  the  collection  of  the  British  Museum. 

British  Museum,  London,  xlii  +  1-567  pp. 

- .  18786.  Notes  on  recent  additions  to  the  collection  of  Chiroptera  in  the  Museum 

d’Histoire  Naturelle  at  Paris,  with  description  of  new  and  rare  species.  Proc.  Zool. 
Soc.  London,  pp.  873-880. 

Elliot,  D.  G.  1901.  A  synopsis  of  the  mammals  of  North  America  and  the  adjacent  seas. 
Field  Columbian  Mus.,  Zool.  Ser.,  2:xv+  1-471. 

- .  1904.  The  land  and  sea  mammals  of  Middle  America  and  the  West  Indies. 

Field  Columbian  Mus.,  Zool.  Ser.,  4:xiii +  441-850. 

- .  1905a.  A  check  list  of  mammals  of  the  North  American  continent,  the  West 

Indies  and  neighboring  seas.  Field  Columbian  Mus.,  Zool,  Ser.,  6:iv+  1-761. 

- .  19056.  Descriptions  of  apparently  new  species  and  subspecies  of  mammals  from 

Mexico  and  San  Domingo.  Proc.  Biol.  Soc.  Washington,  18:233-236. 

- .  1906.  Descriptions  of  an  apparently  new  species  of  monkey  of  the  genus 

Presbytis  from  Sumatra,  and  a  bat  of  the  genus  Dermanura  from  Mexico.  Proc. 
Biol.  Soc.  Washington,  19:49-50. 

- .  1917.  A  checklist  of  mammals  of  the  North  American  continent  the  West  Indies 

and  the  neighboring  seas.  Amer.  Mus.  Nat.  Hist.,  New  York,  supple.,  iv+  192  pp. 
Flower,  W.  H.,  and  R.  Lydekker.  1891.  An  introduction  to  the  study  of  mammals 
living  and  extinct.  Adam  and  Charles  Black,  London,  xvi+  1-763  pp. 

Felten,  H.  1956a.  Fledermause  (Mammalia:  Chiroptera)  aus  El  Salvador.  Teil  3. 
Senckenbergiana  Biol.,  37:179-212. 

- .  19566.  Ene  neue  unterart  von  Trachups  cirrhosus  (Mammalia,  Chiroptera)  aus 

Brasilien.  Senckenbergiana  Biol.,  37:369-370. 


76 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


- .  1956c.  Fledermause  (Mammalia,  Chiroptera)  aus  El  Salvador.  Teil  4.  Senck- 

enbergiana  Biol.,  37:341-367. 

- .  1956 d.  Quiropteros  (Mammalia,  Chiroptera)  en  El  Salvador.  Comun.  Inst. 

Trop.  Invest.  Cient.,  5:153-170. 

Findley,  J.  S.,  A.  H.  Harris,  D.  E.  Wilson,  and  C.  Jones.  1975.  Mammals  of  New  Mexico. 

Univ.  New  Mexico  Press,  Albuquerque,  xxii  +  360  pp. 

Gardner,  A.  L.  1962a.  A  new  bat  of  the  genus  Glossophaga  from  Mexico.  Los  Angeles 
Co.  Mus.,  Contrib.  Sci.,  54:1-7. 

- .  19626.  Bat  records  from  the  Mexican  states  of  Colima  and  Nayarit.  J.  Mamm., 

43:102-103. 

- .  1976.  The  distributional  status  of  some  Peruvian  mammals.  Occas.  Papers 

Mus.  Zool.,  Louisiana  State  Univ.,  48: 1-18. 

Gardner,  A.  L.,  and  D.  C.  Carter.  1972a.  A  new  stenodermine  bat  (Phyllostomatidae) 
from  Peru.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  2:1-4. 

- .  19726.  A  review  of  the  Peruvian  species  of  Vampyrops  (Chiroptera:  Phyllosto¬ 
matidae).  J.  Mamm.,  53:72-82. 

Gardner,  A.  L.,  and  J.  P.  O'Neill.  1969.  The  taxonomic  status  of  Sturnirci  bidens 
(Chiroptera:  Phyllostomatidae)  with  notes  on  its  karyotype  and  life  history.  Occas. 
Papers  Mus.  Zool.,  Louisiana  State  Univ.,  38:1-8. 

- .  1971.  A  new  species  of  Sturnirci  (Chiroptera:  Phyllostomatidae)  from  Peru. 

Occas.  Papers  Mus.  Zool.,  Louisiana  State  Univ.,  42:1-7. 

Gardner,  A.  L.,  and  J.  L.  Patton.  1972.  New  species  of  Philander  (Marsupialia: 

Didelphidae)  and  Mimon  (Chiroptera:  Phyllostomatidae)  from  Peru.  Occas. 
Papers  Mus.  Zool.,  Louisiana  State  Univ.,  43:1-12. 

Gardner,  A.  L.,  R.  K.  LaVal,  and  D.  E.  Wilson.  1970.  The  distributional  status  of  some 
Costa  Rican  bats.  J.  Mamm.,  51:712-729. 

Genoways,  H.  H.,  and  R.  J.  Baker.  1972.  Stenoderma  rufum.  Mammalian  Species, 
18:1-4. 

Genoways,  H.  H.,  and  J.  K.  Jones,  Jr.  1968.  Notes  on  bats  from  the  Mexican  state  of 
Zacatecas.  J.  Mamm.,  49:743-745. 

- .  1975.  Additional  records  of  the  stenodermine  bat,  Sturnira  thomasi,  from  the 

Lesser  Antillean  island  of  Guadeloupe.  J.  Mamm.,  56:924-925. 

Genoways,  H.  H.,  R.  J.  Baker,  and  W.  B.  Wyatt.  1973.  Nongeographic  variation  in  the 
long-nosed  bat,  Choeroniscus  intermedins.  Bull.  S.  California  Acad.  Sci., 
72:106-107. 

Goldman,  E.  A.  1914a.  Descriptions  of  five  new  mammals  from  Panama.  Smithsonian 
Misc.  Coll.,  63(5):  1-7. 

- .  19146.  A  new  bat  of  the  genus  Mimon  from  Mexico.  Proc.  Biol.  Soc.  Wash¬ 
ington,  27:75-76. 

- .  1917.  New  mammals  from  North  and  Middle  America.  Proc.  Biol.  Soc. 

Washington,  30:107-116. 

- .  1925.  A  new  bat  of  the  genus  Trachops  from  Guatemala.  Proc.  Biol.  Soc. 

Washington,  38:23-24. 

Goodwin,  G.  G.  1933.  The  external  characters  of  Brachyphylla  pumila  Miller.  J. 
Mamm.,  14:154-155. 

- .  1934.  Mammals  collected  by  A.  W.  Anthony  in  Guatemala,  1924-1928.  Bull. 

Amer.  Mus.  Nat.  Hist.,  68:1-60. 

- .  1938.  A  new  genus  of  bat  from  Costa  Rica.  Amer.  Mus.  Novit.,  976:1-2. 

- .  1940.  Three  new  bats  from  Honduras  and  the  first  record  of  Enchisthenes  harti 

(Thomas)  for  North  America.  Amer.  Mus.  Novit.,  1075:1-3. 

- .  1942a.  Mammals  of  Honduras.  Bull.  Amer.  Mus.  Nat.  Hist.,  79:107-195. 

- .  19426.  A  summary  of  recognizable  species  of  Tonatia  with  descriptions  of  two 

new  species.  J.  Mamm.,  23:204-209. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


77 


- .  1946.  Mammals  of  Costa  Rica.  Bull.  Amer.  Mus.  Nat.  Hist.,  87:271-471. 

- .  1953.  Catalogue  of  type  specimens  of  Recent  mammals  in  the  American  Museum 

of  Natural  History.  Bull.  Amer.  Mus.  Nat.  Hist.,  102:207-412. 

- .  1954.  Mammals  from  Mexico  collected  by  Marian  Martin  for  the  American 

Museum  of  Natural  History.  Amer.  Mus.  Novit.  1689:1-16. 

- .  1958.  Three  new  bats  from  Trinidad.  Amer.  Mus.  Novit.  1877:1-6. 

- .  1963.  American  bats  of  the  genus  Vampyressa,  with  the  description  of  a  new 

species.  Amer.  Mus.  Novit.,  2125:1-24. 

- .  1969.  Mammals  from  the  state  of  Oaxaca,  Mexico,  in  the  American  Museum  of 

Natural  History.  Bull.  Amer.  Mus.  Nat.  Hist.,  141:1-269. 

Goodwin,  G.  G.,  and  A.  M.  Greenhall.  1961.  A  review  of  the  bats  of  Trinidad  and 
Tobago.  Bull.  Amer.  Mus.  Nat.  Hist.,  122:187-301. 

- .  1962.  Two  new  bats  from  Trinidad  with  comments  on  the  status  of  the  genus 

Mesophylla.  Amer.  Mus.  Novit.,  2080:1-18. 

Gray,  J.  E.  1834.  Characters  of  a  new  genus  of  bats  ( Brachyphylla ),  obtained  by  the 
Society  from  the  collection  of  the  late  Rev.  Landsdown  Guilding. .  Proc.  Zool. 
Soc.  London,  pp.  122-123. 

Gundlach,  J.  1872.  Catalogo  de  los  mamfferos  Cubanos.  An.  Soc.  Espanola  Hist. 
Nat.,  1:231-258. 

- .  1877.  Contribucion  a  la  mamalogia  Cubana.  G.  Montiel  and  Co.,  Havana, 


53pp. 

Grinnell,  H.  W.  1918.  A  synopsis  of  the  bats  of  California.  Univ.  California  Publ. 
Zool.,  17:223-404. 

Hahn,  W.  L.  1905.  A  new  bat  from  Mexico.  Proc.  Biol.  Soc.  Washington,  18:247-248. 

- .  1907.  A  review  of  the  bats  of  the  genus  Hemiderma.  Proc.  U.S.  Nat.  Mus., 

32:103-118 

Hall,  E.  R.  1946.  Mammals  of  Nevada.  Univ.  California  Press,  Berkeley  and  Los 
Angeles,  xi  +  7 10  pp. 

J  Hall,  E.  R.,  and  J.  E.  Bee.  1960.  The  red  fig-eating  bat  Stenoderma  rufum  Desmarest 
found  alive  in  the  West  Indies.  Mammalia,  24:67-75. 

Hall,  E.  R.,  and  W.  W.  Dalquest.  1963.  The  mammals  of  Veracruz.  Univ.  Kansas 
Publ.,  Mus.  Nat.  Hist.,  14:165-362. 

Hall,  E.  R.,  and  K.  R.  Kelson.  1959.  The  mammals  of  North  America.  Ronald  Press, 
New  York,  l:xxx  + 1-546+ 79. 

'  Hall,  E.  R„  and  J.  R.  Tamsitt.  1968.  A  new  subspecies  of  the  red  fig-eating  bat  from 
Puerto  Rico.  Life  Sci.  Occas.  Papers,  Royal  Ontario  Mus.,  11:1-5. 

Hall,  E.  R.,  and  B.  Villa-R.  1949.  An  annotated  checklist  of  mammals  of  Michoacan, 
Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  1:431-472. 

Handley,  C.  O.,  Jr.  1960.  Descriptions  of  new  bats  from  Panama.  Proc.  U.S.  Nat.  Mus., 


112:459-479. 

- .  1965.  Descriptions  of  new  bats  ( Chiroderma  and  Artibens )  from  Mexico.  An. 

Inst.  Biol.,  Mexico,  36:297-301. 

- .  1966a  Descriptions  of  new  bats  ( Choeroniscus  and  Rhinophylla )  from  Colombia. 

Proc.  Biol.  Soc.  Washington,  79:83-98. 

- .  19666.  Checklist  of  the  mammals  of  Panama.  Pp.  753-795,  in  Ectoparasites 

of  Panama  (R.  L.  Wenzel  and  V.  J.  Tipton,  eds.),  Field  Mus.  Nat.  Hist.,  Chicago, 
xii  +  861  pp. 

Handley,  C.  O.  Jr.,  and  K.  C.  Ferris.  1972.  Descriptions  of  new  bats  of  the  genus 
Vampyrops.  Proc.  Biol.  Soc.  Washington,  84:519-523. 

Harrison,  D.  L.  1975.  Macrophyllum  macrophyllum.  Mammalian  Species,  62:1-3. 
Harrison,  D.  L.,  and  N.  Pendleton.  1974.  A  second  record  of  Wieds’  long-legged  bat 
( Macrophyllum  macrophyllum  Schinz,  1821,  Chiroptera:  Phyllostomatidae)  in 
El  Salvador,  with  notes  on  the  palate,  reproduction  and  diet  of  the  species. 
Mammalia,  38:689-693. 


78 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Henson,  O.  W.,  Jr.,  and  A.  Novick.  1966.  An  additional  record  of  the  bat  Phyllonycteris 
aphylla.  J.  Mamm.,  47:351-352. 

Hershkovitz,  P.  1949.  Mammals  of  northern  Colombia.  Preliminary  reports  No.  5: 
Bats  (Chiroptera).  Proc.  U.S.  Nat.  Mus.,  99:429-454. 

Hill,  J.  E.  1964.  Notes  on  bats  from  British  Guiana,  with  the  description  of  a  new  genus 
and  species  of  Phyllostomatidae.  Mammalia,  28:553-572. 

Hill,  J.  E.,  and  A.  Bown.  1963.  Occurrence  of  Macrophyllum  in  Ecuador.  J.  Mamm., 
44:588. 

Hoffmeister,  D.  F.  1957.  Review  of  the  long-nosed  bats  of  the  genus  Leptonycteris. 
J.  Mamm.,  38:454-461. 

Homan,  J.  A.,  and  J.  K.  Jones,  Jr.  1975a  Monophyllus  redmani.  Mammalian  Species, 
57:1-3. 

- .  19756.  Monophyllus  plethodon.  Mammalian  Species,  58: 1-2. 

Howe,  H.  F.  1974.  Additional  records  of  Phyllonycteris  aphylla  and  Ariteus  flavescens 
from  Jamaica.  J.  Mamm.,  55:662-663. 

Husson,  A.  M.  1954.  On  Vampyrodes  caraccioloi  (Thomas)  and  some  other  bats  from  the 
Island  of  Tobago  (British  West  Indies).  Zool.  Mededelingen,  33:63-67. 

- .  1958.  Notes  on  the  neotropical  leaf-nosed  bat  Sphaeronycteris  toxophyllum 

Peters.  Arch.  Neerlandaises  Zool.,  13:114-1 19. 

- .  1960.  De  zoogdieren  van  de  Nederlandse  Antillen.  Natuurwetenschaplijke 

werkgroep  Nederlandse  Antillen,  Curacao,  170  pp. 

- .  1962.  The  bats  of  Suriname.  Zool.  Verhand.  Leiden,  58:1-222. 

Jentink,  F.  A.  1893.  On  a  collection  of  bats  from  the  West  Indies.  Leyden  Mus.  Notes, 
15:278-283. 

Jones,  J.  K.,  Jr.  1958.  Pleistocene  bats  from  San  Josecito  Cave,  Nuevo  Leon,  Mexico. 
Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  9:389-396. 

- .  1964.  Bats  from  western  and  southern  Mexico.  Trans.  Kansas  Acad.  Sci., 

67:509-516. 

- .  1966.  Bats  from  Guatemala.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  16:439-472. 

Jones,  J.  K.,  Jr.,  and  T.  Alvarez.  1964.  Additional  records  of  mammals  from  the  Mexican 
state  of  San  Luis  Potosf.  J.  Mamm.,  45:302-303. 

Jones,  J.  K.,  Jr.,  and  W.  Bleier.  1974.  Sanborn’s  long-tongued  bat,  Leptonycteris 
sanborni,  in  El  Salvador.  Mammalia,  38:144-145. 

Jones,  J.  K.,  Jr.,  and  D.  C.  Carter.  1976.  Annotated  checklist,  with  keys  to  subfamilies 
and  genera.  Pp.  7-38,  in  Biology  of  bats  of  the  New  World  family  Phyllosto¬ 
matidae,  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.),  Spec.  Publ. 
Mus.,  Texas  Tech  Univ.,  10:1-218. 

Jones,  J.  K.,  Jr.,  and  P.  B.  Dunnigan.  1965.  Molossops  greenhalli  and  other  bats  from 
Guerrero  and  Oaxaca,  Mexico.  Trans.  Kansas  Acad.  Sci.,  68:461-462. 

Jones,  J.  K.,  Jr.,  and  H.  H.  Genoways.  1973.  Ardops  nichollsi.  Mammalian  Species, 
24:1-2. 

- .  1975.  Sturnira  thomasi.  Mammalian  Species,  68: 1-2. 

Jones,  J.  K.,  Jr.,  and  J.  A.  Homan.  1974.  Hylonycteris  underwoodi.  Mammalian 
Species,  32: 1-2. 

Jones,  J.  K.,  Jr.,  and  T.  E.  Lawlor.  1965.  Mammals  from  Isla  Cozumel,  Mexico,  with 
description  of  a  new  species  of  harvest  mouse.  Univ.  Kansas.  Publ.,  Mus.  Nat. 
Hist.,  16:409-419. 

Jones,  J.  K.,  Jr.,  and  C.  J.  Phillips.  1970.  Comments  on  systematics  and  zoogeography 
of  bats  in  the  Lesser  Antilles.  Studies  on  the  Fauna  of  Curagao  and  other  Caribbean 
Islands,  32:131-145. 

- .  1976.  Bats  of  the  genus  Sturnira  in  the  Lesser  Antilles.  Occas.  Papers  Mus., 

Texas  Tech  Univ.,  40:1-16. 

Jones,  J.  K.,  Jr.,  and  G.  L.  Phillips.  1964.  A  new  subspecies  of  the  fruit-eating  bat, 
Sturnira  ludovici,  from  western  Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist., 
14:475-481. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


79 


Jones,  J.  K.,  Jr.,  and  A.  Schwartz.  1967.  Bredin-Archbold-Smithsonian  Biological 
survey  of  Dominica.  VI.  Synopsis  of  bats  of  the  Antillean  genus  Ardops.  Proc. 
U.S.  Nat.  Mus.,  124(3634):  1-13. 

Jones,  J.  K.,  Jr.,  T.  Alvarez,  and  M.  R.  Lee.  1962.  Noteworthy  mammals  from  Sinaloa, 
Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  14:145-159. 

Jones,  J.  K.,  Jr.,  H.  H.  Genoways,  and  R.  J.  Baker.  1971a.  Morphological  variation  in 
Stenoderma  rufum.  J.  Mamm.,  52:244-247. 

Jones,  J.  K.,  Jr.,  J.  D.  Smith,  and  R.  W.  Turner.  19716.  Noteworthy  records  of  bats  from 
Nicaragua,  with  a  checklist  of  the  chiropteran  fauna  of  the  country.  Occas.  Papers 
Mus.  Nat.  Hist.,  Univ.  Kansas,  2:1-35. 

Jones,  J.  K„  Jr.,  J.  R.  Choate,  and  A.  Cadena.  1972.  Mammals  from  the  Mexican  state 
of  Sinaloa.  II.  Chiroptera.  Occas.  Papers  Mus.  Nat.  Hist.,  Univ.  Kansas, 
6:1-29. 

Jones,  J.  K.,  Jr.,  J.  D.  Smith,  and  H.  H.  Genoways.  1973.  Annotated  checklist  of  mammals 
of  the  Yucatan  Peninsula,  Mexico.  I.  Chiroptera.  Occas.  Papers  Mus.,  Texas 
Tech  Univ.,  13:1-31. 

Koopman,  K.  F.  1958.  A  fossil  vampire  bat  from  Cuba.  Breviora,  Mus.  Comp.  Zool., 
90:1-4. 

- .  1961.  A  collection  of  bats  from  Sinaloa,  with  remarks  on  the  limits  of  the 

neotropical  region  in  northwestern  Mexico.  J.  Mamm.,  42:536-538. 

- .  1968.  Taxonomic  and  distributional  notes  on  Lesser  Antillean  bats.  Amer. 

Mus.  Novit.,  2333:1-13. 

Koopman,  K.  F.,  and  E.  E.  Williams.  1951.  Fossil  chiroptera  collected  by  H.  E.  Anthony 
in  Jamaica,  1919-1920.  Amer.  Mus.  Novit.,  1519:1-29. 

Lichenstein,  H„  and  W.  Peters.  1855.  Uber  neue  merkwurdige  Saugethiere  des 
Koniglichen  zoologischen  Museums.  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin, 
pp.81-99  (for  1854). 

LaVal,  R.  K.  1969.  Records  of  bats  from  Honduras  and  El  Salvador.  J.  Mamm., 
50:819-822. 

- .  1977.  Notes  on  some  Costa  Rican  bats.  Brenesia,  10/1 1:77-83. 

Lay,  D.  M.  1962.  Seis  marmferos  nuevos  para  la  fauna  de  Mexico.  Ann.  Inst.  Bio., 

Mexico,  33:373-377. 

Lima,  J.  O.  1926.  Os  moicegos  de  colleccao  do  Museu  Paulista.  Rev.  Mus.  Paulista, 
Sao  Paulo,  14:43-127. 

Linares,  O.  J.  1969.  Nuevos  murcielagos  para  la  fauna  de  Venezuela  en  al  Museo  de 
Historia  Natural  La  Salle.  Mem.  Soc.  Cienc.  La  Salle,  29(82):37-42. 

Linares,  O.  J.,  and  J.  Ojasti.  1971.  Una  nueva  especie  de  murcielago  del  genero 
Lonchorhina  (Chiroptera:  Phyllostomatidae)  del  sur  de  Venezuela.  Novedades 
Cientificas,  Zool.  Ser.,  36:1-8. 

Lonnberg,  E.  1921.  A  second  contribution  to  the  mammalogy  of  Ecuador  with  some 
remarks  on  Caenolestes.  Ark.  Zool.,  Stockholm,  1 4(4):  1-1 04. 

Lukens,  P.  W.  Jr.,  and  W.  B.  Davis.  1957.  Bats  of  the  Mexican  state  of  Guerrero.  J. 
Mamm.,  38: 1-14. 

Lyon,  M.  W.,  Jr.  1902a.  Description  of  a  new  phyllostome  bat  from  the  Isthmus  of 
Panama.  Proc.  Biol.  Soc.  Washington,  15:83-84. 

- .  19026.  Description  of  a  new  bat  from  Colombia.  Proc.  Biol.  Soc.  Washington, 

15:151-152. 

- .  1906.  Notes  on  the  type  specimen  of  the  bat,  Micronycteris  microtis ,  Miller. 

Ann.Mag.  Nat.  Hist.,  ser.  7,  18:371-372. 

Marinkelle,  C.  J.  1970.  Vampyrops  intermedius  sp.  n.  from  Colombia  (Chiroptera, 
Phyllostomatidae).  Rev.  Brasil  Biol.,  30:49-53. 

Marinkelle,  C.  J.,  and  A.  Cadena.  1971.  Remarks  on  Sturnira  tildae  in  Colombia. 
J.  Mamm.,  52:235-237. 

- .  1972.  Notes  on  bats  new  to  the  fauna  of  Colombia.  Mammalia,  36:50-58. 


80 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Martinez,  L.,  and  B.  Villa-R.  1938.  Contribuciones  al  conocimiento  de  los  murcielagos 
de  Mexico.  I.  An.  Inst.  Biol.,  Mexico,  9:339-360. 

- .  1940.  Segunda  contribucion  al  conocimiento  de  los  murcielagos  Mexicanos.  II. 

Estado  de  Guerrero.  An.  Inst.  Biol.,  Mexico,  11:291-361. 

- .  1941.  Contribucion  al  conocimiento  de  los  murcielagos.  III.  An.  Inst.  Biol., 

Mexico,  12:401-419. 

Matson,  J.  O.,  and  D.  R.  Patten.  1975.  Notes  on  some  bats  from  the  state  of  Zacatecas, 
Mexico.  Contrib.  Sci.,  Los  Angeles  Co.  Mus.  Nat.  Hist.,  263: 1-12. 

Miller,  G.  S„  Jr.  1898.  Descriptions  of  five  new  phyllostome  bats.  Proc.  Acad.  Nat. 
Sci.  Philadelphia,  50:326-337. 

- .  1899.  Two  new  glossophagine  bats  from  the  West  Indies.  Proc.  Biol.  Soc. 

Washington,  13:33-37. 

- .  1900a  The  bats  of  the  genus  Monophyllus.  Proc.  Washington  Acad.  Sci., 

2:31-38. 

- .  19006.  Three  new  bats  from  the  island  of  Curacao.  Proc.  Biol.  Soc.  Washington, 

13:123-127. 

- .  1900c.  Note  on  Micronycteris  brachyotis  (Dobson)  and  M.  microtis  Miller. 

Proc.  Biol.  Soc.  Washington,  13:154-155. 

- .  1902u.  Twenty  new  American  bats.  Proc.  Acad.  Nat.  Sci.  Philadelphia, 

54:389-412. 

- .  19026.  The  external  characters  of  Brachyphylla  nana  Miller.  Proc.  Biol.  Soc. 

Washington,  15:249. 

- .  1904.  Notes  on  the  bats  collected  by  William  Palmer  in  Cuba.  Proc.  U.S.  Nat. 

Mus.,  27:337-348. 

- .  1912.  A  small  collection  of  bats  from  Panama.  Proc.  U.S.  Nat.  Mus.,  42:21-26. 

- .  1913a  Five  new  mammals  from  tropical  America.  Proc.  Biol.  Soc.  Washington, 

26:31-34. 

- .  19136.  Revision  of  the  bats  of  the  genus  Glossophaga.  Proc.  U.S.  Nat.  Mus., 

46:413-429. 

- .  1918.  Three  new  bats  from  Haiti  and  Santo  Domingo.  Proc.  Biol.  Soc. 

Washington,  31:39-40. 

- .  1929.  A  second  collection  of  mammals  from  caves  near  St.  Michel,  Haiti. 

Smithsonian  Misc.  Coll.,  8 1  (9):  1-30. 

- .  1931.  Two  new  South  American  bats.  J.  Mamm.,  12:41 1-412. 

- .  1932.  Two  tropical  bats  new  to  the  fauna  of  Panama.  Proc.  Biol.  Soc.  Washing¬ 
ton,  45: 149. 

Mumford,  R.  E.  1975.  A  specimen  of  Rhinophylla  fischerae  from  Ecuador.  J.  Mamm., 
56:273-274. 

Ojasti,  J.,  and  O.  J.  Linares.  1971.  Adiciones  a  la  fauna  de  murcielagos  de  Venezuela 
con  notas  sobre  las  especies  de  genero  Diclidurus  (Chiroptera).  Acta  Biol. 
Venezuela,  7:421-441. 

Ojasti,  J.,  and  C.  J.  Naranjo.  1974.  First  record  of  Tonatia  nicaraguae  in  Venezuela. 
J.  Mamm.,  55:249. 

Osgood,  W.  H.  1943.  The  mammals  of  Chile.  Field  Mus.  Nat.  Hist.,  Zool.  Ser.,  30:1-268. 
Paradiso,  J.  L.  1967.  A  review  of  the  wrinkle-faced  bats  ( Centurio  senex  Gray),  with 
description  of  a  new  subspecies.  Mammalia,  31:595-604. 

Peters,  W.  1857.  Uber  die  chiropterengattungen  Mormops  und  Phyllostoma.  Monatsb. 
Kon  preuss  Akad.  Wiss.,  Berlin,  pp.  287-310  (for  1856). 

- .  1863.  Nachricht  von  einem  neuen  frugivoren  Flederthiere,  Stenoderma 

( Pygoderma )  microdon  aus  Surinam.  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin, 
pp.  83-85. 

- .  1865a  Fledertheire  ( Vespertilio  soricinus  Pallas,  Choeronycteris  Lichtenst., 

Rhinophylla  pumilio  nov.  gen.,  Artibeus  fallax  nov.  sp.,  A.  concolor  nov.  sp., 
Dermanura  quadrivittatum  nov.  sp.,  Nycteris  grandis  nov.  sp.).  Monatsb. 
Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  351-359. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


81 


- .  18656.  Die  zu  den  Vampyri  gehorigen  Flederthiere  und  die  naturliche  Stellung 

der  Gattung  Antrozous.  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  503-525. 

- .  1865c.  Die  brasilianischen,  von  Spix  beschriebenen  Flederthiere.  Monatsb. 

Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  568-588. 

- .  1865  d.  Einige  weniger  bekannte  Flederthiere  ( Phyllostoma  brachyotum, 

Coelops,  Furia,  Lasionvcteris).  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin, 
pp.  641-648. 

- .  1866a.  Neue  oder  ungenugend  bekannte  Flederthiere  ( Vampyrops ,  Uroderma, 

Chiroderma,  Ametrida,  Tylostoma,  Vespertilio,  Vesperugo)  und  Nager  ( Tylomys, 
Lasiomys).  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  392-41 1. 

- .  18666.  Fernere  Mittheilungen  zur  Kenntnifs  der  Flederthiere,  namentlich  uber 

Arten  der  Leidener  und  Britischen  Museums.  Monatsb.  Kon.  preuss.  Akad.  Wiss., 
Berlin,  pp.  672-681. 

- .  1868.  Die  zu  den  Glossophagcie  gehorigen  Flederthiere  und  eine  neue  Art  der 

Gattung  Col'eura.  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  361-368. 

- .  1869.  Bemerkungen  iiber  neue  oder  weniger  bekannte  Flederthiere,  besonder 

des  Pariser  Museums.  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  391-406. 

- .  1876.  Stenodernui  Geoffroy  und  eine  damit  verwandte  neue  Flederthiere  Gattung, 

Peltorhinus.  Monatsb.  Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  429-434. 

- .  1880.  Eine  Mittheilung  iiber  neue  Flederthiere.  Monatsb.  Kon.  preuss.  Akad. 

Wiss.,  Berlin,  pp.  258-259. 

- .  1882.  Uber  Sphaeronycteris  toxophyllum,  eine  neue  Gattlung  und  Art  der 

frugivoren  blattnasigen  Flederthiere,  aus  dem  tropischen  America.  Monatsb. 
Kon.  preuss.  Akad.  Wiss.,  Berlin,  pp.  987-990. 

Peterson,  R.  L.  1965a.  The  genus  Vampyressa  recorded  from  British  Honduras.  J. 
Mamm.,  46:676. 

- .  19656.  A  review  of  the  bats  of  the  genus  Ametrida ,  family  Phyllostomatidae. 

Life  Sci.  Contrib.,  Royal  Ontario  Mus.,  65:1-12. 

- .  1968.  A  new  bat  of  the  genus  Vampyressa  from  Guyana,  South  America,  with 

a  brief  systematic  review  of  the  genus.  Life  Sci.  Contrib.,  Royal  Ontario  Mus., 
73:1-17. 

- .  1972.  A  second  specimen  of  Vampyressa  brock i  (Stenoderminae:  Phyllosto¬ 
matidae)  from  Guyana,  South  America,  with  further  notes  on  the  systematic 
affinities  of  the  genus.  Canadian  J.  Zool.,  50:457-469. 

Peterson,  R.  L.,  and  P.  Kirmse.  1969.  Notes  on  Vampyrum  spectrum,  the  false  vampire 
bat,  in  Panama.  Canadian  J.  Zool.,  47:140-142. 

Peterson,  R.  L.,  and  J.  R.  Tamsitt.  1968.  A  new  species  of  bat  of  the  genus  Sturnira 
(family  Phyllostomatidae)  from  northwestern  South  America.  Life  Sci.  Occas. 
Papers,  Royal  Ontario  Mus.,  12:1-8. 

Phillips,  C.  J.,  and  J.  K.  Jones,  Jr.  1971.  A  new  subspecies  of  the  long-nosed  bat, 
Hvlonycteris  underwoodi,  from  Mexico.  J.  Mamm.,  52:77-80. 

Pine,  R.  H.  1972.  The  bats  of  the  genus  Carollia.  Texas  A&M  Agric.  Exp.  Sta.,  Tech. 
Monogr.,  8: 1-125. 

Pirlot,  P.  1963.  Algunas  consideraciones  sobre  la  ecolgia  de  los  mamiferos  del  oeste  de 
Venezuela.  Rev.  Universidad  del  Zulia,  Kasmera,  1:169-214. 

- .  1965a.  Deux  formes  nouvelles  de  chiropteres  des  genres  Eumops  et 

Leptonycteris.  Le  Naturaliste  Canadien,  92:5-7. 

- .  19656.  Chiropteres  de  L'est  du  Venezuela.  II.  Delta  de  L’Orenoque.  Mam¬ 
malia,  29:375-389. 

- .  1967.  Nouvelle  recolte  Chiropteres  dans  L'ouest  du  Venezuela.  Mammalia, 

31:260-274. 

- .  1968.  Chiropteres  du  Perou,  specialement  de  haute-Amazone.  Mammalia, 

32:86-96. 

- .  1972.  Chiropteres  de  Moyenne  Amazonie.  Mammalia,  36:71-85. 


82 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Power,  D.  M.,  and  J.  R.  Tamsitt.  1973.  Variation  in  Phyllostomus  discolor  (Chiroptera: 

Phyllostomatidae).  Canadian  J.  Zool.,  51:461-468. 

Ramirez- Pulido,  J.,  and  T.  Alvarez.  1972.  Notes  sobre  los  murcielagos  del  genero 
Leptonycteris  en  Mexico,  con  la  designacion  del  lectotipo  de  L.  yerbubuenae 
Martinez  Y  Villa,  1940.  Southwestern  Nat.  16:249-259. 

Reddell,  J.  R.  1968.  The  hairy-legged  vampire,  Diphylla  ecaudata,  in  Texas.  J.  Mamm., 
49:769. 

Rehn,  J.  A.  G.  1900.  Notes  on  Chiroptera.  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
52:755-759. 

- .  1901.  A  study  of  the  genus  Centurio.  Proc.  Acad.  Sci.  Philadelphia,  53:295-302. 

- .  1902a.  A  new  bat  of  the  genus  Glossophaga.  Proc.  Acad.  Nat.  Sci.  Philadelphia, 

54:37-38. 

- .  19026.  Three  new  American  bats.  Proc.  Acad.  Nat.  Sci.  Philadelphia, 

54:638-641. 

- .  1904.  A  revision  of  the  mammalian  genus  Macrotus.  Proc.  Acad.  Nat.  Sci. 

Philadelphia,  56:427-446. 

Rick,  A.  M.  1968.  Notes  on  bats  from  Tikal,  Guatemala.  J.  Mamm.,  49:5 16-520. 
Robinson,  W.,  and  M.  W.  Lyon,  Jr.  1901.  An  annotated  list  of  mammals  collected  in 
the  vicinity  of  La  Guaire,  Venezuela.  Proc.  U.S.  Nat.  Mus.,  24:135-162. 

Rouk,  C.  S.,  and  D.  C.  Carter.  1972.  A  new  species  of  Vampyrops  (Chiroptera: 

Phyllostomatidae)  from  South  America.  Occas.  Papers  Mus.,  Texas  Tech  Univ., 
1:1-7. 

Russell,  R.  J.  1956.  Artibeus  lituratus  in  Morelos,  Mexico.  J.  Mamm.,  37:283-284. 
Ryan,  R.  M.  1960.  Mamiferos  colectados  en  Guatemala  en  1954.  Acta  Zool.  Mexicana, 
4:1-19. 

Sanborn,  C.  C.  1932.  Neotropical  bats  in  the  Carnegie  Museum.  Ann.  Carnegie  Mus., 
21:171-183. 

- .  1933.  Bats  of  the  genera  Anoura  and  Lonchoglossa.  Field  Mus.  Nat.  Hist., 

Zool.  Ser.,  20:23-27. 

- .  1935.  New  mammals  from  Guatemala  and  Honduras.  Field  Mus.  Nat.  Hist., 

Zool.  Ser.  20:81-85. 

- .  1936.  Records  and  measurements  of  neotropical  bats.  Field  Mus.  Nat.  Hist., 

Zool.  Ser.,  20:93-106. 

- .  1938.  Notes  on  neotropical  bats.  Occas.  Papers  Mus.  Zool.,  Univ.  Michigan, 

373:1-5. 

- .  1941.  Descriptions  and  records  of  neotropical  bats.  Field  Mus.  Nat.  Hist., 

Zool.  Ser.,  27:371-387. 

- .  1943.  External  characters  of  the  bats  of  the  sub-family  Glossophagine.  Field 

Mus.  Nat.  Hist.,  Zool.  Ser.,  24:271-277. 

- .  1949a.  Bats  of  the  genus  Micronycteris  and  its  sub-genera.  Fieldiana  Zool., 

31:215-233. 

- .  19496.  Mammals  from  the  Rio  Ucayali,  Peru.  J.  Mamm.,  30:277-288. 

- .  1951.  Mammals  from  Marcapata,  southeastern  Peru.  Publ.  Mus.  Hist.  Nat. 

“Javier  Prado”,  ser.  A,  Zool.,  6:1-26. 

- .  1953.  Mammals  from  the  departments  of  Cuzco  and  Puno,  Peru.  Publ.  Mus. 

Hist.  Nat.  “Javier  Prado”,  ser.  A,  Zool.,  12:1-8. 

- .  1954.  Bats  from  Chimanta-Tepui,  Venezuela  with  remarks  on  Choeroniscus. 

Fieldiana  Zool.,  34:289-293. 

- .  1955.  Remarks  on  the  bats  of  the  genus  Vampyrops.  Fieldiana,  Zool.,  37:403- 

413. 

Saussure,  H.  de.  1860a.  Note  sur  quelques  mammiferes  de  Mexique.  Rev.  Mag. 
Zool.,  Paris,  ser.  2,  12:377-383. 

- .  18606.  Note  sur  quelques  mammiferes  du  Mexique.  Rev.  Mag.  Zool.,  Paris, 

ser.  2,  12:425-431. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


83 


- .  1860c.  Note  sur  quelques  mammiferes  du  Mexique.  Rev.  Mag.  Zool.,  Paris, 

ser.  2,  12:479-494. 

Schaldach,  W.  J.,  and  C.  A.  McLaughlin.  1960.  A  new  genus  and  species  of  glossophagine 
bat  from  Colima,  Mexico.  Los  Angeles  Co.  Mus.,  Contrib.  Sci.,  37:1-8. 

Schwartz,  A.,  and  J.  K.  Jones,  Jr.  1967.  Bredin-Archbold-Smithsonian  biological 
survey  of  Dominica.  VII.  Review  of  bats  of  the  endemic  Antillean  genus 
Monophyllus.  Proc.  U.S.  Nat.  Mus.,  124:(3635):  1-20. 

Shamel,  H.  H.  1927.  A  new  bat  from  Colombia.  Proc.  Biol.  Soc.  Washington, 
40:129-130. 

- .  1931.  Bats  from  the  Bahamas.  J.  Washington  Acad.  Sci.,  21:251-253. 

Sherman,  H.  B.  1955.  A  record  of  Lasiurus  and  of  Vampyrops  from  Paraguay.  J. 
Mamm.,  36:130. 

Silva-Taboada,  G.  1974.  Fossil  chiroptera  from  cave  deposits  in  central  Cuba,  with 
description  of  two  new  species  (genera  Pteronotus  and  Mormoops)  and  the  first 
West  Indian  record  of  Mormoops  megalophylla.  Acta  Zool.  Cracoviensia, 
19:33-73. 

- .  1976.  Historialy  actualizacion  taxonomica  de  algunas  especies  Antillanas  de 

murcielagos  de  los  generos  Pteronotus,  Brachyphylla,  Lasiurus,  y  Antrozous. 
Poeyana,  153:1-24. 

Smith,  J.  D.,  and  H.  H.  Genoways.  1974.  Bats  of  Margarita  Island,  Venezuela,  with 
zoogeographic  comments.  Bull.  S.  California  Acad.  Sci.,  73:64-79. 

Spenrath,  C.  A.,  and  R.  K.  LaVal.  1970.  Records  of  bats  from  Queretaro  and  San  Luis 
PotosI,  Mexico.  J.  Mamm.,  51:395-396. 

Stains,  H.  J.  1957.  A  new  bat  (genus  Leptonycteris)  from  Coahuila.  Univ.  Kansas 
Publ.,  Mus.  Nat.  Hist.,  9:353-356. 

Starrett,  A.  1969.  A  new  species  of  Anoura  (Chiroptera:  Phyllostomatidae)  from  Costa 
Rica.  Los  Angeles  Co.  Mus.,  Contrib.  Sci.,  157:1-9. 

- .  1976.  Comments  on  bats  newly  recorded  from  Costa  Rica.  Los  Angeles  Co. 

Mus.,  Contrib.  Sci.,  277:1-5. 

Starrett,  A.,  and  R.  S.  Casebeer.  1968.  Records  of  bats  from  Costa  Rica.  Los  Angeles 
Co.  Mus.,  Contrib.  Sci.,  148:1-21. 

Starrett,  A.,  and  L.  de  la  Torre.  1964.  Notes  on  a  collection  of  bats  from  Central 
America,  with  the  third  record  for  Cvttarops  alecto  Thomas.  Zoologica,  New 
York,  49:53-63. 

Stephens,  F.  1906.  California  mammals.  West  Coast  Publ.  Co.,  San  Diego,  California, 
351  pp. 

Taddei,  V.  A.  1975a.  Phyllostomidae  (Chiroptera)  do  Norte-ocidental  do  Estado  de 
Sao  Paulo  I-Phyllostominae.  Ciencia  e  Cultura,  27:621-632. 

- .  19756.  Phyllostomidae  (Chiroptera)  do  Norte-ocidental  do  Estado  de  Sao 

Paulo  II-Glossophaginae;  Carolliinae;  Sturnirinae.  Ciencia  e  Cultura,  27:723-734. 

Tamsitt,  J.  R.,  and  D.  Valdivieso.  1962.  Desmodus  rotundus  rotundus  from  a  high 
altitude  in  southern  Colombia.  J.  Mamm.,  43:106-107. 

- .  1963a.  Records  and  observations  on  Colombian  bats.  J.  Mamm.,  44: 168-180. 

- .  19636.  Notes  on  bats  from  Leticia,  Amazonas,  Colombia.  J.  Mamm.,  44:263. 

- .  1965a.  The  male  reproductive  cycle  of  the  bat  Artibeus  lituratus.  Amer. 

Midland  Nat.,  73:150-160. 

- .  19656.  Reproduction  of  the  female  big  fruit-eating  bat,  Artibeus  lituratus 

palmarum,  in  Colombia.  Caribbean  J.  Sci.,  5:157-166. 

- .  1966a.  Bats  from  Colombia  in  the  Swedish  Museum  of  Natural  History,  Stock¬ 
holm.  Mammalia,  30:97-104. 

- .  19666.  Taxonomic  comments  on  Anoura  caudifer,  Artibeus  lituratus  and 

Molossus  molossus.  J.  Mamm.,  47:230-238. 

- .  1966c.  Parturition  in  the  red  fig-eating  bat,  Stenoderma  rufum.  J.  Mamm., 

47:352-353. 


84 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Thomas,  M.  E.,  and  D.  N.  McMurray.  1974.  Observations  on  Sturnira  aratathomasi 
from  Colombia.  J.  Mamm.,  55:834-836. 

Thomas,  O.  1889.  Description  of  a  new  stenodermatous  bat  from  Trinidad.  Ann.  Mag. 
Nat.  Hist.,  ser.  6,  7:167-170. 

- .  1891a.  Descriptions  of  three  new  bats  in  the  British  Museum  Collection.  Ann. 

Mag.  Nat.  Hist.,  ser.  6,  7:527-530. 

- .  18916.  Note  on  Chiroderma  villosum,  Peters,  with  the  description  of  a  new 

species  of  the  genus.  Ann.  Mus.  Civ.  Genova,  2a.  ser.,  10:881-883. 

- .  1892.  Description  of  a  new  bat  of  the  genus  Artibeus  from  Trinidad.  Ann. 

Mag.  Nat.  Hist.,  ser.  6,  10:408-410. 

- .  1893.  On  some  mammals  from  central  Peru.  Proc.  Zool.  Soc.  London,  pp. 

333-341. 

- .  1894.  Description  of  a  new  bat  of  the  genus  Stenoderma  from  Montserrat.  Proc. 

Zool.  Soc.  London,  pp.  132-133. 

- .  1895.  On  small  mammals  from  Nicaragua  and  Bogota.  Ann.  Mag.  Nat.  Hist., 

ser.  6,  9:55-60. 

- .  1896.  On  new  mammals  from  the  Neotropical  region.  Ann.  Mag.  Nat.  Hist., 

ser.  6,  18:301-314. 

- .  1900.  Descriptions  of  new  Neotropical  mammals.  Ann.  Mag.  Nat.  Hist., 

ser.  7,  5:269-276. 

- .  1901a.  New  Myotis,  Artibeus,  Sylvilagus,  and  Metachirus  from  Central  and 

South  America.  Ann.  Mag.  Nat.  Hist.,  ser.  7,  7:541-545. 

- .  19016.  On  a  collection  of  mammals  from  Kanuku  Mountains,  British  Guiana. 

Ann.  Mag.  Nat.  Hist.,  ser.  7,  8:139-154. 

- .  1901c.  On  a  collection  of  bats  from  Para.  Ann.  Mag.  Nat.  Hist.,  ser.  7, 

8:189-193. 

- .  1903a.  Two  new  glossophagine  bats  from  Central  America.  Ann.  Mag.  Nat. 

Hist.,  ser.  7,  11:286-289. 

- .  19036.  New  mammals  from  Chiriqui.  Ann.  Mag.  Nat.  Hist.,  ser  7,  1 1:376-382. 

- .  1903c.  Notes  on  South  America  monkeys,  bats,  carnivores,  and  rodents,  with 

descriptions  of  new  species.  Ann.  Mag.  Nat.  Hist.,  ser.  7,  12:455-464. 

- .  1909.  Notes  on  some  South-American  mammals,  with  descriptions  of  new 

species.  Ann.  Mag.  Nat.  Hist.,  ser.  8,  4:230-242. 

- .  1910.  Mammals  from  the  River  Supinaam,  Demerara,  presented  by  Mr.  F.  V. 

McConnell  to  the  British  Museum.  Ann.  Mag.  Nat.  Hist.,  ser.  8,  6:184-189. 

- .  1912a.  Three  small  mammals  from  S.  America.  Ann.  Mag.  Nat.  Hist.,  ser.  8, 

9:408-410. 

- .  19126.  New  bats  and  rodents  from  S.  America.  Ann.  Mag.  Nat.  Hist.,  ser.  8, 

10:403-411. 

- .  1913.  A  new  genus  of  glossophagine  bat  from  Colombia.  Ann.  Mag.  Nat. 

Hist.,  ser.  8,  12:270-271. 

- .  1914.  Four  new  small  mammals  from  Venezuela.  Ann.  Mag.  Nat.  Hist., 

ser.  8,  14:410-414. 

- .  1915.  A  new  genus  of  phyllostome  bats  and  a  new  Rhipidomys  from  Ecuador. 

Ann.  Mag.  Nat.  Hist.,  ser.  8,  16:310-312. 

- .  1926.  The  Godman-Thomas  Expedition  to  Peru.  III.  On  mammals  collected 

by  Mr.  R.  W.  Hendee  in  the  Chachapoyas  Region  of  north  Peru.  Ann.  Mag. 
Nat.  Hist.,  ser.  9,  18:156-167. 

- .  1928a.  A  new  genus  and  species  of  glossophagine  bat,  with  a  new  subdivision  of 

the  genus,  Choeronycteris.  Ann.  Mag.  Nat.  Hist.,  ser.  10,  1:120-123. 

- .  19286.  The  Godman-Thomas  Expedition  to  Peru.  VIII.  On  mammals  obtained 

by  Mr.  Hendee  at  Pebas  and  Iquitos,  Upper  Amazons.  Ann.  Mag.  Nat.  Hist., 
ser.  10,  2:285-294. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


85 


Tomes,  R.  F.  1863.  On  a  new  genus  and  species  of  leaf-nosed  bats  in  the  museum  at  Fort 
Pitt.  Proc.  Zool.  Soc.  London,  pp.  81-84. 

Tuttle,  M.  D.  1970.  Distribution  and  zoogeography  of  Peruvian  bats,  with  comments 
on  natural  history.  Univ.  Kansas  Sci.  Bull.,  49:45-86. 

Valdez,  R.,  and  R.  K.  LaVal.  1971.  Records  of  bats  from  Honduras  and  Nicaragua.  J. 
Mamm.,  52:247-250. 

Valdivieso,  D.  1964.  La  fauna  quireoptera  del  deparamento  de  Cundinamarca,  Colombia. 
Rev.  Biol.  Trop.,  12:19-45. 

Valdivieso,  D.,  and  J.  R.  Tamsitt.  1962.  First  records  of  the  pale  spear-nosed  bat  in 
Colombia.  J.  Mamm.,  43:422-423. 

Villa-R.,  B.  1953.  Mamiferos  silvestres  del  Valle  de  Mexico.  An.  Inst.  Biol.,  Mexico, 
23:269-492. 

- .  1962.  Nota  acerca  de  la  distribucion  de  los  murcielagos  Euderma  maculatum 

(J.  A.  Allen)  y  Chiroderma  isthmicum  Miller  en  Mexico.  An.  Inst.  Biol., 
Mexico,  33:379-384. 

- .  1963.  Reflexiones  acerca  de  la  posicion  taxonomica  de  los  murcielagos 

siricoteros  de  Mexico,  genero  Glossophaga.  An.  Inst.  Biol.,  Mexico,  34:381-391. 

- .  1965.  Diaemus  youngi  (Jentink)  el  vampiro,  overo,  en  el  sur  de  Tamaulipas, 

Mexico.  Ann.  Inst.  Biol.,  Mexico,  35:127-128. 

- .  1967.  Los  murcielagos  de  Mexico.  Univ.  Nacional  Autonoma  Mexico,  Inst. 

Biol.,  xvi  +  491  pp. 

Villa-R.,  B.,  and  M.  Villa  Cornejo.  1969.  Algunos  murcielagos  del  norte  de  Argentina. 

Pp.  407-428,  in  Contributions  in  mammalogy  (J.  K.  Jones,  Jr.,  ed.),  Misc.  Publ. 
Mus.  Nat.  Hist.,  Univ.  Kansas,  51:1-428. 

Walton,  D.  W.  1963.  A  collection  of  the  bat  Lonchophylla  robusta  Miller  from  Costa 
Rica.  Tulane  Studies,  Zool.,  10:87-90. 

Ward,  H.  L.  1891.  Descriptions  of  three  new  species  of  Mexican  bats.  Amer.  Nat., 
25:743-753. 

Watkins,  L.  C.,  J.  K.  Jones,  Jr.,  and  H.  H.  Genoways.  1972.  Bats  of  Jalisco,  Mexico. 
Spec.  Publ.  Mus.,  Texas  Tech.  Univ.,  1:1-44. 

Woloszyn,  B.  W.,  and  N.  A.  Mayo.  1974.  Postglacial  remains  of  a  vampire  bat 
(Chiroptera:  Desmodus)  from  Cuba.  Acta  Zool  Cracoviensia,  19:253-265. 


86 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Selected  measurements  of  phyllostomatid  bats.  Museum  acronyms  used  are  as 
follows:  AMNH,  American  Museum  of  Natural  History ;  AS,  Albert  Schwartz  Collection; 
BMNH,  British  Museum  ( Natural  History),  CM,  Carnegie  Museum  of  Natural  History, 
COLU,  Department  of  Biology,  Colgate  University;  FHKSC,  Museum  of  the  High  Plains, 
Fort  Hays  Kansas  State  College;  KU,  Museum  of  Natural  History,  University  of  Kansas; 
LACM,  Natural  History  Museum  of  Los  Angeles  County;  LSU,  Museum  of  Zoology, 
Louisiana  State  University;  ROM,  Royal  Ontario  Museum;  TCWC,  Texas  Cooperative 
Wildlife  Collection,  Texas  A&M  University;  TTU,  The  Museum,  Texas  Tech  University; 

USNM,  National  Museum  of  Natural  History. 


Museum,  catalogue 

no.,  and  sex  Locality 


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Phyllostomatinae 


Chrotopterus  auritus 


KU  23661  2 

Veracruz 

78.9 

36.7 

31.2 

19.2 

5.9 

13.4 

12.9 

12.0 

KU  93385  2 

Yucatan 

78.7 

36.2 

31.1 

18.5 

6.3 

13.3 

12.7 

11.5 

USNM  305204  2 

Panama 

83.1 

37.8 

31.8 

19.5 

6.2 

14.0 

13.5 

12.4 

USNM  335156  2 

Panama 

82.5 

37.1 

31.7 

19.6 

6.2 

14.5 

13.2 

12.6 

TTU  9339  6 

Veracruz 

81.1 

35.7 

30.4 

18.5 

5.9 

12.9 

13.0 

12.0 

KU  23622  d 

Veracruz 

79.1 

35.7 

30.6 

18.1 

6.1 

12.9 

13.0 

11.6 

KU  93383  S 

Yucatan 

80.8 

36.0 

31.0 

18.2 

6.3 

13.3 

12.9 

11.6 

KU  105962  6 

Nicaragua 

79.8 

36.2 

31.0 

18.2 

6.0 

13.2 

12.6 

11.3 

Lonchorhina  aurita 

USNM  305186  2 

Panama 

50.5 

20.0 

17.9 

10.4 

4.8 

8.4 

6.3 

7.0 

TTU  5320  V 

Trinidad 

47.1 

20.8 

19.1 

10.4 

4.9 

8.9 

6.7 

7.1 

TTU  5322  2 

Trinidad 

50.3 

20.5 

18.6 

10.8 

4.9 

8.8 

6.6 

7.0 

TTU  8984  2 

Trinidad 

51.1 

20.6 

18.7 

10.8 

4.9 

8.9 

6.6 

7.0 

TTU  5321  6 

Trinidad 

49.0 

20.7 

18.9 

10.5 

4.9 

8.7 

6.6 

7.0 

TTU  5323  J 

Trinidad 

50.0 

20.4 

19.0 

10.8 

4.8 

8.7 

6.6 

7.1 

TTU  9827  6 

Trinidad 

49.9 

20.7 

18.7 

10.4 

5.0 

8.7 

6.6 

7.1 

TTU  9829  <3 

Trinidad 

49.8 

20.5 

18.7 

10.4 

4.9 

8.7 

6.6 

7.0 

Lonchorhina  orinocensis 

USNM  373254  2 

Venezuela 

42.3 

19.0 

16.4 

9.3 

4.0 

8.0 

5.9 

6.0 

USNM  373255  2 

Venezuela 

42.2 

19.1 

16.4 

9.6 

4.1 

8.2 

5.9 

6.0 

USNM  373256  2 

Venezuela 

43.7 

19.2 

16.8 

9.6 

4.2 

8.0 

6.0 

6.0 

USNM  373260  2 

Venezuela 

41.5 

18.3 

16.2 

9.1 

3.8 

8.2 

5.7 

5.7 

USNM  373248(3 

Venezuela 

41.4 

19.5 

17.0 

9.7 

4.2 

8.3 

6.0 

5.9 

USNM  373249(3 

Venezuela 

42.6 

19.5 

17.0 

9.6 

4.0 

8.0 

6.3 

6.1 

USNM  373257d 

Venezuela 

41.5 

19.3 

17.0 

9.7 

4.0 

8.3 

6.0 

5.9 

USNM  373258(3 

Venezuela 

43.0 

19.5 

16.8 

9.8 

4.0 

8.1 

6.1 

6.1 

Macrophylt um  macrophyll ton 

AMNH  177666  2 

Nicaragua 

36.9 

17.1 

14.5 

9.2 

3.4 

8.2 

5.5 

6.1 

AMNH  177669  2 

Nicaragua 

36.2 

17.0 

14.7 

9.5 

3.1 

7.8 

5.7 

6.4 

AMNH  177670  2 

Nicaragua 

37.4 

17.4 

14.7 

9.5 

3.0 

8.0 

5.5 

6.2 

AMNH  177671  2 

Nicaragua 

37.4 

17.1 

14.7 

9.4 

3.0 

8.1 

5.6 

6.2 

KU  70478  3 

Nicaragua 

35.6 

16.6 

13.6 

9.2 

3.0 

7.8 

5.2 

6.1 

USNM  311944(3 

Panama 

35.0 

16.8 

14.2 

8.9 

3.2 

7.8 

5.5 

6.1 

USNM  312963  6 

Panama 

37.2 

17.7 

14.9 

9.8 

3.2 

8.0 

5.7 

6.7 

USNM  315212(3 

Panama 

34.3 

17.2 

14.2 

10.0 

3.2 

8.0 

5.7 

6.8 

Macrolus  californicus 

FHKSC  2442  2 

Arizona 

49.0 

22.7 

19.9 

10.4 

3.3 

8.1 

8.9 

7.4 

TTU  10529  2 

Sonora 

49.4 

22.3 

19.9 

10.8 

3.5 

8.1 

9.0 

7.0 

TTU  10584  2 

Sonora 

51.8 

22.9 

20.7 

11.2 

3.5 

9.0 

8.8 

7.1 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


87 


Appendix  1. — Continued. 


TTU  10588  2 

Sonora 

51.7 

23.0 

20.4 

11.2 

3.7 

8.4 

8.9 

7.0 

FHKSC  1994  6 

Arizona 

50.7 

23.2 

20.5 

11.4 

3.8 

8.5 

8.7 

7.5 

TTU  10582  6 

Sonora 

49.7 

23.8 

20.1 

11.7 

3.5 

8.5 

9.5 

7.4 

TTU  10585  6 

Sonora 

48.3 

22.6 

20.2 

10.6 

3.6 

8.1 

8.9 

7.0 

TTU  10587  6 

Sonora 

50.0 

23.2 

20.1 

11.1 

3.6 

8.3 

9.2 

7.3 

Macrotus  waterhousii 

TTU  10566  2 

Sonora 

49.2 

22.2 

19.5 

10.6 

3.9 

8.5 

8.4 

7.2 

TTU  21470  2 

Jamaica 

53.9 

25.3 

21.5 

12.2 

4.1 

9.2 

9.4 

7.8 

TTU  21471  2 

Jamaica 

55.0 

25.8 

21.9 

12.5 

4.2 

9.2 

9.7 

8.0 

TTU  21505  2 

Jamaica 

54.1 

26.0 

22.0 

12.0 

4.2 

8.8 

9.6 

7.5 

TTU  6267  6 

Sonora 

47.2 

23.2 

20.0 

11.1 

4.2 

8.6 

8.9 

7.5 

TTU  10564  6 

Sonora 

49.6 

23.0 

20.0 

11.2 

4.1 

8.6 

8.8 

7.4 

TTU  10565  6 

Sonora 

48.3 

22.4 

19.4 

10.9 

4.1 

8.5 

8.6 

7.4 

TTU  21501  6 

Jamaica 

54.7 

26.4 

22.0 

12.4 

4.4 

9.5 

9.7 

7.9 

Micronycteris  behni 

BMNH  69.5.13.3  2 

Peru 

4.7 

8.1 

7.2 

Micronycteris  brachyolis 

USNM  323059  2 

Panama 

42.2 

22.4 

19.3 

11.4 

5.0 

9.0 

8.3 

7.3 

TTU  5237  2 

Trinidad 

39.4 

21.3 

18.5 

10.2 

5.0 

8.6 

8.1 

6.7 

TTU  5315  2 

Trinidad 

40.9 

21.6 

19.0 

10.4 

5.1 

8.4 

8.6 

6.9 

AMNH  175633  2 

Trinidad 

40.3 

21.4 

18.7 

10.5 

5i0 

8.5 

8.1 

6.9 

USNM  245153  (3 

Guatemala 

40.9 

21.7 

19.3 

10.7 

5.1 

8.7 

8.2 

6.9 

USNM  306546<3 

Panama 

40.8 

22.8 

19.9 

11.1 

5.2 

8.9 

8.2 

7.0 

USNM  323060<3 

Panama 

40.2 

21.3 

18.7 

10.8 

4.8 

8.4 

7.9 

6.9 

TTU  5314<3 

Trinidad 

39.4 

21.9 

19.2 

10.5 

5.2 

8.9 

8.2 

7.0 

Micronycteris  daviesi 

BMNH  64.767  2 

Guyana 

57.1 

27.3 

23.7 

13.3 

6.5 

10.8 

11.0 

9.3 

USNM  335104(3 

Panama 

54.0 

27.3 

23.5 

13.2 

6.7 

10.5 

10.7 

9.2 

USNM  460089(3 

Brazil 

54.7 

26.1 

22.8 

12.8 

6.2 

10.6 

10.5 

9.1 

M icronycteris  h  irsuta 

TTU  13158  2 

Nicaragua 

42.5 

23.8 

20.6 

11.8 

5.0 

8.4 

9.4 

7.3 

CM  2659  2 

Colombia 

42.9 

23.0 

19.9 

11.3 

4.7 

8.9 

8.8 

6.9 

USNM  418876  2 

Venezuela 

42.2 

24.0 

20.6 

11.6 

4.8 

8.6 

9.3 

7.5 

TTU  5299  2 

Trinidad 

43.0 

23.8 

20.6 

11.8 

5.2 

8.8 

9.4 

7.5 

TTU  13155(3 

Nicaragua 

39.5 

22.8 

19.4 

11.0 

4.7 

8.5 

8.7 

7.2 

TTU  5410(3 

Trinidad 

42.1 

24.0 

20.2 

11.6 

5.0 

8.9 

9.2 

7.4 

TTU  5449  (3 

Trinidad 

42.3 

23.7 

20.3 

11.3 

4.9 

8.7 

8.9 

7.2 

TTU  101 16  6 

Trinidad 

42.7 

24.3 

20.7 

11.5 

5.0 

8.5 

9.2 

7.3 

Micronycteris  megalotis 

KU  70474  2 

Nicaragua 

38.0 

20.2 

17.4 

9.7 

4.2 

8.0 

7.5 

6.5 

KU  97407  2 

Nicaragua 

33.6 

19.6 

17.2 

9.2 

4.1 

7.7 

7.1 

6.2 

KU  97409  2 

Nicaragua 

36.7 

19.6 

17.1 

9.0 

3.9 

7.7 

7.3 

6.3 

KU  114772  2 

Nicaragua 

34.7 

18.6 

16.3 

9.1 

4.0 

7.4 

7.0 

6.0 

TTU  5438  (3 

Trinidad 

32.6 

18.4 

16.0 

8.9 

4.0 

7.5 

6.9 

6.0 

TTU  5446  (3 

Trinidad 

35.5 

19.1 

16.3 

8.7 

3.9 

7.5 

7.0 

5.9 

TTU  5495  (3 

Trinidad 

32.2 

18.7 

16.2 

8.6 

3.8 

7.4 

6.9 

6.0 

TTU  9788  (3 

Trinidad 

33.3 

18.8 

16.1 

8.8 

4.1 

7.5 

7.1 

5.9 

Micronycteris  minuta 

TTU  5226  2 

Trinidad 

36.5 

18.5 

15.9 

8.6 

4.3 

7.6 

6.6 

5.7 

TTU  5437  2 

Trinidad 

35.6 

18.6 

15.8 

8.6 

4.0 

7.5 

6.5 

5.7 

TTU  5443  2 

Trinidad 

34.6 

18.4 

16.0 

8.4 

4.0 

7.6 

6.5 

5.5 

TTU  5444  2 

Trinidad 

36.5 

18.8 

16.4 

8.5 

4.0 

7.5 

6.7 

5.6 

TTU  5225  (3 

Trinidad 

35.3 

18.8 

16.5 

8.6 

4.2 

7.4 

6.8 

5.6 

TTU  5239  (3 

Trinidad 

35.2 

18.7 

16.5 

8.7 

4.1 

7.6 

6.4 

5.5 

TTU  5294  J 

Trinidad 

34.9 

18.3 

16.0 

8.7 

4.2 

7.5 

6.7 

5.5 

TTU  5295  (3 

Trinidad 

35.5 

19.0 

16.2 

8.3 

4.0 

7.4 

6.7 

5.5 

88 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. —  Continued. 


Micronycteris  nicefori 


TTU  5257  9 

Trinidad 

40.2 

22.0 

19.6 

9.8 

4.5 

8.3 

7.8 

6.3 

TTU  5297  9 

Trinidad 

40.0 

21.4 

19.3 

9.5 

4.1 

8.2 

7.7 

6.0 

TTU  5298  9 

Trinidad 

38.8 

21.2 

19.9 

9.4 

4.2 

7.6 

7.6 

6.3 

TTU  8954  9 

Trinidad 

38.4 

21.1 

19.0 

9.8 

4.4 

8.3 

7.5 

6.3 

TTU  8963  c? 

Trinidad 

36.8 

21.1 

18.9 

9.5 

4.0 

8.0 

7.6 

6.2 

TTU  8964  6 

Trinidad 

37.1 

20.7 

18.2 

9.5 

4.3 

8.3 

7.2 

6.3 

TTU  8965  c? 

Trinidad 

36.3 

20.8 

18.7 

9.6 

4.2 

8.2 

7.6 

6.1 

TTU  8966  c? 

Trinidad 

39.1 

20.8 

18.4 

9.3 

4.0 

8.5 

7.5 

6.3 

Micronycteris  pnsilla 

AMNH  78830  c? 

Brazil 

34.3 

15.2 

8.9 

4.2 

7.6 

6.7 

5.7 

AMNH  78831  c? 

Brazil 

33.7 

17.8 

15.4 

4.5 

7.8 

6.8 

6.0 

Micronycteris  schmidtorum 

USNM  388704  9 

Venezuela 

34.7 

19.5 

16.9 

9.3 

4.2 

7.5 

7.5 

6.1 

USNM  388713  9 

Venezuela 

34.2 

19.8 

17.1 

9.2 

4.4 

7.7 

7.4 

6.0 

USNM  407257  9 

Venezuela 

33.9 

19.4 

17.1 

9.4 

4.2 

7.9 

7.2 

6.1 

USNM  415210  9 

Venezuela 

35.1 

18.9 

17.0 

9.1 

4.1 

7.4 

7.4 

5.9 

AMNH  130715  c? 

Venezuela 

36.9 

20.0 

17.7 

9.7 

4.2 

7.8 

7.8 

6.5 

AMNH  130718  c? 

Venezuela 

36.0 

20.1 

17.4 

9.5 

4.0 

7.6 

7.5 

6.4 

AMNH  130725  c? 

Venezuela 

36.8 

20.2 

17.8 

9.8 

4.1 

7.6 

7.8 

6.7 

USNM  444235  <? 

Venezuela 

37.3 

20.2 

17.7 

9.6 

4.2 

7.6 

7.9 

6.6 

Micronycteris 

sylvestris 

KU  96970  9 

Nayarit 

43.8 

21.2 

18.7 

10.2 

4.9 

8.5 

8.4 

6.7 

KU  23646  9 

Veracruz 

42.1 

20.5 

18.7 

10.4 

4.8 

8.5 

8.4 

7.0 

USNM  396399  9 

Panama 

42.0 

19.8 

10.7 

4.5 

8.7 

7.9 

7.2 

KU  23651  3 

Veracruz 

41.2 

21.0 

18.7 

10.2 

4.9 

8.6 

8.3 

7.0 

KU  23653  £ 

Veracruz 

40.3 

20.7 

18.5 

10.1 

4.9 

8.4 

8.1 

6.9 

KU  29594  £ 

Veracruz 

38.3 

21.0 

18.7 

10.2 

4.9 

8.8 

8.5 

6.9 

BMNH  96.10.1.2  £ 

Costa  Rica 

39.5 

20.1 

17.1 

9.9 

4.5 

8.2 

7.9 

6.8 

Minton  bennettii 

BMNH  3.7.1.153  9 

Brazil 

56.6 

26.1 

22.8 

13.8 

4.6 

9.8 

9.4 

9.1 

BMNH  3.7.1.155  9 

Brazil 

56.0 

25.4 

21.8 

4.6 

9.8 

9.0 

8.6 

USNM  391027  9 

Brazil 

58.4 

26.3 

22.9 

14.1 

4.7 

10.0 

9.4 

9.5 

BMNH  65.618  c? 

Guyana 

53.1 

25.6 

22.0 

13.9 

4.5 

9.6 

9.0 

9.5 

USNM  123393  c? 

Brazil 

53.7 

24.5 

21.8 

4.7 

9.1 

9.1 

9.3 

Mimon  coz 

umelae 

KU  23658  9 

Veracruz 

57.5 

26.7 

23.3 

14.3 

4.8 

10.1 

9.5 

9.7 

KU  32092  9 

Veracruz 

55.1 

26.1 

22.6 

14.2 

4.4 

10.0 

9.5 

9.8 

KU  91548  9 

Yucatan 

59.0 

27.0 

23.0 

13.8 

4.5 

9.6 

9.5 

9.3 

KU  93380  9 

Yucatan 

54.9 

25.2 

22.1 

13.6 

4.4 

9.8 

9.1 

8.8 

KU  19171  c? 

Veracruz 

55.1 

26.5 

22.6 

13.3 

4.5 

9.9 

9.4 

9.4 

KU  23656  c? 

Veracruz 

55.4 

26.0 

22.2 

13.8 

4.6 

9.9 

9.4 

9.7 

KU  91546  c? 

Yucatan 

56.4 

25.6 

22.5 

13.1 

4.4 

9.5 

9.3 

8.8 

TTU  9340  c? 

Yucatan 

56.6 

26.0 

22.7 

13.8 

4.7 

9.8 

9.1 

9.1 

Mimon  crenulutum 

USNM  371497  9 

Venezuela 

48.8 

21.3 

18.5 

11.7 

3.9 

8.7 

7.6 

8.4 

USNM  371503  9 

Venezuela 

50.4 

21.9 

18.9 

12.1 

3.8 

8.8 

7.7 

7.9 

TTU  5340  9 

Trinidad 

45.9 

22.0 

19.2 

12.5 

4.0 

8.4 

7.9 

9.0 

TTU  5374  9 

Trinidad 

48.7 

22.6 

19.5 

12.0 

4.3 

8.6 

7.7 

8.4 

TTU  5264  <3 

Trinidad 

47.0 

21.6 

18.7 

11.6 

3.9 

7.9 

7.5 

8.5 

TTU  5375  <? 

Trinidad 

51.0 

22.4 

19.6 

11.8 

4.2 

8.7 

7.6 

8.6 

TTU  5379  c? 

Trinidad 

47.3 

21.6 

18.5 

11.7 

3.9 

8.4 

7.7 

8.4 

TTU  5460  c? 

Trinidad 

48.2 

21.9 

19.0 

11.7 

4.1 

8.3 

7.7 

8.5 

Mimon  koepckeae 

LSU  1 6447  9 

Peru* 

47.4 

21.9 

18.9 

11.6 

4.2 

8.6 

7.5 

8.2 

LSU  15675  c? 

Peru 

46.3 

21.7 

18.5 

11.4 

4.1 

8.3 

7.1 

8.0 

LSU  15676  c? 

Peru 

49.8 

21.5 

18.9 

11.7 

3.9 

8.2 

7.4 

8.1 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


89 


Appendix  1. — Continued. 


USNM  388843  9 

Venezuela 

Phylloderma  stenops 

71.8  30.9  26.0 

14.9 

8.9 

12.7 

9.7 

9.7 

USNM  388844  9 

Venezuela 

71.6 

32.2 

28.3 

16.0 

9.2 

13.4 

10.0 

10.2 

USNM  388848  9 

Venezuela 

71.2 

30.3 

26.3 

14.8 

9.0 

12.9 

9.7 

9.4 

TTU  5318  9 

Trinidad 

65.8 

30.0 

25.3 

14.6 

8.9 

12.1 

9.6 

9.8 

USNM  335144  <J 

Panama 

73.4 

31.8 

27.3 

15.6 

9.1 

13.0 

10.2 

9.6 

USNM  388842  6 

Venezuela 

72.2 

31.2 

27.4 

14.7 

8.9 

12.7 

9.8 

9.6 

USNM  388845  6 

Venezuela 

69.5 

30.3 

26.5 

14.7 

8.7 

12.7 

9.6 

9.4 

USNM  388846  i 

Venezuela 

73.3 

30.8 

26.5 

16.3 

9.0 

13.1 

10.3 

10.6 

KU  114811  9 

Nicaragua 

Phyllostomus  discolor 

61.7  30.8  27.1 

15.0 

6.5 

12.1 

9.5 

10.3 

KU  114812  9 

Nicaragua 

64.9 

31.8 

26.8 

16.1 

6.6 

12.2 

9.9 

10.5 

KU  1 148 13  9 

Nicaragua 

61.6 

29.5 

26.2 

15.3 

6.2 

11.7 

9.5 

10.0 

TTU  5452  9 

Trinidad 

59.9 

29.7 

25.6 

15.4 

6.2 

12.1 

9.0 

9.6 

KU  110701  6 

Nicaragua 

62.9 

31.0 

27.3 

15.5 

6.6 

11.8 

9.8 

10.0 

KU  1 10702  c? 

N  icaragua 

64.3 

32.1 

28.5 

16.4 

6.4 

12.6 

10.0 

10.4 

KU  1 14800  6 

Nicaragua 

61.8 

31.7 

28.5 

16.1 

6.6 

12.3 

9.8 

10.4 

TTU  5412  6 

Trinidad 

60.9 

30.5 

27.0 

15.4 

6.3 

11.9 

9.6 

9.8 

Phyllostomus  elongalus 


USNM  364304  9 

Peru 

67.5 

30.6 

26.3 

16.1 

5.4 

10.9 

10.6 

11.2 

USNM  364306  9 

Peru 

66.2 

30.0 

25.2 

16.7 

5.4 

11.1 

10.3 

11.4 

USNM  364310  9 

Peru 

64.3 

29.0 

25.2 

16.6 

5.3 

10.9 

10.2 

11.3 

USNM  499015  9 

Peru 

64.6 

29.0 

25.4 

16.3 

5.4 

10.8 

10.0 

11.3 

USNM  483339  6 

Colombia 

60.8 

28.9 

24.7 

15.4 

5.1 

10.5 

10.3 

11.0 

USNM  361515  6 

Brazil 

64.5 

30.5 

25.6 

16.5 

5.3 

10.9 

10.3 

11.5 

USNM  364303  <J 

Peru 

67.2 

30.2 

25.8 

16.9 

5.7 

11.2 

10.4 

1 1.3 

USNM  364305  6 

Peru 

67.7 

29.1 

25.5 

16.5 

5.6 

10.8 

10.1 

11.0 

Phyllostomus  hastatus 


KU  1 10716  9 

Nicaragua 

90.4 

39.3 

33.9 

21.0 

7.3 

14.9 

13.2 

13.5 

KU  110717  9 

Nicaragua 

92.6 

40.7 

34.3 

21.3 

7.1 

14.6 

14.1 

14.2 

KU  110720  9 

Nicaragua 

88.2 

40.8 

34.5 

21.9 

7.3 

15.1 

13.9 

14.4 

CM  2667  9 

Colombia 

84.7 

38.9 

32.5 

20.7 

7.2 

14.2 

13.0 

14.0 

KU  110718d 

Nicaragua 

91.1 

41.5 

35.8 

22.4 

7.7 

15.2 

14.3 

14.3 

KU  110719d 

Nicaragua 

94.4 

43.1 

36.1 

23.2 

7.7 

15.5 

13.8 

14.3 

ROM  31469(3 

Trinidad 

82.8 

37.6 

32.1 

19.9 

6.8 

13.8 

12.8 

12.8 

ROM  50233  d 

Brazil 

86.9 

39.0 

32.5 

21.0 

7.4 

14.1 

12.9 

13.9 

Phyllostomus  latifolius 


BMNH 

1.6.4.44  9 

Guyana 

59.6 

28.0 

23.5 

15.1 

4.9 

10.3 

10.0 

10.3 

BMNH 

1.6.4.45  9 

Guyana 

59.8 

28.4 

24.0 

15.1 

5.2 

10.4 

9.9 

10.7 

BMNH 

1.6.4.40(3 

Guyana 

58.7 

28.3 

24.1 

15.5 

5.1 

10.3 

10.0 

10.5 

BMNH 

1.6.4.41  (3 

Guyana 

59.2 

28.3 

24.4 

4.8 

10.3 

10.4 

10.6 

BMNH 

1.6.4.42(3 

Guyana 

58.9 

28.6 

24.5 

15.8 

5.1 

10.4 

10.3 

11.2 

BMNH 

1.6.4.43(3 

Guyana 

58.5 

28.2 

24.1 

15.8 

5.0 

10.5 

10.0 

10.9 

Tonatia  hidens 


USNM  315218  9 

Panama 

58.8 

28.9 

24.0 

14.0 

5.7 

10.7 

9.7 

8.9 

TTU  5260  9 

Trinidad 

55.8 

27.9 

23.7 

13.7 

5.3 

10.2 

9.6 

8.3 

TTU  9774  9 

Trinidad 

54.8 

28.3 

24.1 

14.3 

5.4 

10.6 

9.3 

8.4 

TTU  9778  9 

Trinidad 

55.2 

28.3 

23.5 

14.0 

5.2 

10.4 

9.6 

8.6 

TTU  13108(3 

Nicaragua 

57.0 

28.9 

24.5 

14.5 

5.7 

10.6 

10.4 

9.4 

TTU  5261  (3 

Trinidad 

55.1 

28.6 

24.2 

14.5 

5.6 

10.6 

9.6 

8.5 

TTU  5338  (3 

Trinidad 

54.9 

27.6 

23.1 

13.9 

5.1 

10.7 

9.4 

8.4 

TTU  5339(3 

Trinidad 

51.5 

27.3 

23.1 

13.8 

5.6 

10.6 

9.5 

8.6 

Tonatia  hrasiliense 


AMNH  95497  9 

Peru 

35.7 

20.0 

16.6 

9.4 

3.3 

8.1 

6.8 

6.5 

AMNH  95498  6 

Brazil 

35.8 

20.0 

17.0 

9.5 

3.2 

7.9 

6.8 

6.3 

LSU  16440(3 

Peru 

37.9 

20.8 

17.0 

9.4 

3.2 

8.3 

6.8 

6.1 

90 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


Tonatia  carrikeri 


AMNH  30180  9 

Venezuela 

46.0 

25.2 

20.3 

11.5 

3.6 

9.4 

8.0 

7.8 

AMNH  30183  9 

Venezuela 

46.8 

24.8 

20.0 

10.8 

3.6 

9.3 

8.3 

7.5 

AMNH  209322  9 

Bolivia 

45.6 

24.5 

20.2 

11.1 

3.6 

9.4 

8.1 

7.7 

AMNH  30181  6 

Venezuela 

48.4 

25.8 

21.5 

12.2 

3.9 

9.7 

8.6 

8.0 

ROM  67468  6 

Guyana 

43.9 

23.9 

19.5 

11.5 

3.7 

9.4 

7.8 

7.3 

Tonatia  minutu 


USNM  314221  9 

Panama 

33.3 

18.9 

15.8 

8.8 

2.9 

7.6 

6.7 

5.8 

USNM  362457  9 

Panama 

34.0 

19.2 

16.3 

9.0 

2.9 

7.8 

6.8 

6.2 

USNM  362458  9 

Panama 

35.0 

19.2 

16.0 

9.2 

3.0 

7.6 

6.7 

6.1 

TTU  5238  9 

Trinidad 

35.8 

20.1 

16.9 

9.6 

2.9 

8.0 

6.9 

6.2 

TTU  5222  c? 

Trinidad 

36.3 

20.2 

16.8 

9.6 

3.2 

8.4 

7.0 

6.1 

TTU  5309  <? 

Trinidad 

34.5 

20.2 

16.8 

9.6 

3.1 

8.2 

6.7 

6.4 

TTU  5422  c? 

Trinidad 

35.5 

20.6 

17.6 

10.0 

3.4 

8.5 

7.0 

6.7 

TTU  10119c? 

Trinidad 

35.2 

20.8 

17.3 

10.0 

3.3 

8.4 

6.9 

6.4 

Tonatia  silvicola 


USNM  306549  9 

Panama 

51.6 

27.0 

22.8 

12.9 

3.9 

10.5 

8.9 

8.1 

USNM  309357  9 

Panama 

50.0 

26.4 

22.2 

12.7 

3.9 

10.1 

9.5 

8.7 

USNM  323068  9 

Panama 

53.3 

26.7 

22.6 

12.9 

3.9 

10.2 

9.0 

8.3 

USNM  364278  9 

Peru 

55.0 

28.7 

23.6 

13.1 

4.0 

10.4 

9.8 

9.0 

USNM  323074  c? 

Panama 

54.7 

27.9 

23.1 

13.5 

4.1 

10.6 

9.3 

8.8 

USNM  323076c? 

Panama 

53.5 

27.8 

23.3 

13.3 

4.1 

10.5 

9.2 

8.7 

USNM  407291  6 

Venezuela 

51.4 

28.3 

23.6 

13.7 

4.3 

11.1 

9.7 

8.6 

USNM  364275  c ? 

Peru 

55.2 

30.4 

24.8 

14.1 

4.1 

10.8 

10.4 

9.6 

Tonatia  venezuelae 


USNM  102919  9 

Venezuela 

39.8 

21.5 

17.9 

10.5 

3.1 

8.3 

7.5 

6.9 

USNM  142567  9 

Venezuela 

38.9 

21.7 

17.9 

10.0 

3.2 

8.3 

7.4 

6.7 

BMNH  11.5.25.41  c? 

Venezuela 

39.1 

21.5 

17.7 

10.6 

3.4 

8.6 

7.4 

7.0 

Trachops  cirrhosus 


KU  93381  9 

Campeche 

57.9 

27.8 

24.1 

13.6 

5.0 

11.1 

9.7 

9.7 

TTU  13172  9 

Costa  Rica 

60.1 

28.0 

24.2 

13.8 

5.3 

11.5 

10.3 

9.7 

TTU  9777  9 

Trinidad 

60.1 

29.0 

25.4 

14.8 

5.3 

11.9 

10.8 

10.4 

TTU  9780  9 

Trinidad 

61.2 

29.6 

25.7 

14.9 

5.2 

11.7 

11.4 

10.5 

TTU  6077  c? 

Oaxaca 

59.3 

28.2 

24.5 

13.8 

5.0 

11.5 

10.0 

9.7 

TTU  6115  c? 

Chiapas 

59.5 

28.2 

24.5 

13.5 

4.9 

11.1 

10.1 

9.8 

KU  114818c? 

Nicaragua 

57.3 

27.6 

24.2 

13.9 

4.8 

11.4 

9.8 

9.4 

TTU  9779  c? 

Trinidad 

60.7 

30.4 

26.4 

15.4 

5.5 

12.1 

11.1 

10.8 

Vampyrum  spectrum 


USNM  335161  9 

Panama 

106.0 

51.9 

43.1 

24.5 

8.5 

15.9 

20.2 

14.8 

USNM  335162  9 

Panama 

107.1 

53.6 

43.7 

25.4 

7.9 

15.6 

19.8 

15.0 

TTU  5357  9 

Trinidad 

102.0 

51.2 

42.9 

23.3 

8.0 

15.8 

20.9 

14.5 

TTU  9837  9 

Trinidad 

103.3 

51.6 

44.0 

23.4 

7.7 

15.7 

21.1 

15.4 

AMNH  28993  c? 

Nicaragua 

105.7 

50.4 

42.3 

24.5 

8.0 

15.6 

19.7 

14.4 

KU  88190  c? 

Costa  Rica 

110.4 

50.7 

43.0 

23.4 

8.1 

15.8 

19.9 

14.3 

TTU  9836  c? 

Trinidad 

106.1 

52.4 

44.1 

24.2 

7.8 

15.8 

21.1 

15.2 

TTU  11439  c? 

Trinidad 

107.1 

52.0 

43.2 

25.2 

8.4 

16.4 

20.6 

15.4 

Glossophaginae 

Anoura  brevirostrum 


AMNH  214324  9 

Peru 

39.8 

23.5 

22.5 

9.6 

5.0 

9.4 

8.3 

5.7 

AMNH  233263  9 

Peru 

38.9 

23.3 

22.3 

9.4 

4.9 

9.1 

8.0 

5.4 

TCWC  11881  9 

Peru 

38.0 

23.1 

22.3 

9.4 

4.8 

9.2 

7.7 

5.3 

TCWC  11882  9 

Peru 

40.0 

23.1 

22.0 

9.5 

4.6 

9.2 

8.2 

5.6 

LSU  17941  c? 

Peru 

40.2 

23.3 

22.6 

10.3 

5.0 

9.3 

8.0 

5.7 

TCWC  11880  c? 

Peru 

39.6 

23.3 

22.5 

10.0 

4.8 

9.3 

8.1 

5.4 

Anoura  caudifer 

USNM  373705  9 

Venezuela 

38.5 

24.6 

23.8 

9.5 

4.6 

9.3 

9.1 

5.5 

USNM  373761  9 

Venezuela 

36.0 

22.0 

21.3 

8.9 

4.3 

8.6 

8.0 

5.3 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


91 


Appendix  1. — Continued. 


USNM  389076  9 

Venezuela 

35.1 

22.1 

21.4 

9.1 

4.6 

8.9 

8.2 

5.6 

USNM  389108  9 

Venezuela 

36.4 

23.0 

22.3 

9.2 

4.6 

8.9 

8.3 

5.3 

USNM  370109  6 

Venezuela 

37.2 

24.0 

23.2 

9.4 

4.6 

8.8 

8.8 

5.3 

USNM  373704  3 

Venezuela 

37.4 

24.3 

23.4 

10.1 

4.6 

9.2 

8.9 

5.8 

USNM  385771  <? 

Venezuela 

36.8 

22.0 

21.5 

9.3 

4.5 

9.0 

8.0 

5.3 

USNM  385773  c? 

Venezuela 

37.2 

22.0 

21.3 

9.3 

4.4 

9.0 

8.0 

5.4 

Anoura  cull  rata 

USNM  309400  9 

Panama 

43.8 

26.4 

25.5 

10.9 

5.0 

10.0 

9.3 

6.0 

USNM  319249  9 

Panama 

41.7 

26.3 

25.5 

10.3 

5.1 

10.0 

9.2 

6.0 

USNM  419465  9 

Venezuela 

41.4 

25.4 

24.6 

10.6 

5.0 

9.8 

8.9 

6.2 

USNM  419466  9 

Venezuela 

41.1 

25.6 

24.5 

10.4 

5.0 

10.0 

8.7 

6.2 

USNM  309396  c? 

Panama 

43.0 

26.4 

25.6 

10.7 

5.3 

10.3 

9.1 

5.7 

USNM  309397  c? 

Panama 

44.3 

26.6 

25.8 

11.0 

5.3 

10.0 

9.4 

6.1 

USNM  309398  c? 

Panama 

43.6 

26.7 

26.0 

11.1 

5.3 

10.0 

9.3 

6.1 

USNM  337991  c? 

Panama 

42.3 

26.2 

25.4 

11.0 

5.3 

10.0 

9.3 

6.3 

Anoura  geoffroyi 

USNM  362594  9 

Panama 

43.7 

26.3 

25.7 

11.0 

4.9 

9.8 

10.1 

6.3 

USNM  385802  9 

Venezuela 

42.7 

25.0 

24.1 

10.7 

4.9 

9.7 

9.5 

6.2 

TTU  5825  9 

Trinidad 

42.7 

25.0 

24.2 

10.8 

4.8 

9.7 

9.5 

6.3 

TTU  8977  9 

Trinidad 

42.0 

24.8 

24.1 

10.6 

5.1 

9.8 

9.3 

6.2 

USNM  385852  c? 

Venezuela 

42.0 

25.3 

25.1 

10.8 

4.8 

9.7 

9.5 

6.0 

TTU  5370  c? 

Trinidad 

41.0 

24.9 

24.1 

10.8 

5.1 

9.6 

9.2 

6.1 

TTU  5823  c? 

Trinidad 

43.0 

24.7 

24.2 

11.3 

5.1 

9.8 

9.2 

6.3 

TTU  5826  c? 

Trinidad 

40.5 

24.5 

24.0 

11.0 

4.9 

9.8 

9.0 

6.3 

Anoura  werckleae 

LACM  25438  9 

Costa  Rica 

43.1 

26.1 

25.3 

10.5 

5.2 

10.1 

9.3 

6.0 

LACM  15186  c? 

Costa  Rica** 

40.7 

25.8 

25.1 

10.8 

5.3 

10.2 

9.0 

6.1 

Choeroniscus  godmani 

KU  90650  9 

Sinaloa 

33.8 

20.7 

20.0 

3.2 

8.2 

7.3 

4.2 

AMNH  186162  9 

Oaxaca 

35.1 

20.8 

20.1 

3.2 

8.4 

7.3 

4.2 

USNM  337550  9 

Nicaragua 

34.4 

21.2 

20.6 

3.3 

8.1 

7.7 

4.3 

USNM  337551  9 

Nicaragua 

33.8 

20.6 

20.4 

3.5 

8.0 

7.6 

4.2 

KU  102370  c? 

Chiapas 

33.4 

19.7 

18.8 

2.9 

7.9 

6.9 

4.2 

AMNH  172778  c? 

Oaxaca 

32.6 

19.3 

18.8 

3.0 

8.3 

6.7 

4.0 

AMNH  172779  c? 

Oaxaca 

33.1 

18.9 

18.2 

2.9 

8.0 

6.7 

3.9 

AMNH  208869  c? 

Oaxaca 

32.3 

19.2 

18.5 

2.9 

8.3 

6.4 

4.0 

Choeroniscus  inca 

AMNH  140471  9 

Guyana 

37.3 

24.5 

24.1 

3.8 

8.5 

8.3 

4.7 

BMNH  12.9.5.2  9 

Peru 

33.1 

3.8 

8.5 

7.6 

4.5 

Choeroniscus  intermedins 

TTU  5319  9 

Trinidad 

34.2 

23.1 

22.8 

3.8 

8.5 

7.8 

4.6 

TTU  5496  9 

Trinidad 

34.9 

23.2 

22.8 

3.5 

8.3 

8.0 

4.5 

TTU  9006  9 

Trinidad 

34.8 

22.6 

22.5 

3.5 

8.4 

8.1 

4.4 

TTU  9007  9 

Trinidad 

36.0 

23.6 

23.0 

4.0 

8.7 

8.1 

4.6 

TTU  8994  c? 

Trinidad 

34.1 

22.8 

21.8 

3.3 

8.8 

7.6 

4.3 

TTU  8995  c ? 

Trinidad 

35.0 

21.7 

21.3 

3.2 

8.4 

7.1 

4.4 

TTU  8998  c? 

Trinidad 

35.4 

21.2 

20.7 

3.2 

8.2 

7.5 

4.2 

TTU  8999  c? 

Trinidad 

35.7 

22.4 

21.9 

3.6 

8.2 

7.8 

4.7 

Choeroniscus 

minor 

AMNH  69152  9 

Guyana 

36.0 

22.7 

21.6 

3.7 

8.2 

7.7 

4.2 

USMN  361573  9 

Brazil 

33.7 

23.2 

22.5 

3.6 

8.2 

8.6 

4.6 

USNM  361574  9 

Brazil 

35.7 

22.7 

22.2 

3.4 

8.3 

8.3 

4.4 

USMN  460100  9 

Brazil 

35.7 

23.6 

22.8 

3.6 

8.7 

8.2 

4.5 

Choeroniscus  periosus 

AMNH  217038  9 

Colombia 

40.4 

30.0 

29.2 

4.8 

9.3 

10.5 

5.0 

USNM  344918  9 

Colombia 

41.2 

30.2 

29.5 

4.9 

9.8 

10.9 

5.3 

92 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


Choeronycteris  mexicana 


TTU  6288  9 

Sonora 

45.8 

30.8 

29.8 

4.0 

9.9 

11.4 

5.6 

TTU  6360  9 

Sonora 

46.3 

29.8 

28.6 

3.9 

10.0 

11.0 

5.9 

TTU  6447  9 

Sonora 

42.4 

29.7 

28.8 

4.2 

10.0 

11.5 

5.6 

TTU  10122  9 

Tamaulipas 

45.9 

29.4 

28.5 

4.0 

9.6 

11.0 

5.7 

KU  31863  S 

Jalisco 

45.6 

30.3 

29.0 

4.1 

9.6 

11.5 

5.6 

KU  38250  <3 

Jalisco 

45.3 

29.5 

28.6 

4.0 

9.4 

11.3 

5.3 

KU  107192d 

Jalisco 

43.0 

29.4 

28.5 

3.7 

9.4 

11.0 

5.2 

KU  107194c3 

Jalisco 

43.6 

30.1 

28.9 

3.8 

9.7 

11.6 

5.5 

Glossophaga  alticola 


KU  70624  9 

Nicaragua 

38.3 

21.0 

19.4 

9.6 

4.5 

8.9 

7.0 

5.6 

KU  70628  9 

Nicaragua 

38.1 

20.4 

18.8 

9.7 

4.5 

8.9 

7.0 

5.7 

KU  105966  9 

Nicaragua 

37.3 

21.2 

19.6 

9.8 

4.6 

8.8 

7.1 

5.4 

KU  114819  9 

Nicaragua 

36.8 

20.9 

19.3 

9.5 

4.4 

8.6 

7.1 

5.6 

KU  105964(3 

Nicaragua 

34.0 

20.0 

18.7 

9.4 

4.3 

8.5 

6.8 

5.4 

KU  105967(5 

Nicaragua 

35.8 

20.1 

18.4 

9.4 

4.6 

8.9 

6.9 

5.3 

KU  114820(3 

Nicaragua 

36.6 

19.8 

18.3 

9.3 

4.4 

8.7 

6.7 

5.2 

KU  114822(3 

Nicaragua 

36.3 

20.7 

19.0 

9.8 

4.5 

9.0 

7.1 

5.8 

Glossophaga  commissarisi 


KU  105972  9 

Nicaragua 

32.4 

19.8 

18.4 

9.3 

4.4 

8.3 

6.8 

5.3 

KU  105975  9 

Nicaragua 

32.7 

20.2 

18.8 

9.6 

4.5 

8.2 

6.9 

5.5 

KU  110770  9 

Nicaragua 

33.3 

20.3 

18.8 

9.6 

4.5 

8.4 

6.9 

5.4 

KU  110775  9 

Nicaragua 

34.5 

20.4 

19.0 

9.3 

4.3 

8.1 

7.1 

5.4 

KU  110730(3 

Nicaragua 

33.9 

20.8 

19.3 

9.8 

4.7 

8.4 

7.0 

5.6 

KU  110733  (3 

Nicaragua 

31.1 

20.6 

18.8 

9.9 

4.7 

8.9 

6.9 

5.3 

KU  110734(3 

Nicaragua 

34.6 

20.3 

18.8 

9.9 

4.5 

8.4 

6.7 

5.5 

KU  110767(3 

Nicaragua 

35.6 

20.7 

19.1 

9.4 

4.4 

8.5 

6.7 

5.1 

Glossophaga  longirostris 


TTU  9338  9 

Grenada 

38.6 

23.1 

21.5 

9.4 

4.7 

8.6 

7.9 

5.8 

KU  118105  9 

Venezuela 

38.0 

22.8 

21.4 

10.1 

4.4 

8.8 

8.0 

5.5 

KU  118117  9 

Venezuela 

38.6 

23.3 

21.6 

10.1 

4.6 

8.8 

8.1 

5.9 

KU  118123  9 

Venezuela 

39.5 

23.3 

22.0 

9.9 

4.5 

8.8 

8.2 

6.0 

KU  110073(3 

Grenada 

37.5 

23.1 

21.5 

10.2 

4.5 

8.6 

7.9 

5.7 

KU  118114(3 

Venezuela 

36.4 

22.2 

21.1 

9.8 

4.5 

8.8 

7.6 

5.9 

KU  118115  <3 

Venezuela 

37.6 

23.0 

21.2 

9.8 

4.4 

8.8 

7.7 

5.8 

KU  118116(3 

Venezuela 

36.3 

22.8 

21.4 

10.1 

4.7 

9.4 

8.0 

5.8 

Glossophaga  soricina 


KU  106015  9 

Nicaragua 

36.5 

21.0 

19.7 

9.1 

4.6 

8.2 

7.3 

5.4 

KU  106018  9 

Nicaragua 

36.7 

21.4 

19.9 

9.3 

4.5 

8.6 

6.9 

5.2 

KU  106019  9 

Nicaragua 

36.5 

21.5 

19.9 

9.6 

4.6 

8.5 

7.2 

5.3 

KU  106020  9 

Nicaragua 

36.0 

21.9 

20.6 

10.0 

4.9 

8.9 

7.7 

5.7 

KU  106008(3 

Nicaragua 

36.7 

21.4 

19.7 

9.7 

4.6 

8.8 

7.0 

5.5 

KU  106016(3 

Nicaragua 

35.0 

20.9 

19.2 

9.2 

4.7 

8.5 

7.0 

5.3 

KU  106021  (3 

Nicaragua 

34.5 

21.1 

19.3 

9.4 

4.5 

8.4 

7.0 

5.3 

KU  106022  (3 

Nicaragua 

36.8 

21.7 

20.1 

9.7 

4.5 

8.6 

7.3 

5.4 

Hylonycleris  underwood i 


KU  108603  9 

Jalisco 

36.3 

20.6 

20.0 

4.0 

8.1 

7.2 

4.2 

KU  108605  9 

Jalisco 

33.0 

20.6 

20.0 

3.9 

8.1 

7.0 

4.2 

KU  98140  9 

Oaxaca 

33.9 

23.0 

22.0 

4.5 

8.8 

8.2 

4.9 

TTU  13142  9 

Costa  Rica 

32.2 

21.9 

21.2 

4.0 

8.1 

7.6 

4.5 

KU  108604(3 

Jalisco 

31.6 

20.0 

19.0 

4.1 

8.1 

6.7 

4.0 

KU  108606(3 

Jalisco 

32.5 

20.3 

19.5 

3.8 

8.0 

7.0 

4.2 

KU  23709  <3 

Veracruz 

33.1 

21.6 

20.8 

4.1 

8.6 

7.4 

4.6 

KU  98139(3 

Oaxaca 

33.5 

21.5 

20.8 

4.2 

8.2 

7.5 

4.6 

Leplonycteris  curasoae 


USNM  444799  9 

Venezuela 

53.7 

28.1 

26.8 

11.2 

5.2 

9.8 

9.6 

7.0 

USNM  444800  9 

Venezuela 

53.2 

27.9 

26.5 

10.9 

5.0 

10.2 

9.4 

7.2 

USNM  444802  9 

Venezuela 

54.4 

27.5 

26.7 

11.0 

4.7 

9.6 

9.3 

7.3 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


93 


Appendix  1. — Continued. 


USNM  444803  9 

Venezuela 

54.0 

27.8 

26.9 

10.8 

4.8 

9.9 

9.5 

6.9 

USNM  444734  6 

Venezuela 

50.6 

21 A 

26.1 

11.3 

4.9 

10.0 

9.4 

7.2 

USNM  444736  <J 

Venezuela 

52.6 

27.1 

26.1 

11.2 

4.7 

10.1 

9.1 

7.0 

USNM  444739(3 

Venezuela 

53.3 

27.4 

26.5 

11.1 

5.1 

9.9 

9.1 

7.0 

USNM  444740cJ 

Venezuela 

53.8 

27.9 

26.6 

11.2 

5.2 

10.3 

9.5 

7.1 

Leptonycteris  nivalis 

TTU  6565  9 

Texas 

58.2 

27.8 

27.1 

11.5 

5.3 

11.0 

9.6 

7.1 

KU  33068  9 

Coahuila 

52.0 

28.9 

27.5 

11.2 

5.2 

11.0 

9.6 

7.0 

KU  33070  9 

Coahuila 

50.6 

27.5 

26.8 

11.3 

5.5 

10.7 

9.2 

7.1 

KU  33071  9 

Coahuila 

52.9 

29.1 

27.5 

11.4 

5.6 

11.0 

9.4 

6.7 

TTU  9208  (3 

Texas 

56.7 

27.7 

26.8 

11.0 

5.5 

10.6 

9.0 

6.7 

KU  98378(3 

Nuevo  Leon 

56.3 

28.1 

27.1 

11.4 

5.3 

10.7 

9.0 

6.6 

KU  98379d 

Nuevo  Leon 

54.8 

28.4 

26.8 

11.4 

5.5 

10.9 

9.0 

7.0 

KU  98413  d 

Nuevo  Leon 

56.8 

27.5 

26.3 

11.0 

4.9 

10.5 

8.9 

7.0 

Leptonycteris 

sanborni 

TTU  6564  9 

Sonora 

53.4 

27.1 

25.9 

10.6 

4.8 

10.0 

8.9 

6.9 

TTU  10603  9 

Sonora 

54.8 

27.5 

26.6 

10.6 

4.8 

9.9 

9.1 

6.8 

TTU  10604  9 

Sonora 

50.9 

26.7 

25.6 

10.4 

4.7 

9.8 

9.0 

6.6 

TTU  10605  9 

Sonora 

50.0 

26.1 

25.5 

10.3 

4.6 

9.8 

8.4 

6.5 

KU  33349(3 

Jalisco 

51.3 

25.9 

25.0 

10.7 

4.4 

9.5 

8.3 

6.1 

KU  34148(3 

Jalisco 

53.1 

26.4 

25.3 

10.6 

4.3 

9.6 

9.0 

6.6 

KU  34149(3 

Jalisco 

51.6 

26.4 

25.8 

11.0 

4.7 

9.9 

8.7 

6.5 

KU  34222(3 

Jalisco 

51.8 

27.1 

26.0 

10.8 

5.0 

9.9 

9.0 

6.6 

Lichonycteris 

degener 

AMNH  95118  9 

Brazil 

18.4 

17.9 

4.3 

8.4 

6.0 

4.4 

AMNH  95485  9 

Brazil 

32.4 

USNM  239520  9 

Brazil 

18.8 

18.2 

3.8 

7.9 

6.0 

4.2 

Lichonycteris  ohscura 

TTU  13124  9 

Nicaragua 

31.7 

19.2 

18.0 

4.1 

8.0 

6.2 

4.4 

TTU  13125  9 

Nicaragua 

32.2 

18.4 

17.6 

4.0 

8.1 

5.9 

4.4 

TTU  13126  9 

Nicaragua 

32.6 

18.8 

18.2 

4.0 

7.7 

6.3 

4.4 

TTU  13128  9 

Nicaragua 

33.0 

18.8 

17.9 

4.2 

7.9 

6.0 

4.4 

KU  110785  (3 

Nicaragua 

30.7 

18.2 

17.0 

3.9 

7.9 

5.7 

4.3 

TTU  13117(3 

Nicaragua 

30.3 

18.0 

16.8 

3.9 

8.1 

5.5 

4.2 

TTU  13127(3 

Nicaragua 

32.1 

18.5 

17.3 

4.0 

8.2 

5.8 

4.1 

TTU  18967(3 

Nicaragua 

31.9 

17.9 

16.9 

3.9 

7.9 

5.5 

4.5 

Lionycteris  spurrelli 

USNM  385702  9 

Venezuela 

37.1 

19.5 

17.7 

3.8 

7.9 

5.9 

5.0 

USNM  385704  9 

Venezuela 

36.8 

20.3 

18.8 

4.2 

7.9 

6.4 

5.3 

USNM  385705  9 

Venezuela 

35.3 

20.7 

19.0 

4.0 

8.1 

6.3 

5.1 

USNM  385706  9 

Venezuela 

34.8 

19.5 

17.5 

4.1 

7.5 

6.2 

5.1 

BMNH  13.8.10.1  (3 

Colombia 

32.5 

18.9 

17.1 

3.7 

8.0 

5.9 

4.6 

USNM  385698(3 

Venezuela 

33.4 

19.3 

17.8 

4.0 

8.2 

6.0 

4.9 

USNM  385699(3 

Venezuela 

35.2 

19.5 

18.0 

4.0 

7.9 

6.0 

4.7 

USNM  239477  6 

Brazil 

35.2 

19.5 

18.0 

4.2 

8.0 

6.0 

5.1 

Lonchophylla 

concava 

TCWC  9826  9 

Costa  Rica 

33.7 

23.0 

21.5 

4.4 

8.7 

7.4 

5.1 

TCWC  9827  9 

Costa  Rica 

33.7 

22.8 

21.5 

4.6 

9.1 

7.6 

5.3 

TCWC  22528  9 

Costa  Rica 

33.5 

22.5 

20.9 

4.3 

8.8 

7.7 

5.0 

USNM  309389  9 

Panama 

34.4 

23.6 

22.0 

4.5 

8.9 

7.9 

5.4 

TCWC  9828  (3 

Costa  Rica 

34.0 

22.8 

21.5 

4.4 

8.9 

7.4 

5.2 

TCWC  22526  c3 

Costa  Rica 

34.4 

23.3 

21.6 

4.4 

8.8 

7.6 

5.2 

TCWC  22527  6 

Costa  Rica 

33.7 

23.1 

21.7 

4.3 

8.8 

7.5 

5.0 

USNM  179621  6 

Panama 

33.5 

23.5 

22.1 

4.5 

9.0 

7.9 

5.5 

Lonchophylla 

hesperia 

TCWC  11899  9 

Peru 

38.4 

27.4 

26.1 

4.6 

9.2 

8.9 

5.6 

TCWC  23274  9 

Peru 

38.7 

26.0 

24.5 

4.8 

9.0 

8.3 

5.4 

USNM  283177(3 

Peru 

36.0 

25.5 

24.5 

4.8 

9.1 

8.6 

5.8 

94 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


Lonchophylla  mordax 


BMNH  3.9.5.32  3 

Brazil 

34.6 

23.1 

21.5 

4.3 

8.3 

7.7 

5.1 

BMNH  3.9.5.33  3 

Brazil 

34.6 

23.7 

22.2 

4.3 

8.5 

8.3 

5.3 

BMNH  3.9.5.34  <3 

Brazil 

34.3 

23.8 

21.7 

4.3 

9.1 

8.0 

5.3 

USNM  283008  <3 

Brazil 

33.7 

22.7 

20.4 

4.0 

8.2 

7.6 

4.8 

Lonchophylla  rohiislu 


TCWC  18945  9 

Nicaragua 

41.8 

26.4 

24.9 

5.4 

10.2 

9.7 

6.5 

USNM  305237  9 

Panama 

42.4 

26.9 

25.1 

5.4 

10.5 

9.7 

6.9 

USNM  483361  9 

Colombia 

44.3 

26.9 

24.8 

5.1 

10.1 

9.4 

7.0 

TCWC  1  1879  9 

Peru 

45.0 

27.4 

25.6 

5.1 

10.4 

9.9 

6.3 

TCWC  18944c3 

Nicaragua 

41.0 

26.5 

24.8 

5.4 

10.2 

10.0 

6.7 

TTU  13137(3 

Costa  Rica 

43.0 

27.4 

25.8 

5.2 

10.3 

9.8 

6.5 

TTU  13138c3 

Costa  Rica 

45.1 

27.1 

25.4 

5.3 

10.3 

9.8 

7.0 

AMNH  230214(3 

Peru 

45.2 

27.0 

25.9 

5.0 

9.8 

10.1 

6.4 

Lonchophylla  thomasi 


USNM  335180  9 

Panama 

32.0 

21.7 

20.3 

4.1 

8.0 

7.0 

5.1 

USNM  483363  9 

Colombia 

31.4 

21.3 

19.8 

4.2 

8.3 

6.7 

5.3 

ROM  33112  9 

Guyana 

32.4 

21.2 

19.4 

4.2 

8.3 

6.7 

5.2 

AMNH  210688  9 

Bolivia 

31.8 

21.8 

20.2 

4.2 

8.0 

6.8 

5.2 

USNM  483359(3 

Colombia 

31.0 

21.7 

19.7 

4.2 

8.6 

6.9 

5.4 

AMNH  16120(3 

Venezuela 

31.2 

20.8 

19.1 

4.2 

8.5 

6.5 

5.1 

ROM  31607(3 

Guyana 

31.9 

20.2 

18.7 

4.2 

8.3 

6.2 

5.0 

ROM  33986(3 

Guyana 

33.2 

20.4 

18.9 

4.2 

8.3 

6.4 

5.0 

Monophyllus  plelhodon 


TTU  20798  9 

Guadeloupe 

41.2 

23.5 

22.0 

10.0 

4.5 

9.5 

7.9 

5.4 

TTU  20799  9 

Guadeloupe 

41.7 

23.5 

21.6 

10.0 

4.6 

9.5 

8.2 

5.6 

KU  104771  9 

Dominica 

40.2 

22.8 

21.2 

9.6 

4.4 

9.2 

7.8 

5.2 

KU  110088  9 

St.  Vincent 

41.4 

23.0 

21.5 

9.4 

4.6 

9.3 

8.0 

5.4 

TTU  20795  3 

Guadeloupe 

40.1 

23.5 

21.4 

10.3 

4.5 

9.5 

7.8 

5.5 

TTU  20796(3 

Guadeloupe 

42.8 

23.7 

21.9 

10.4 

4.8 

9.6 

8.0 

5.5 

TTU  20800  d 

Guadeloupe 

41.8 

23.3 

21.7 

10.2 

4.6 

9.3 

7.9 

5.6 

TTU  9337(3 

Dominica 

40.9 

23.3 

21.5 

10.4 

4.5 

9.7 

7.7 

5.6 

Monophyllus  redmani 


TTU  22544  9 

Haiti 

39.6 

22.0 

20.7 

8.8 

4.2 

8.8 

7.9 

5.1 

TTU  22545  9 

Haiti 

40.0 

21.7 

20.2 

9.1 

4.3 

9.0 

7.8 

5.0 

TTU  22546  9 

Haiti 

39.6 

21.3 

19.8 

9.0 

4.3 

8.9 

7.8 

5.0 

TTU  22547  9 

Haiti 

39.6 

21.5 

20.0 

9.1 

4.2 

9.1 

7.8 

4.8 

TTU  22537(3 

Haiti 

39.8 

21.2 

20.0 

9.2 

3.9 

8.7 

7.8 

4.9 

TTU  22548(3 

Haiti 

41.4 

21.8 

20.4 

9.2 

4.1 

9.1 

7.9 

5.0 

TTU  22549  (3 

Haiti 

40.8 

22.0 

20.6 

9.4 

4.2 

9.2 

7.8 

4.9 

TTU  22552  (3 

Haiti 

41.0 

22.3 

20.7 

9.3 

4.3 

9.0 

7.8 

5.1 

Musonycteris  harrisoni 


LACM  1  1487  9 

Colima 

41.8 

32.0 

30.8 

4.4 

9.2 

12.5 

4.9 

LACM  11488  9 

Colima 

41.5 

31.7 

30.5 

4.6 

9.2 

11.6 

4.7 

USNM  314689  9 

Colima 

42.7 

32.2 

31.0 

4.0 

9.0 

12.2 

4.8 

USNM  324971  9 

Colima 

42.4 

31.5 

30.5 

4.2 

9.0 

11.7 

4.8 

AMNH  235179(3 

Colima 

42.4 

34.4 

32.9 

4.0 

9.1 

12.3 

4.5 

BMNH61. 1612(3 

Colima 

42.3 

34.4 

33.1 

4.4 

9.0 

13.2 

4.9 

KU  98874  3 

Colima 

40.8 

34.5 

33.3 

4.1 

9.1 

13.6 

5.0 

TTU  9307  6 

Colima 

42.2 

33.3 

32.2 

4.0 

8.2 

12.8 

4.8 

Plalalina  genovensium 

USNM  268765  9 

Peru 

32.7 

30.2 

5.1 

10.3 

5.5 

BMNH  27.11.19.38(3 

Peru 

46.1 

32.7 

30.3 

4.9 

10.3 

10.7 

5.3 

FMNH  24336  6 

Peru 

48.5 

31.1 

29.4 

4.6 

9.6 

10.2 

5.5 

MCZ  34843  3 

Peru 

49.6 

31.9 

29.9 

4.7 

9.5 

10.7 

5.8 

MCZ  32948  3 

Peru 

50.0 

32.6 

30.0 

4.8 

9.5 

11.2 

5.7 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


95 


Appendix  1. — Continued. 


Scleronycleris  ega 


BMNH  7.1.1.671  9 

Brazil 

34.7 

4.3 

8.7 

7.5 

4.8 

USNM  407889  c} 

Venezuela 

35.0 

22.0 

21.2 

4.5 

8.8 

7.7 

5.0 

Carolliinae 

Carollia  brevicauda 

KU  110866  9 

Nicaragua 

38.9 

22.0 

19.4 

5.2 

9.2 

7.0 

7.6 

KU  1 10870  9 

Nicaragua 

41.9 

22.3 

19.7 

5.5 

9.4 

7.0 

8.0 

KU  1 10875  9 

Nicaragua 

39.8 

22.5 

19.5 

5.1 

9.1 

6.7 

7.2 

KU  110878  9 

Nicaragua 

39.7 

22.6 

19.6 

5.5 

9.3 

7.2 

7.7 

KU  1  10873  c} 

Nicaragua 

39.9 

22.5 

19.9 

5.6 

9.6 

6.7 

7.6 

KU  110874  c} 

Nicaragua 

41.3 

23.4 

20.4 

5.2 

9.5 

7.7 

8.1 

KU  110876  c} 

Nicaragua 

39.0 

22.7 

20.2 

5.7 

9.5 

7.1 

8.1 

KU  110877  c} 

Nicaragua 

38.6 

21.6 

18.9 

5.2 

9.5 

6.8 

7.5 

Carollia  custanea 

KU  110890  9 

Nicaragua 

36.5 

19.0 

17.0 

5.2 

8.5 

6.0 

6.9 

KU  114871  9 

Nicaragua 

35.8 

19.4 

17.1 

5.2 

8.6 

6.3 

7.1 

KU  114873  9 

Nicaragua 

35.8 

19.4 

17.0 

5.2 

8.9 

6.3 

6.9 

KU  114880  9 

Nicaragua 

35.2 

19.5 

17.0 

5.1 

9.0 

6.1 

6.7 

KU  110889  c} 

Nicaragua 

35.2 

19.4 

17.0 

5.2 

8.9 

6.0 

6.5 

KU  110892  c} 

Nicaragua 

35.5 

19.7 

17.2 

5.1 

8.6 

6.0 

6.9 

KU  114872  c} 

Nicaragua 

35.9 

19.9 

17.4 

5.1 

8.8 

6.3 

6.8 

KU  114881  c } 

Nicaragua 

36.3 

19.7 

17.0 

5.2 

8.6 

6.3 

7.0 

Carollia  perspicillata 

KU  97645  9 

Nicaragua 

42.3 

23.7 

20.7 

5.4 

9.6 

7.7 

7.8 

KU  110791  9 

Nicaragua 

44.2 

24.3 

21.3 

5.6 

9.7 

7.8 

7.8 

KU  114895  9 

Nicaragua 

42.7 

23.4 

20.5 

5.3 

9.5 

7.5 

7.7 

KU  114896  9 

Nicaragua 

44.8 

23.6 

21.3 

5.5 

9.6 

7.8 

8.2 

KU  110793  c } 

Nicaragua 

44.8 

24.4 

21.4 

5.7 

9.7 

8.0 

8.2 

KU  1 10805  rf 

Nicaragua 

43.8 

24.0 

21.2 

6.0 

10.1 

7.6 

7.8 

KU  110806  c} 

Nicaragua 

43.0 

23.9 

20.7 

5.3 

9.6 

7.8 

7.8 

KU  114897  c} 

Nicaragua 

44.2 

24.4 

21.5 

5.8 

9.9 

7.7 

8.5 

Carollia  subrufa 

KU  114906  9 

N  icaragua 

37.1 

21.3 

18.9 

5.3 

9.2 

6.7 

7.7 

KU  114908  9 

Nicaragua 

39.5 

21.1 

18.8 

5.1 

9.0 

6.7 

7.5 

KU  114915  9 

Nicaragua 

38.4 

21.0 

18.5 

5.0 

8.9 

6.6 

7.5 

KU  114916  9 

Nicaragua 

38.9 

20.8 

18.4 

5.2 

9.0 

6.6 

7.4 

KU  1 14905  6 

Nicaragua 

39.5 

21.5 

19.0 

5.3 

9.4 

6.9 

7.5 

KU  114912c } 

Nicaragua 

38.1 

21.5 

19.2 

5.3 

9.3 

6.9 

7.7 

KU  1 14913  6 

Nicaragua 

38.0 

21.7 

19.3 

5.3 

9.2 

6.7 

7.7 

KU  114914c } 

Nicaragua 

38.7 

21.6 

19.1 

5.3 

9.0 

6.6 

7.5 

Rliinophylla  alethina 

USNM  483445  9 

Colombia 

36.1 

20.4 

17.8 

5.4 

8.8 

5.2 

7.1 

USNM  483446  9 

Colombia 

35.4 

20.4 

17.9 

5.5 

8.9 

4.9 

6.8 

USNM  483447  9 

Colombia 

33.5 

19.0 

16.7 

5.3 

8.8 

4.7 

6.8 

USNM  483449  9 

Colombia 

37.5 

21.3 

18.4 

5.4 

9.0 

5.1 

6.5 

USNM  324988  c} 

Colombia 

35.7 

19.9 

17.3 

5.3 

8.9 

4.9 

6.7 

USNM  483448<} 

Colombia 

34.5 

20.0 

17.4 

5.4 

9.1 

4.8 

6.5 

Rb  inophylla  fisch 

terae 

AMNH  94557  9 

Brazil 

30.5 

16.8 

14.6 

4.8 

7.8 

4.5 

5.9 

TCWC  12102  9 

Peru 

30.0 

17.0 

14.7 

5.1 

7.9 

4.3 

6.1 

USNM  364385  9 

Peru 

30.5 

17.0 

14.8 

5.3 

7.9 

4.2 

6.3 

USNM  364386  9 

Peru 

30.0 

17.0 

14.7 

5.1 

7.6 

4.3 

6.2 

AMNH  94555  c } 

Brazil 

30.6 

16.8 

14.7 

4.7 

7.4 

4.4 

6.1 

TCWC  12096  c} 

Peru 

29.0 

16.2 

14.1 

4.8 

7.9 

4.2 

5.7 

TCWC  12097  c } 

Peru 

29.8 

16.8 

14.5 

5.0 

8.1 

4.3 

5.9 

96 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


Rhinophylla  pnmilio 


USNM  386528  5 

Venezuela 

34.0 

18.7 

16.5 

5.7 

8.3 

4.9 

6.4 

USNM  386530  9 

Venezuela 

34.5 

19.2 

17.1 

5.5 

8.5 

5.2 

6.5 

USNM  386531  9 

Venezuela 

34.8 

19.4 

17.4 

5.5 

8.2 

5.4 

6.6 

USNM  386532  9 

Venezuela 

34.4 

19.8 

17.6 

5.6 

8.7 

5.3 

6.8 

USNM  386539.3 

Venezuela 

34.3 

19.4 

17.5 

5.6 

8.2 

5.2 

6.3 

USNM  386551  c5 

Venezuela 

32.4 

19.1 

16.8 

5.5 

8.4 

5.1 

6.5 

USNM  393674(3 

Brazil 

32.3 

19.3 

16.9 

5.5 

8.2 

5.1 

6.5 

USNM  393676(3 

Brazil 

33.6 

18.9 

16.9 

5.4 

8.2 

4.8 

6.3 

Stenoderminae 

Ametrida  centurio 

TTU  8814  9 

Trinidad 

32.9 

16.4 

13.7 

11.4 

4.1 

8.7 

4.9 

8.1 

TTU  8815  9 

Trinidad 

31.1 

16.0 

13.5 

10.8 

4.0 

8.4 

4.7 

7.7 

TTU  8816  9 

Trinidad 

31.7 

16.5 

13.6 

11.2 

4.4 

8.2 

4.9 

7.9 

TTU  8817  9 

Trinidad 

33.1 

16.7 

13.8 

1 1.4 

4.4 

8.7 

4.8 

8.1 

TTU  5215c3 

Trinidad 

25.2 

15.1 

12.2 

10.3 

4.5 

8.3 

4.0 

7.0 

TTU  8888(3 

Trinidad 

25.5 

15.4 

12.1 

10.6 

4.0 

8.4 

4.1 

7.3 

TTU  9545  (3 

Trinidad 

26.0 

14.9 

11.7 

10.4 

4.0 

8.4 

4.2 

7.3 

TTU  9548  (3 

Trinidad 

24.7 

14.9 

11.3 

10.7 

3.8 

8.6 

4.0 

7.1 

Ardops  rtichollsi 

TTU  20802  9 

Guadeloupe 

49.3 

23.5 

19.9 

15.0 

5.6 

10.2 

7.4 

10.0 

TTU  20820  9 

Guadeloupe 

48.8 

23.2 

20.2 

15.0 

5.8 

10.5 

7.5 

10.1 

TTU  20821  9 

Guadeloupe 

50.8 

23.4 

20.2 

15.3 

5.8 

10.7 

7.5 

10.3 

TTU  20822  9 

Guadeloupe 

51.4 

24.4 

20.8 

15.8 

5.7 

10.6 

7.8 

10.4 

TTU  20806  (3 

Guadeloupe 

47.9 

22.3 

18.7 

14.9 

5.9 

10.6 

6.8 

9.7 

TTU  20808  (3 

Guadeloupe 

47.3 

22.6 

19.3 

15.0 

5.7 

10.7 

7.1 

9.8 

TTU  20809  J 

Guadeloupe 

47.4 

22.3 

19.4 

15.0 

5.8 

10.4 

7.0 

9.6 

TTU  20824  6 

Guadeloupe 

49.6 

22.4 

19.4 

14.7 

5.6 

10.4 

7.1 

9.8 

Ariteus  flavescens 

TTU  21721  9 

Jamaica 

42.7 

20.6 

17.3 

14.2 

4.9 

9.8 

5.9 

9.1 

TTU  21773  9 

Jamaica 

41.3 

19.8 

17.1 

13.9 

4.7 

9.6 

5.9 

8.9 

TTU  21777  9 

Jamaica 

43.0 

21.3 

17.9 

14.5 

5.2 

10.3 

6.2 

9.3 

TTU  21782  9 

Jamaica 

43.1 

20.4 

17.4 

14.4 

4.7 

9.8 

5.9 

8.9 

TTU  21763  (3 

Jamaica 

37.8 

18.5 

15.2 

12.9 

4.5 

9.4 

5.4 

8.2 

TTU  21769(3 

Jamaica 

38.7 

19.3 

15.5 

13.2 

4.7 

9.7 

5.5 

8.5 

TTU  21774(3 

Jamaica 

39.8 

18.6 

15.7 

13.0 

4.6 

9.2 

5.3 

8.2 

TTU  21781  (3 

Jamaica 

38.1 

19.2 

16.0 

13.6 

4.7 

9.5 

5.4 

8.4 

Artibeus  aztecus 

TTU  12907  9 

Costa  Rica 

48.0 

23.2 

20.6 

13.8 

5.5 

10.3 

7.5 

10.6 

TTU  12911  9 

Costa  Rica 

46.6 

22.9 

20.3 

13.8 

5.5 

10.0 

7.5 

10.5 

TTU  12913  9 

Costa  Rica 

44.6 

22.3 

19.7 

12.9 

5.1 

9.8 

7.3 

10.1 

TTU  12914  9 

Costa  Rica 

45.3 

23.1 

20.7 

13.8 

5.3 

10.4 

7.6 

10.6 

KU  94141  (3 

Sinaloa 

42.9 

22.0 

19.6 

13.3 

5.9 

9.8 

7.0 

9.3 

KU  94142  (3 

Sinaloa 

44.2 

22.0 

19.7 

12.7 

5.4 

9.8 

7.0 

9.0 

TTU  12908(3 

Costa  Rica 

46.5 

22.6 

20.1 

13.3 

5.4 

10.2 

7.3 

10.7 

TTU  12910(3 

Costa  Rica 

42.1 

21.8 

19.0 

12.8 

5.0 

9.7 

7.2 

9.8 

Artibeus  cinereus 

TTU  5335  9 

Trinidad 

37.6 

20.3 

18.2 

11.3 

4.6 

9.0 

6.5 

8.2 

TTU  5352  9 

Trinidad 

39.2 

20.0 

18.1 

12.2 

5.0 

8.6 

6.3 

8.4 

TTU  5769  9 

Trinidad 

40.6 

21.2 

18.8 

12.2 

4.7 

9.2 

6.8 

8.5 

TTU  5859  9 

Trinidad 

40.2 

20.5 

18.3 

12.1 

4.9 

8.5 

6.5 

8.6 

TTU  5229  (3 

Trinidad 

39.4 

20.4 

18.5 

11.2 

4.8 

8.9 

6.6 

8.8 

TTU  5230(3 

Trinidad 

38.2 

20.9 

18.5 

11.8 

4.9 

9.0 

6.8 

8.6 

TTU  5541  (5 

Trinidad 

41.8 

21.1 

19.2 

12.3 

5.0 

9.0 

6.8 

9.0 

TTU  9015  (3 

Trinidad 

40.4 

20.8 

18.5 

11.6 

5.1 

8.9 

6.5 

8.7 

Artibeus  concolor 

ROM  36827  9 

Guyana 

48.2 

21.7 

18.9 

13.5 

5.6 

9.9 

6.7 

9.5 

ROM  36830  9 

Guyana 

47.4 

22.0 

19.5 

13.1 

5.1 

10.0 

7.3 

9.4 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


97 


Appendix  1. — Continued. 


ROM  36847  9 

Guyana 

46.3 

22.5 

19.7 

13.6 

5.4 

10.0 

7.0 

9.5 

ROM  60446  9 

Guyana 

48.8 

22.4 

19.8 

13.0 

5.3 

9.4 

7.0 

9.4 

ROM  57444  6 

Guyana 

46.1 

20.6 

17.8 

12.6 

5.5 

9.4 

6.8 

9.1 

ROM  59925  6 

Guyana 

48.4 

21.5 

18.8 

13.1 

5.3 

10.0 

6.8 

9.1 

ROM  66581  & 

Guyana 

49.4 

21.5 

19.0 

13.0 

5.4 

9.1 

7.2 

9.5 

ROM  67478  & 

Guyana 

45.0 

21.3 

18.4 

12.8 

5.3 

9.6 

6.8 

9.2 

Artibeus  glaucus 

AMNH  2I436I  9 

Peru 

38.1 

20.0 

17.9 

11.6 

5.0 

9.1 

6.2 

8.4 

AMNH  233750  9 

Peru 

40.8 

20.6 

18.4 

11.5 

5.4 

9.1 

6.5 

8.4 

AMNH  233751  9 

Peru 

41.5 

20.1 

17.7 

11.5 

5.4 

8.9 

6.3 

8.1 

AMNH  233775  9 

Peru 

40.1 

19.6 

17.2 

11.2 

4.7 

8.9 

6.1 

8.1 

AMNH  214363  & 

Peru 

37.5 

19.0 

17.4 

10.8 

4.7 

8.6 

6.2 

8.2 

AMNH  233755  <3 

Peru 

41.1 

20.5 

18.1 

11.7 

5.2 

9.3 

6.5 

8.4 

AMNH  233763  <3 

Peru 

40.7 

20.2 

17.7 

11.7 

5.0 

9.1 

6.3 

8.6 

AMNH  233771  c 3 

Peru 

41.0 

20.3 

17.9 

11.7 

5.0 

9.3 

6.3 

8.5 

Artibeus  hirsutus 

TTU  8700  9 

Jalisco 

56.0 

27.8 

24.5 

16.8 

6.8 

11.7 

10.0 

12.2 

TTU  8701  9 

Jalisco 

55.0 

26.8 

23.4 

16.8 

6.5 

11.9 

9.4 

11.6 

TTU  8703  9 

Jalisco 

55.2 

27.6 

24.4 

17.3 

6.8 

12.3 

9.7 

11.9 

TTU  8704  9 

Jalisco 

56.9 

27.3 

23.9 

16.8 

6.7 

11.8 

9.6 

11.6 

TTU  8702  (3 

Jalisco 

56.0 

27.1 

23.6 

17.0 

6.9 

12.2 

9.5 

11.8 

TTU  10592c3 

Jalisco 

55.2 

26.7 

23.7 

16.5 

6.8 

12.3 

9.8 

11.4 

TTU  10593  6 

Jalisco 

53.0 

27.0 

23.8 

16.4 

6.9 

12.0 

9.7 

11.6 

TTU  10596d 

Jalisco 

57.3 

26.3 

23.0 

15.7 

6.7 

12.0 

9.8 

11.3 

Artibeus  inopinatus 

TCWC9517  9 

Honduras 

52.8 

26.1 

22.2 

16.2 

5.6 

11.6 

9.0 

10.9 

TTU  7685  9 

Honduras 

50.3 

25.7 

21.9 

15.8 

5.4 

11.2 

8.6 

10.4 

TTU  7686  9 

Honduras 

52.0 

25.8 

22.2 

15.6 

5.4 

1 1.4 

8.8 

10.7 

TTU  12915  9 

Nicaragua 

51.1 

25.3 

21.7 

15.4 

5.4 

11.2 

8.6 

10.6 

TTU  7688  (3 

Honduras 

50.0 

25.9 

22.4 

15.6 

5.4 

11.7 

8.9 

10.7 

TTU  7689  (3 

Honduras 

50.0 

25.2 

21.5 

15.7 

5.3 

11.3 

8.7 

10.6 

TTU  7690  <3 

Honduras 

50.2 

25.2 

21.8 

15.6 

5.6 

1  1.4 

8.8 

10.7 

TTU  129163 

Nicaragua 

50.0 

25.6 

21.7 

15.5 

5.4 

1  1.4 

8.6 

10.6 

Artibeus  jamaicensis 

AS  5234  9 

Jamaica 

61.4 

29.5 

26.1 

17.1 

7.2 

12.8 

10.4 

13.0 

AS  5236  9 

Jamaica 

57.0 

28.3 

24.7 

17.0 

7.1 

12.1 

9.5 

12.4 

KU  97801  9 

Nicaragua 

60.1 

29.3 

25.7 

17.4 

6.9 

12.7 

9.8 

12.0 

KU  97802  9 

Nicaragua 

56.4 

27.9 

24.3 

17.0 

7.2 

12.2 

9.5 

12.1 

COLU  316  3 

Jamaica 

59.2 

28.7 

24.8 

17.3 

7.2 

12.6 

10.0 

12.8 

COLU  323  3 

Jamaica 

57.3 

27.8 

24.5 

16.8 

6.7 

12.0 

9.6 

12.1 

AMNH  28335  3 

Nicaragua 

56.4 

29.4 

25.7 

16.9 

7.0 

12.4 

10.4 

12.5 

KU  115030  3 

Nicaragua 

58.8 

28.8 

24.9 

17.7 

7.3 

12.9 

9.7 

12.9 

Artibeus  lituratus 

KU  115967  9 

Nicaragua 

67.3 

31.7 

27.7 

19.2 

6.4 

13.2 

10.4 

12.9 

KU  115068  9 

Nicaragua 

72.6 

31.9 

28.8 

19.1 

6.6 

13.7 

11.1 

13.6 

KU  115069  9 

Nicaragua 

70.5 

31.1 

27.3 

18.9 

6.3 

13.9 

10.4 

13.5 

KU  115072  9 

Nicaragua 

71.1 

32.1 

28.3 

19.9 

6.5 

14.3 

1 1.2 

13.8 

KU  115062  3 

Nicaragua 

69.3 

31.7 

27.8 

19.5 

7.0 

14.2 

10.9 

13.0 

KU  115065  3 

Nicaragua 

72.8 

31.1 

27.1 

19.0 

6.7 

14.0 

10.1 

12.9 

KU  115070  3 

Nicaragua 

73.0 

31.9 

27.9 

19.6 

6.4 

14.0 

11.1 

13.6 

KU  115071  3 

Nicaragua 

69.3 

31.0 

27.2 

18.9 

6.6 

13.6 

11.0 

13.0 

Artibeus  phaeotis 

KU  106145  9 

Nicaragua 

35.1 

18.3 

15.9 

10.6 

4.5 

8.6 

5.6 

7.7 

KU  106146  9 

Nicaragua 

34.2 

18.3 

16.0 

11.3 

4.8 

8.5 

5.7 

7.5 

KU  106153  9 

Nicaragua 

35.7 

19.4 

17.2 

11.8 

4.7 

9.0 

5.8 

7.7 

KU  106155  9 

Nicaragua 

34.9 

18.5 

16.1 

11.1 

4.8 

8.9 

5.6 

7.7 

KU  106147  3 

Nicaragua 

34.9 

18.3 

15.8 

11.1 

4.8 

9.0 

5.6 

7.5 

KU  106148  3 

Nicaragua 

37.2 

18.1 

15.7 

10.9 

4.5 

8.4 

5.6 

7.7 

98 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. —  Continued. 


KU  105149  <3 

N  icaragua 

36.9 

18.3 

16.0 

10.8 

4.3 

8.8 

5.7 

7.4 

KU  106150  6 

Nicaragua 

36.3 

19.0 

17.0 

11.5 

4.5 

8.7 

5.8 

7.7 

Artibeus  toltecus 

TTU  73519 

Tamaulipas 

39.4 

20.5 

17.9 

12.2 

4.7 

9.2 

6.6 

8.9 

TTU  7354  9 

Tamaulipas 

37.5 

20.8 

18.1 

12.5 

5.1 

9.3 

6.6 

8.9 

TTU  7355  9 

Tamaulipas 

39.7 

21.5 

18.7 

12.5 

4.9 

9.4 

6.7 

9.0 

TTU  12930  9 

Honduras 

40.3 

21.4 

19.4 

12.7 

5.7 

9.7 

6.9 

8.7 

TTU  8163  i 

San  Luis  Polosi 

36.9 

19.5 

17.2 

12.6 

5.0 

9.4 

6.4 

8.9 

TTU  12929  6 

Honduras 

40.6 

21.0 

18.7 

12.0 

5.6 

9.3 

6.9 

8.8 

TTU  12931  6 

El  Salvador 

39.0 

20.3 

17.9 

11.7 

5.0 

9.2 

6.7 

9.0 

TTU  12932  6 

El  Salvador 

40.2 

19.7 

17.4 

11.5 

4.8 

9.2 

6.2 

8.7 

Artibeus  walsoni 

KU  82102  9 

Guatemala 

37.8 

19.7 

17.8 

11.2 

4.6 

8.8 

6.4 

8.2 

TTU  12964  9 

Honduras 

36.3 

19.0 

16.3 

11.2 

4.7 

8.7 

5.8 

8.4 

TTU  12967  9 

Honduras 

38.8 

19.8 

17.3 

12.1 

5.0 

8.9 

6.1 

8.6 

KU  111171  9 

Nicaragua 

38.5 

19.9 

17.6 

11.5 

4.9 

8.7 

6.6 

8.4 

TTU  12962  6 

Honduras 

37.5 

19.1 

16.6 

11.7 

4.7 

9.0 

6.0 

8.5 

TTU  12963  <3 

Honduras 

37.6 

19.8 

17.1 

11.8 

4.8 

8.8 

6.0 

8.5 

TTU  12934(3 

Nicaragua 

39.3 

20.0 

17.7 

11.3 

4.9 

8.5 

6.2 

8.0 

TTU  12948(3 

Nicaragua 

37.9 

19.5 

17.4 

11.4 

4.8 

8.6 

6.2 

8.3 

Centurio  sene x 

FHKSC  9813  9 

Chiapas 

45.7 

18.5 

15.1 

15.0 

5.8 

10.4 

4.8 

10.7 

TTU  13076  9 

Honduras 

42.6 

18.9 

14.5 

14.9 

5.7 

9.3 

4.8 

10.6 

KU  115113  9 

Nicaragua 

42.6 

19.0 

14.8 

14.9 

5.9 

9.8 

5.0 

10.6 

KU  115114  9 

Nicaragua 

43.5 

18.9 

15.0 

15.0 

6.0 

10.0 

4.7 

10.6 

FHKSC  9812(3 

Chiapas 

42.0 

18.7 

14.5 

14.8 

5.5 

10.0 

4.7 

10.5 

KU  115108(3 

Nicaragua 

41.6 

18.7 

14.6 

14.5 

5.6 

10.6 

4.6 

10.3 

KU  115111  (3 

Nicaragua 

42.7 

19.0 

14.5 

15.0 

5.7 

10.8 

4.6 

10.4 

TTU  5221  (3 

Trinidad 

44.1 

19.8 

15.2 

15.8 

6.1 

10.5 

5.1 

11.1 

Chiroderma 

doriae 

BMNH  9.1 1.19.15  9 

Brazil 

53.7 

28.0 

25.9 

17.8 

6.1 

11.2 

10.3 

13.6 

TTU  30707  (3 

Brazil 

28.1 

25.8 

17.6 

6.4 

12.0 

10.0 

13.0 

TTU  30708  c3 

Brazil 

28.8 

26.3 

17.9 

6.2 

12.0 

10.2 

13.4 

TTU  30709  9 

Brazil 

29.0 

26.4 

18.1 

6.3 

12.5 

10.2 

13.5 

Chiroderma  improvisum 

TTU  19900(3 

Guadeloupe 

57.5 

29.9 

27.7 

18.9 

6.5 

12.2 

10.7 

7.2 

Chiroderma  salvini 

USNM  33871  1  9 

Colima 

46.1 

24.2 

22.0 

15.2 

6.2 

10.6 

8.6 

1  1.4 

TCWC  17499  9 

Guatemala 

47.8 

26.4 

23.8 

16.1 

6.0 

11.0 

9.2 

11.5 

TTU  12809  9 

Honduras 

51.8 

27.6 

24.8 

16.9 

6.1 

11.2 

9.5 

12.1 

AMNH  142484  9 

Costa  Rica 

51.5 

27.6 

24.8 

17.5 

6.3 

11.6 

10.1 

13.0 

TTU  6123  (3 

Colima 

43.6 

24.5 

21.9 

15.0 

5.8 

10.5 

8.4 

11.1 

TTU  12800(3 

Honduras 

48.0 

26.6 

24.1 

16.2 

6.2 

11.0 

9.4 

11.7 

TTU  12801  c3 

Honduras 

45.6 

26.0 

23.6 

16.0 

5.7 

11.0 

9.1 

11.8 

TTU  12802c3 

Honduras 

49.4 

26.6 

24.2 

16.6 

6.2 

11.2 

9.3 

12.2 

Chiroderma  trinitatum 

TTU  5223  9 

Trinidad 

41.5 

22.7 

19.8 

13.4 

5.4 

9.5 

7.8 

10.6 

TTU  5224  9 

Trinidad 

38.0 

22.5 

20.0 

13.3 

5.3 

9.4 

7.5 

10.1 

TTU  5336  9 

Trinidad 

40.3 

22.4 

19.7 

13.8 

5.4 

9.5 

7.5 

10.1 

TTU  5382  9 

Trinidad 

38.8 

22.5 

19.6 

13.7 

5.4 

9.6 

7.4 

10.2 

TTU  5487  (3 

Trinidad 

39.0 

22.4 

19.8 

13.5 

5.5 

9.7 

7.4 

10.3 

TTU  5675  (3 

Trinidad 

38.7 

22.5 

19.8 

13.8 

5.2 

9.5 

7.4 

9.6 

TTU  8989  (3 

Trinidad 

39.5 

22.3 

19.8 

13.6 

5.3 

9.6 

7.5 

9.7 

TTU  9014  (3 

Trinidad 

39.1 

22.2 

19.5 

13.5 

5.5 

9.4 

7.4 

10.0 

Chiroderma  villosum 

TTU  5289  9 

Trinidad 

46.5 

26.0 

23.4 

16.4 

5.7 

10.8 

9.1 

11.6 

TTU  5321  9 

Trinidad 

45.3 

25.0 

22.4 

16.0 

5.6 

11.0 

8.7 

11.5 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


99 


Appendix  1. — Continued. 


TTU  5353  9 

Trinidad 

47.9 

26.6 

23.6 

16.5 

5.8 

11.0 

9.1 

12.0 

TTU  5354  9 

Trinidad 

47.2 

26.2 

23.4 

17.0 

6.2 

10.4 

9.0 

12.0 

TTU  5262  c? 

Trinidad 

45.9 

26.4 

23.3 

16.4 

6.2 

11.3 

9.1 

11.5 

TTU  5276  c? 

Trinidad 

46.0 

25.3 

22.4 

15.7 

5.9 

10.5 

8.5 

11.4 

TTU  5668  <5 

Trinidad 

44.3 

26.0 

22.9 

15.7 

6.1 

10.8 

9.0 

11.6 

TTU  9016  c? 

Trinidad 

46.8 

26.5 

22.8 

15.1 

5.8 

10.6 

8.6 

10.9 

Ectophylla 

alba 

KU  88025  9 

Costa  Rica 

28.1 

16.4 

15.1 

9.8 

4.0 

7.3 

6.0 

7.4 

USNM  335318  9 

Panama 

29.1 

16.4 

15.5 

10.0 

4.2 

7.5 

6.0 

7.2 

USNM  335320  9 

Panama 

28.9 

16.7 

15.5 

10.0 

4.0 

7.7 

6.0 

7.3 

USNM  335322  9 

Panama 

29.4 

16.3 

15.2 

10.3 

4.3 

8.0 

6.0 

7.6 

TCWC  19372<? 

Honduras 

28.4 

17.1 

15.7 

10.1 

4.2 

7.9 

6.1 

7.3 

TCWC  19373  c? 

Honduras 

28.5 

17.1 

15.7 

10.3 

4.2 

7.8 

6.3 

7.4 

USNM  315563  c? 

Panama 

28.4 

16.9 

15.4 

10.3 

4.3 

7.8 

6.1 

7.5 

USNM  319426c? 

Panama 

28.7 

16.5 

15.4 

9.9 

4.2 

7.5 

6.0 

7.2 

Enchisthenes 

hurtii 

AMNH  206872  9 

Oaxaca 

40.1 

21.1 

19.1 

13.0 

6.1 

9.5 

6.8 

9.0 

KU  102600  9 

Chiapas 

39.5 

20.7 

18.7 

12.2 

5.9 

9.3 

6.6 

8.5 

TTU  5371  9 

Trinidad 

38.6 

20.5 

18.3 

12.0 

5.7 

9.4 

6.8 

8.7 

AMNH  233798  9 

Peru 

36.7 

20.3 

18.6 

12.4 

5.9 

9.6 

7.0 

8.6 

KU  97039  c? 

Jalisco 

39.8 

21.0 

18.9 

12.9 

5.9 

9.8 

6.7 

8.9 

AMNH  126239  c? 

Honduras 

36.5 

20.9 

18.6 

11.3 

5.4 

9.4 

7.1 

8.6 

BMNH  92.9.7.8  c? 

Trinidad 

37.1 

20.4 

18.5 

12.0 

6.1 

9.1 

7.2 

8.4 

AMNH  233599  c? 

Peru 

39.6 

20.9 

18.9 

12.1 

5.7 

9.6 

6.7 

8.0 

Mesophylta  macconnelli 

TTU  5359  9 

Trinidad 

32.6 

18.6 

16.6 

10.7 

4.6 

8.2 

6.2 

7.6 

TTU  5475  9 

Trinidad 

31.5 

18.2 

16.3 

10.4 

4.5 

7.8 

6.3 

7.4 

TTU  9786  9 

Trinidad 

33.5 

19.0 

16.7 

10.8 

4.8 

8.3 

6.5 

7.7 

BMNH  1.6.4.64  9 

Guyana 

30.0 

17.7 

15.5 

10.1 

4.4 

7.8 

6.0 

7.0 

TTU  5211  c? 

Trinidad 

32.0 

18.5 

16.5 

10.6 

4.7 

8.2 

6.1 

7.5 

TTU  5212  c? 

Trinidad 

31.5 

18.5 

16.3 

10.8 

4.5 

8.3 

6.2 

7.5 

TTU  5213  c? 

Trinidad 

32.3 

18.7 

16.6 

11.0 

4.6 

8.2 

6.2 

7.7 

BMNH  70.1008  c? 

Brazil 

29.5 

17.7 

15.4 

9.8 

4.2 

7.7 

5.8 

7.0 

Phyllops  falcatus 

AMNH  176190  9 

Cuba 

44.0 

20.9 

18.9 

14.2 

5.6 

10.0 

6.0 

8.7 

USNM  143844  9 

Cuba 

43.3 

20.8 

18.7 

14.1 

5.3 

10.0 

6.2 

8.5 

BMNH  c? 

Cuba 

42.9 

5.3 

5.8 

8.1 

Phyllops  halt 

iensis 

TTU  22675  9 

Haiti 

41.8 

20.3 

18.3 

13.7 

5.7 

10.0 

5.9 

8.2 

TTU  22676  9 

Haiti 

43.8 

20.7 

18.3 

13.6 

5.4 

10.1 

6.2 

8.5 

TTU  22677  9 

Haiti 

44.0 

20.4 

18.4 

13.8 

5.7 

10.3 

6.1 

8.4 

TTU  22678  9 

Haiti 

42.8 

20.5 

18.3 

13.2 

5.5 

9.9 

6.1 

8.3 

TTU  22697  c? 

Haiti 

39.0 

19.4 

17.2 

12.5 

5.2 

9.6 

5.5 

7.8 

TTU  22698  c? 

Haiti 

40.2 

19.4 

17.5 

12.9 

5.3 

9.6 

5.6 

7.9 

TTU  22699  c? 

Haiti 

40.9 

19.5 

17.4 

13.2 

5.7 

9.9 

5.8 

7.9 

TTU  22700  c? 

Haiti 

42.1 

19.7 

17.4 

13.3 

5.4 

9.9 

5.6 

8.1 

Phyllops  veins 

AMNH  41001  ? 

Cuba 

18.1 

13.5 

5.4 

5.9 

7.9 

AMNH  41002  ? 

Cuba 

19.5 

17.3 

13.0 

5.2 

9.7 

5.4 

7.3 

AMNH  41003? 

Cuba 

20.1 

18.0 

5.3 

10.0 

5.5 

7.5 

AMNH  41005  ? 

Cuba 

17.0 

5.0 

10.0 

5.3 

7.3 

Pygoderma  hilabialum 

AMNH  234288  9 

Paraguay 

38.9 

20.5 

17.5 

14.0 

7.4 

9.9 

6.0 

8.0 

AMNH  234290  9 

Paraguay 

37.6 

20.9 

17.5 

14.3 

8.0 

10.1 

5.7 

8.0 

AMNH  234292  9 

Paraguay 

39.8 

21.0 

17.9 

14.7 

7.7 

10.3 

6.1 

8.4 

KU  92656  9 

Paraguay 

39.5 

20.2 

17.4 

14.1 

7.4 

10.1 

6.0 

7.9 

AMNH  234291  c? 

Paraguay 

36.4 

20.1 

16.5 

13.2 

7.2 

10.0 

5.4 

7.1 

100 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


AMNH  234294  c? 

Paraguay 

36.6 

20.0 

16.8 

13.4 

7.3 

10.0 

5.3 

7.2 

AMNH  234297  c? 

Paraguay 

36.2 

20.5 

17.2 

13.7 

7.7 

10.4 

5.5 

7.5 

AMNH  234298  c? 

Paraguay 

37.0 

19.9 

17.0 

13.7 

7.5 

10.3 

5.4 

7.3 

Sphaeronycleris 

toxophyllum 

TCWC  28252  9 

Venezuela 

39.5 

17.2 

14.2 

12.1 

5.6 

9.5 

4.7 

7.9 

USNM  370848  9 

Venezuela 

40.0 

17.4 

14.5 

12.2 

5.7 

9.4 

4.6 

7.9 

USNM  370849  9 

Venezuela 

40.1 

17.2 

14.5 

12.3 

5.7 

9.2 

4.7 

7.8 

AMNH  209704  9 

Bolivia 

39.6 

17.5 

14.6 

12.1 

5.6 

9.0 

4.4 

8.0 

TTU  10227  c? 

Colombia 

36.6 

16.1 

13.8 

11.7 

5.5 

8.9 

4.4 

7.2 

USNM  405688  c? 

Venezuela 

37.0 

16.8 

13.9 

12.2 

5.6 

9.5 

4.3 

7.3 

USNM  409233  c? 

Venezuela 

37.3 

16.5 

13.4 

11.9 

5.6 

8.9 

4.4 

7.4 

AMNH  209741  c? 

Bolivia 

38.7 

16.9 

13.8 

12.4 

5.7 

9.0 

4.2 

7.6 

Stenoderma  rufum 

TTU  8876  9 

Puerto  Rico 

49.0 

23.0 

19.4 

15.5 

5.7 

11.4 

7.2 

10.1 

TTU  8879  9 

Puerto  Rico 

49.0 

22.5 

19.1 

15.2 

5.7 

10.6 

6.8 

9.7 

TTU  8880  9 

Puerto  Rico 

51.2 

23.5 

19.8 

15.8 

5.7 

11.4 

7.0 

10.2 

TTU  8884  9 

Puerto  Rico 

50.3 

22.9 

19.4 

15.3 

5.7 

10.7 

7.0 

10.0 

TTU  8860<? 

Puerto  Rico 

46.5 

22.2 

18.5 

15.0 

5.5 

10.5 

6.6 

9.7 

TTU  8861  <J 

Puerto  Rico 

47.1 

22.5 

19.0 

14.9 

5.6 

10.6 

6.6 

9.7 

TTU  8864  $ 

Puerto  Rico 

46.1 

22.0 

18.0 

14.4 

5.2 

10.2 

6.2 

9.7 

TTU  8865  <? 

Puerto  Rico 

48.5 

22.5 

18.7 

14.9 

5.4 

10.7 

6.3 

9.5 

Sturnira  aratathomasi 

ROM  70874  9 

Colombia 

58.0 

29.1 

25.4 

17.2 

7.5 

12.9 

7.6 

10.2 

USNM  501064  9 

Colombia 

57.5 

28.5 

25.5 

16.9 

6.9 

12.5 

7.7 

10.1 

USNM  501066  9 

Colombia 

56.8 

28.8 

25.0 

16.7 

7.2 

12.5 

7.4 

9.7 

ROM  46349  9 

Ecuador 

60.5 

29.7 

26.2 

17.5 

7.2 

12.8 

8.1 

10.5 

ROM  70875  <? 

Colombia 

57.7 

29.4 

26.5 

16.7 

7.2 

12.8 

7.8 

10.4 

ROM  70876  e? 

Colombia 

54.8 

28.8 

25.2 

16.8 

7.0 

12.7 

7.6 

10.2 

USNM  395158  c? 

Colombia 

57.1 

29.4 

26.5 

17.5 

7.3 

13.0 

7.9 

10.1 

USNM  501065  c? 

Colombia 

57.5 

28.8 

25.9 

16.5 

6.9 

12.3 

7.7 

10.0 

Sturnira  i 

bide  ns 

USNM  386557  9 

Venezuela 

39.3 

21.2 

18.9 

11.7 

5.5 

9.4 

6.0 

6.8 

USNM  386558  9 

Venezuela 

40.2 

21.6 

19.7 

11.7 

5.3 

9.7 

6.0 

6.8 

USNM  386560  9 

Venezuela 

39.7 

22.1 

19.6 

12.0 

5.5 

9.8 

6.1 

7.1 

USNM  386562  9 

Venezuela 

40.8 

21.7 

19.5 

12.0 

5.5 

9.6 

6.0 

6.9 

USNM  386559  c? 

Venezuela 

39.7 

21.2 

19.0 

11.9 

5.4 

9.6 

5.7 

6.9 

USNM  386567  c? 

Venezuela 

39.7 

21.3 

18.7 

11.7 

5.4 

9.5 

5.9 

7.0 

USNM  386570  c? 

Venezuela 

39.5 

21.0 

18.7 

11.7 

5.3 

9.6 

5.9 

6.6 

AMNH  214349  c? 

Peru 

41.2 

21.3 

18.7 

11.7 

5.4 

9.7 

5.9 

6.7 

Sturnira  erythromos 

ROM  67254  9 

Colombia 

39.3 

21.3 

18.6 

12.7 

5.7 

10.0 

6.0 

8.0 

ROM  67267  9 

Colombia 

41.1 

20.7 

18.3 

12.0 

5.3 

9.5 

5.9 

7.5 

USNM  483451  9 

Colombia 

40.6 

21.5 

19.0 

12.7 

6.0 

9.9 

5.8 

7.5 

USNM  483452  9 

Colombia 

40.6 

21.0 

18.9 

12.5 

6.1 

9.7 

6.0 

7.4 

ROM  67270  c? 

Colombia 

41.6 

21.4 

19.2 

12.1 

6.0 

9.6 

5.9 

7.4 

BMNH  15.7.1 1.13  c? 

Ecuador 

40.8 

22.0 

19.4 

12.9 

6.0 

10.4 

6.3 

8.0 

Sturnira 

1  ilium 

TTU  5367  9 

Trinidad 

42.5 

23.4 

20.4 

13.6 

6.0 

10.3 

6.4 

8.4 

TTU  5407  9 

Trinidad 

43.9 

22.9 

20.2 

13.7 

5.8 

10.5 

6.4 

8.2 

TTU  5669  9 

Trinidad 

42.4 

22.7 

19.9 

13.4 

5.6 

10.1 

6.5 

8.0 

TTU  5670  9 

Trinidad 

41.4 

22.8 

19.9 

13.6 

6.2 

10.4 

6.3 

8.0 

TTU  5408  c? 

Trinidad 

43.2 

23.1 

20.4 

13.8 

6.0 

10.4 

6.6 

8.2 

TTU  5415  c? 

Trinidad 

41.9 

23.2 

20.3 

13.7 

6.3 

10.5 

6.3 

7.9 

TTU  5775  c? 

Trinidad 

41.3 

22.9 

20.4 

13.6 

6.4 

10.5 

6.8 

8.5 

TTU  5776  c? 

Trinidad 

42.7 

22.4 

19.6 

13.4 

5.9 

10.5 

6.4 

8.0 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


101 


TTU  15543  9 
TTU  15546  9 
TCWC  14359  9 
TCWC  14360  9 
TTU  7341  £ 
TTU  6124  £ 
TTU  6125  £ 
KU  97689  £ 


AMNH  214347  9 
TCWC  27474  9 
LSU  16518  9 
LSU  19031  9 
LSU  16517(3 
LSU  19027  £ 
LSU  19028<J 


BMNH  69.1263  9 
TCWC  10034(3 
TCWC  10035c3 
TCWC  10041  £ 
TCWC  10042  £ 


AMNH  219138  9 
LSU  16521  9 
LSU  16522  9 
LSU  16524  9 
AMNH  219171  £ 
AMNH  219172  (3 
AMNH  219173  £ 
TCWC  28071  £ 


TTU  19904  9 
TTU  19905  9 
TTU  19906  9 
TTU  19907  9 
AMNH  234950(5 
USNM  361883  (3 


TTU  5406  9 
TTU  5667  9 
TTU  5786  9 
TTU  5791  9 
TTU  5337  £ 
TTU  5372  £ 
TTU  5402  £ 
TTU  5454(3 


KU  114985  9 
TTU  5327  9 
TTU  5485  9 
TTU  5813  9 
KU  114986(5 
TTU  5254(3 
TTU  5300(3 
TTU  5301  £ 


Appendix  1. — Continued. 


Sturnira  ludovici 


Hidalgo 

44.1 

24.0 

20.8 

13.0 

6.1 

10.3 

6.5 

8.0 

Hidalgo 

43.2 

24.0 

20.8 

13.5 

6.1 

10.2 

6.5 

8.0 

Guatemala 

45.1 

23.2 

20.1 

13.2 

5.6 

10.3 

6.3 

8.0 

Guatemala 

46.9 

24.4 

21.6 

13.9 

6.1 

10.6 

6.6 

8.3 

Tamaulipas 

42.5 

23.9 

21.0 

13.6 

6.0 

10.6 

6.4 

8.3 

Jalisco 

44.3 

23.4 

20.3 

13.9 

6.1 

10.0 

6.2 

7.9 

Jalisco 

43.2 

23.2 

20.1 

12.7 

6.0 

10.1 

6.2 

7.8 

Nicaragua 

45.1 

24.0 

21.4 

14.2 

6.2 

10.4 

6.5 

8.3 

Sturnira  magna 

Peru 

59.2 

29.0 

25.3 

16.7 

6.9 

12.0 

7.1 

9.0 

Peru 

57.7 

27.9 

24.7 

16.4 

7.0 

11.8 

7.2 

8.8 

Peru 

57.7 

29.1 

25.6 

17.2 

7.0 

12.5 

7.4 

9.9 

Peru 

57.4 

28.5 

24.4 

16.0 

6.8 

11.5 

7.3 

8.8 

Peru 

57.0 

29.5 

25.6 

17.2 

7.0 

11.9 

7.4 

9.3 

Peru 

55.4 

28.5 

24.7 

17.0 

7.0 

12.1 

7.1 

9.1 

Peru 

56.0 

28.8 

24.9 

16.9 

6.9 

12.2 

7.5 

9.3 

Sturnira  mordax 

Costa  Rica 

46.2 

25.8 

22.4 

13.1 

5.9 

10.6 

6.7 

7.8 

Costa  Rica 

48.3 

26.1 

22.9 

13.8 

6.1 

10.9 

6.9 

8.2 

Costa  Rica 

46.1 

25.5 

22.0 

13.3 

5.9 

11.0 

6.7 

7.9 

Costa  Rica 

47.7 

25.7 

22.4 

13.3 

6.0 

10.7 

6.7 

7.8 

Costa  Rica 

48.3 

26.3 

23.1 

13.7 

6.2 

10.9 

6.9 

8.0 

Sturnira 

nana 

Peru 

34.7 

18.8 

16.6 

10.2 

4.6 

8.2 

4.8 

5.8 

Peru 

34.8 

18.5 

16.5 

10.0 

4.7 

8.1 

4.7 

5.6 

Peru 

33.7 

18.9 

16.6 

9.8 

4.8 

8.3 

4.8 

5.7 

Peru 

34.1 

19.0 

16.8 

10.1 

4.6 

8.5 

4.9 

6.0 

Peru 

34.5 

18.7 

16.5 

10.1 

4.7 

8.5 

4.7 

5.5 

Peru 

35.4 

18.8 

16.5 

10.1 

4.6 

8.5 

4.9 

5.8 

Peru 

35.0 

18.5 

16.3 

9.7 

4.7 

8.2 

4.7 

5.6 

Peru 

32.6 

18.4 

16.1 

9.9 

4.7 

8.1 

4.8 

5.7 

Sturnira  thomasi 

Guadeloupe 

45.9 

25.3 

23.3 

12.1 

5.7 

9.8 

7.0 

8.1 

Guadeloupe 

46.4 

24.4 

22.4 

11.9 

5.6 

9.5 

6.7 

7.7 

Guadeloupe 

46.1 

24.9 

22.9 

12.2 

5.5 

9.8 

6.9 

8.0 

Guadeloupe 

47.7 

25.1 

23.6 

12.5 

5.9 

9.6 

6.9 

8.0 

Guadeloupe 

46.5 

25.1 

23.7 

12.2 

5.7 

9.5 

6.7 

8.2 

Guadeloupe 

48.1 

26.2 

24.7 

12.7 

6.0 

9.9 

7.7 

8.2 

Sturnira  tildae 

Trinidad 

44.0 

23.6 

21.1 

14.6 

6.0 

10.7 

6.8 

8.1 

Trinidad 

44.1 

23.9 

21.5 

14.3 

6.1 

10.8 

6.9 

8.5 

Trinidad 

44.7 

24.3 

21.8 

14.3 

5.7 

10.6 

7.1 

8.3 

Trinidad 

43.4 

22.8 

20.2 

13.7 

6.1 

10.4 

6.6 

7.8 

Trinidad 

44.7 

23.9 

21.1 

14.2 

6.3 

10.7 

7.1 

8.5 

Trinidad 

44.5 

23.1 

20.5 

14.0 

5.9 

10.6 

6.9 

8.2 

Trinidad 

46.3 

24.4 

22.2 

14.7 

6.6 

10.7 

7.4 

8.6 

Trinidad 

44.2 

23.7 

21.5 

14.6 

6.5 

11.0 

6.8 

8.8 

Uroderma  bilobatum 

Nicaragua 

41.6 

22.8 

20.2 

13.0 

5.4 

9.5 

7.9 

9.3 

Trinidad 

42.1 

23.6 

20.8 

12.8 

4.6 

9.5 

8.2 

9.4 

Trinidad 

39.6 

23.0 

20.6 

12.5 

5.3 

9.3 

7.9 

9.3 

Trinidad 

42.4 

23.6 

21.4 

13.0 

5.6 

9.4 

8.5 

9.9 

Nicaragua 

43.0 

22.6 

19.9 

12.7 

5.3 

9.9 

7.8 

9.1 

Trinidad 

43.1 

24.7 

22.1 

13.4 

5.7 

9.9 

8.6 

9.9 

Trinidad 

40.5 

23.7 

21.6 

13.1 

5.4 

9.7 

8.3 

9.5 

Trinidad 

41.4 

24.0 

21.0 

12.8 

5.5 

9.4 

7.9 

8.9 

102 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


Uroderma  magnirostrum 


TTU  17111  9 

El  Salvador 

42.3 

23.0 

20.8 

12.7 

5.7 

9.7 

7.6 

8.9 

KU  114987  9 

Nicaragua 

45.1 

23.9 

21.8 

13.1 

5.8 

9.3 

8.3 

9.1 

TTU  9080  9 

Colombia 

42.6 

23.1 

21.0 

12.6 

5.6 

9.3 

8.1 

9.2 

TTU  9517  9 

Colombia 

41.8 

22.4 

20.3 

12.2 

5.6 

9.3 

7.8 

8.9 

TCWC  17189  6 

Honduras 

41.0 

22.5 

20.5 

12.6 

5.6 

9.5 

7.6 

8.6 

KU  106109  c? 

Nicaragua 

41.6 

23.6 

21.7 

12.9 

5.5 

9.5 

7.7 

8.7 

TTU  9054  c? 

Colombia 

43.6 

24.0 

21.7 

13.5 

6.0 

10.3 

8.1 

9.5 

TTU  9056  c? 

Colombia 

43.4 

23.5 

21.3 

12.9 

5.3 

9.7 

8.0 

9.1 

Vampyressa  bidens 


ROM  59895  9 

Guyana 

36.0 

20.2 

17.2 

11.8 

5.4 

9.1 

6.3 

8.3 

ROM  66587  9 

Guyana 

38.2 

20.5 

17.8 

12.3 

5.3 

9.0 

6.4 

8.9 

AMNH  208072  9 

Peru 

36.6 

20.4 

17.8 

12.1 

5.1 

8.8 

6.5 

8.9 

TCWC  27508  9 

Peru 

36.4 

19.5 

16.8 

11.4 

4.8 

8.6 

5.9 

8.1 

AMNH  98780.? 

Peru 

39.1 

20.0 

17.1 

12.2 

5.2 

9.1 

6.1 

8.6 

TCWC  27503  c? 

Peru 

35.6 

20.0 

17.3 

11.7 

5.2 

9.3 

6.2 

8.5 

TCWC  27505  <? 

Peru 

35.3 

19.8 

17.1 

11.2 

5.2 

8.4 

6.3 

8.2 

TCWC  27506  c? 

Peru 

35.5 

20.2 

17.3 

12.2 

5.4 

9.1 

6.3 

8.3 

Vampyressa  brocki 


TTU  8827  9 

Colombia 

35.4 

18.4 

16.0 

10.9 

4.9 

8.4 

5.7 

7.9 

TTU  8832  9 

Colombia 

32.1 

18.3 

15.8 

10.8 

4.7 

8.4 

5.7 

7.6 

TTU  9047  9 

Colombia 

33.2 

18.4 

16.2 

10.7 

5.1 

8.0 

5.7 

7.8 

ROM  38515  9 

Guyana 

33.0 

17.7 

15.5 

10.4 

4.7 

7.8 

5.6 

7.6 

Vampyressa  melissa 


BMNH  26.5.3.4  9 

Peru 

37.1 

21.5 

19.6 

12.8 

5.0 

9.0 

6.8 

9.6 

LSU  16580  9 

Peru 

39.2 

22.2 

20.4 

13.2 

5.2 

9.5 

7.1 

9.5 

LSU  16583  9 

Peru 

38.2 

21.8 

20.1 

12.9 

5.1 

8.9 

6.7 

9.2 

LSU  19100  9 

Peru 

37.3 

21.3 

19.8 

13.1 

5.1 

8.9 

6.6 

9.1 

USNM  319283  c? 

Panama 

37.9 

22.8 

21.2 

12.0 

5.1 

8.8 

7.6 

9.1 

USNM  319284  c? 

Panama 

36.5 

22.8 

21.3 

12.1 

5.2 

9.1 

7.5 

9.2 

USNM  319285  c? 

Panama 

37.8 

22.8 

21.3 

12.3 

5.1 

9.0 

7.7 

9.3 

AMNH  233769  c? 

Peru 

36.5 

21.9 

20.0 

13.1 

5.2 

9.4 

6.9 

9.4 

Vampyressa  nymphaea 


KU  115005  9 

N  icaragua 

36.2 

21.1 

18.4 

12.3 

4.7 

9.2 

7.0 

8.6 

TCWC  19368  9 

Nicaragua 

35.7 

21.2 

18.0 

12.1 

4.9 

9.2 

7.1 

8.7 

TTU  12611  9 

Nicaragua 

37.9 

21.6 

19.1 

13.0 

4.6 

9.4 

7.5 

9.4 

USNM  483687  9 

Colombia 

39.0 

21.6 

18.7 

12.2 

4.8 

9.3 

7.0 

8.8 

TCWC  19367  c? 

Nicaragua 

38.2 

21.7 

18.7 

12.4 

4.5 

9.3 

7.1 

8.8 

TTU  12612  c? 

Nicaragua 

37.4 

22.0 

19.0 

12.8 

5.0 

9.5 

7.5 

9.1 

AMNH  233189  c? 

Colombia 

37.0 

21.2 

18.4 

11.9 

4.9 

9.3 

6.8 

8.9 

BMNH  9.7.17.40  <? 

Colombia 

34.9 

21.0 

18.3 

12.2 

4.7 

9.2 

7.2 

8.8 

Vampyressa  pusilla 


KU  114082  9 

Nicaragua 

31.2 

18.2 

16.5 

10.3 

4.7 

8.1 

5.7 

7.7 

KU  114084  9 

Nicaragua 

30.0 

17.9 

16.2 

10.2 

4.7 

8.5 

5.5 

7.8 

KU  114085  9 

Nicaragua 

31.1 

18.7 

16.7 

10.8 

4.7 

8.3 

6.0 

7.9 

KU  114086  9 

Nicaragua 

30.1 

18.2 

16.5 

10.6 

4.6 

7.8 

6.0 

7.8 

TTU  12894  c? 

Honduras 

29.9 

18.0 

15.9 

10.5 

4.9 

8.2 

5.8 

8.0 

KU  114083  c? 

Nicaragua 

31.9 

18.6 

16.0 

10.5 

4.7 

8.5 

6.0 

7.9 

TTU  9431  c? 

Colombia 

31.9 

18.1 

16.0 

10.5 

4.5 

7.7 

5.5 

7.3 

TTU  9480  <? 

Colombia 

32.5 

18.6 

16.7 

10.5 

4.6 

8.1 

5.7 

7.5 

Vampyrodes  caraccioli 


KU  11  1033  9 

Nicaragua 

53.7 

28.1 

24.3 

17.0 

6.6 

11.6 

9.6 

12.2 

KU  111035  9 

Nicaragua 

53.9 

28.3 

24.8 

17.8 

6.9 

11.8 

9.9 

12.4 

TTU  5288  9 

Trinidad 

49.5 

25.7 

22.7 

16.1 

6.5 

10.8 

9.1 

11.4 

TTU  5355  9 

Trinidad 

49.5 

26.3 

22.6 

16.2 

6.1 

11.0 

9.1 

11.4 

KU  111034  c? 

Nicaragua 

53.3 

28.3 

24.9 

17.8 

6.7 

11.9 

9.7 

12.7 

TTU  5366  c? 

Trinidad 

46.8 

25.9 

22.5 

16.0 

6.2 

10.9 

9.0 

11.2 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


103 


Appendix  1. — Continued. 


TTU  5373  & 

Trinidad 

47.2 

25.8 

22.4 

16.2 

6.2 

10.8 

8.6 

11.3 

TTU  5509  6 

Trinidad 

47.4 

26.0 

22.5 

16.1 

6.0 

11.2 

9.0 

11.5 

Vampyrops 

aurarius 

USNM  387157  9 

Venezuela 

52.0 

28.2 

25.2 

16.0 

6.5 

11.3 

10.6 

12.5 

USNM  387159  9 

Venezuela 

51.9 

28.8 

25.5 

17.0 

6.6 

11.6 

10.8 

13.0 

USNM  387171  9 

Venezuela 

52.5 

29.1 

26.0 

16.8 

6.6 

11.8 

10.5 

12.1 

USNM  387172  9 

Venezuela 

53.4 

29.9 

27.0 

17.8 

6.6 

11.9 

11.0 

13.0 

USNM  387153  6 

Venezuela 

52.4 

28.8 

25.9 

16.5 

6.6 

11.8 

10.6 

12.3 

USNM  387154  <3 

Venezuela 

51.1 

29.5 

26.9 

16.8 

6.8 

11.8 

11.0 

12.8 

USNM  387155  c? 

Venezuela 

51.4 

29.3 

25.9 

16.6 

6.7 

12.2 

10.9 

12.6 

USNM  387161  c? 

Venezuela 

50.4 

28.4 

25.9 

16.8 

6.6 

11.4 

10.5 

12.4 

Vampyrops  brachycephalus 

TCWC  29658  9 

Brazil 

37.2 

20.8 

18.1 

11.8 

5.1 

9.0 

6.8 

8.2 

AMNH  230639  9 

Peru 

36.7 

21.2 

18.5 

12.5 

5.4 

9.3 

7.0 

8.8 

TCWC  12184  9 

Peru 

37.8 

20.8 

18.4 

12.2 

5.1 

9.2 

7.0 

8.6 

TCWC  12185  9 

Peru 

36.9 

20.1 

18.4 

12.5 

5.4 

9.5 

7.0 

8.9 

TCWC  29657  c? 

Brazil 

37.5 

20.9 

18.3 

12.5 

5.5 

9.3 

7.1 

8.8 

TCWC  12177  6 

Peru 

37.5 

20.5 

18.1 

11.8 

5.3 

9.2 

6.9 

8.2 

TCWC  12178  6 

Peru 

36.8 

21.0 

18.3 

12.3 

5.3 

8.9 

7.0 

8.6 

TCWC  12193  c? 

Peru 

40.7 

21.9 

19.2 

13.4 

5.7 

9.6 

7.8 

9.9 

Vampyrops  dorsalis 


AMNH  235778  9 

Colombia 

49.1 

28.3 

25.2 

16.8 

6.5 

11.7 

10.6 

11.5 

AMNH  235779  9 

Colombia 

47.2 

27.2 

24.0 

16.1 

6.8 

11.7 

10.0 

11.4 

AMNH  233614  9 

Peru 

48.0 

26.7 

24.0 

15.4 

6.0 

11.1 

10.4 

11.8 

AMNH  233615  9 

Peru 

50.5 

26.7 

24.2 

16.0 

6.1 

10.8 

10.3 

12.1 

AMNH  233186  6 

Colombia 

50.7 

29.2 

25.9 

17.6 

6.5 

12.3 

11.6 

11.9 

AMNH  233187  c? 

Colombia 

49.6 

28.8 

25.6 

17.0 

6.4 

11.7 

10.4 

11.5 

BMNH  99.12.5.1  c? 

Ecuador 

48.2 

27.4 

25.2 

15.3 

6.3 

10.8 

10.6 

11.5 

AMNH  214356c? 

Peru 

49.3 

28.0 

25.0 

17.0 

7.0 

11.4 

10.5 

13.0 

Vampyrops  helleri 


KU  106131  9 

Nicaragua 

38.0 

22.3 

19.8 

11.9 

5.2 

8.8 

7.5 

8.7 

KU  106133  9 

Nicaragua 

36.3 

21.8 

19.5 

12.1 

5.2 

9.2 

7.9 

9.0 

KU  106134  9 

Nicaragua 

37.1 

21.4 

19.0 

11.8 

5.2 

8.9 

7.3 

8.4 

FHKSC  8839  9 

Colombia 

37.0 

22.5 

20.2 

12.7 

5.3 

9.0 

7.8 

9.0 

FHKSC  9734  c? 

Chiapas 

38.6 

22.8 

20.5 

13.0 

5.5 

9.3 

7.9 

9.3 

KU  106129  c? 

Nicaragua 

37.6 

21.7 

19.3 

12.2 

5.6 

10.0 

7.5 

8.8 

KU  106130  c? 

Nicaragua 

38.5 

21.2 

18.8 

12.1 

5.2 

8.9 

7.4 

8.4 

KU  106131  c? 

Nicaragua 

38.7 

22.3 

20.2 

12.3 

5.5 

9.2 

7.7 

8.9 

Vumpyrops  infuscus 


AMNH  67661  9 

Ecuador 

55.9 

30.6 

27.4 

18.3 

6.3 

12.4 

10.9 

13.8 

AMNH  67664  9 

Ecuador 

55.0 

29.9 

26.2 

17.6 

6.5 

12.0 

11.1 

13.4 

AMNH  236131  9 

Peru 

54.9 

30.5 

27.3 

17.8 

6.5 

12.3 

11.5 

13.1 

AMNH  236132  9 

Peru 

55.0 

30.1 

26.9 

17.6 

6.4 

12.2 

11.3 

12.8 

TTU  9494  c? 

Colombia 

53.4 

29.5 

26.7 

17.5 

6.8 

11.7 

11.6 

13.8 

AMNH  67662  c? 

Ecuador 

55.5 

30.5 

27.3 

17.8 

7.0 

12.2 

11.3 

13.8 

AMNH  67663  c? 

Ecuador 

54.6 

30.9 

27.2 

18.1 

6.8 

12.1 

12.0 

13.7 

AMNH  233729  c ? 

Peru 

56.9 

30.5 

27.3 

18.6 

6.8 

12.4 

11.6 

13.8 

Vampyrops  lineutus 


AMNH  37013  9 

Brazil 

48.5 

25.5 

22.5 

14.3 

6.3 

10.7 

9.1 

10.5 

AMNH  37015  9 

Brazil 

47.6 

25.0 

22.4 

14.6 

6.2 

10.2 

8.8 

10.3 

AMNH  37016  9 

Brazil 

46.2 

24.3 

21.8 

14.1 

6.3 

10.4 

8.3 

10.2 

AMNH  205185  9 

Paraguay 

47.0 

25.4 

22.5 

14.3 

6.4 

10.4 

8.8 

10.2 

AMNH  36995  c? 

Brazil 

46.6 

24.5 

22.0 

13.7 

5.9 

10.3 

9.0 

9.8 

AMNH  148666  c? 

Paraguay 

50.1 

25.0 

22.1 

14.0 

6.3 

10.5 

8.6 

10.2 

AMNH  205184  c? 

Paraguay 

44.6 

24.6 

21.9 

14.1 

6.4 

10.6 

8.7 

10.1 

AMNH  234285  <3 

Paraguay 

45.8 

25.2 

22.2 

14.4 

6.4 

10.7 

8.8 

10.1 

104 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


Vampyrops  nigettus 


AMNH  233686  9 

Peru 

43.9 

25.2 

22.8 

14.2 

6.0 

10.3 

9.2 

10.6 

AMNH  233710  9 

Peru 

44.1 

24.6 

22.0 

13.9 

5.9 

10.4 

8.9 

10.1 

AMNH  233716  9 

Peru 

43.1 

25.0 

22.3 

14.8 

6.0 

10.6 

9.3 

10.7 

AMNH  236106  9 

Peru 

44.4 

25.2 

22.2 

14.4 

6.0 

10.8 

9.0 

10.8 

AMNH  214353  c? 

Peru 

43.2 

24.4 

22.2 

13.9 

5.9 

10.3 

9.0 

10.3 

AMNH  233644  c? 

Peru 

41.1 

24.4 

21.8 

13.5 

5.6 

10.3 

9.0 

10.2 

AMNH  233646  c? 

Peru 

43.5 

25.2 

22.8 

14.0 

5.8 

10.4 

9.2 

10.3 

AMNH  2361 1 1  c? 

Peru 

44.3 

25.3 

22.9 

14.4 

5.9 

10.6 

8.8 

10.1 

Vampyrops  recifinus 

BMNH  93.1.9.15  9 

Brazil 

42.1 

23.7 

21.5 

14.0 

5.7 

10.2 

8.7 

10.2 

BMNH  81.3.16.4c? 

Brazil 

40.6 

24.0 

21.3 

14.0 

5.7 

10.2 

8.7 

10.4 

Vampyrops  vil talus 


TTU  12891  9 

Costa  Rica 

63.7 

34.0 

30.6 

19.8 

7.7 

13.4 

13.0 

14.6 

K.U  93925  9 

Panama 

60.1 

32.7 

29.6 

19.1 

7.6 

13.5 

12.7 

14.0 

TTU  9439  9 

Colombia 

64.3 

33.4 

31.1 

19.9 

7.6 

12.8 

13.6 

15.2 

AMNH  233725  9 

Peru 

57.1 

32.9 

30.1 

20.0 

7.4 

13.5 

13.2 

14.8 

TCWC  10051  c? 

Costa  Rica 

61.5 

32.8 

29.6 

19.5 

7.2 

13.2 

12.5 

14.5 

KU  99355  c? 

Panama 

57.8 

32.4 

28.6 

18.8 

7.4 

13.0 

12.5 

14.3 

AMNH  233718  c? 

Peru 

59.5 

32.8 

29.6 

20.7 

7.3 

13.3 

12.7 

15.5 

AMNH  233728  c? 

Peru 

57.6 

31.6 

28.6 

19.0 

6.6 

12.9 

12.9 

14.7 

Brachyphyllinae 

Brachyphylla  cavernarum 


TTU  20972  9 

Guadeloupe 

66.4 

32.1 

28.7 

17.9 

6.3 

13.1 

1 1.2 

12.3 

TTU  20989  9 

Guadeloupe 

63.3 

30.9 

27.4 

16.6 

6.2 

12.7 

10.7 

11.6 

TTU  20991  9 

Guadeloupe 

64.2 

32.3 

28.8 

17.3 

6.3 

13.0 

11.0 

12.1 

TTU  20995  9 

Guadeloupe 

66.0 

31.0 

27.7 

17.2 

6.5 

12.6 

10.7 

11.6 

TTU  20970  c? 

Guadeloupe 

63.5 

31.1 

27.9 

17.2 

6.5 

12.6 

11.0 

11.7 

TTU  20977  c? 

Guadeloupe 

68.7 

32.6 

29.0 

16.9 

6.3 

12.8 

11.0 

11.6 

TTU  20980  c? 

Guadeloupe 

66.4 

31.8 

28.2 

17.6 

6.5 

12.4 

10.7 

12.1 

TTU  20985  <? 

Guadeloupe 

65.3 

31.1 

27.1 

16.6 

6.6 

12.5 

11.2 

11.7 

Brachyphylla  nana 


AMNH  19085  9 

Cuba 

58.1 

28.0 

24.7 

15.0 

6.0 

11.5 

9.0 

10.0 

AMNH  19090  9 

Cuba 

59.1 

28.9 

25.8 

14.7 

6.2 

11.4 

9.8 

10.5 

TTU  22762  9 

Haiti 

58.8 

28.1 

25.3 

14.9 

6.3 

11.7 

9.4 

9.9 

TTU  22764  9 

Haiti 

58.3 

28.1 

24.8 

14.6 

6.4 

11.9 

9.5 

10.1 

AMNH  214390  c? 

Dominican  Republic 

56.7 

28.2 

25.2 

14.6 

6.5 

11.3 

9.5 

9.8 

AMNH  214393  c? 

Dominican  Republic 

57.9 

28.6 

25.1 

14.5 

6.5 

11.8 

9.4 

9.6 

TTU  22760  c? 

Haiti 

58.0 

28.4 

25.1 

15.5 

6.3 

11.7 

9.5 

10.1 

TTU  22761  c? 

Haiti 

58.8 

28.2 

25.0 

14.7 

6.2 

11.2 

9.5 

9.4 

Erophylla  bombifrons 


AMNH  97591  9 

Dominican  Republic 

47.7 

23.8 

21.7 

11.3 

4.6 

10.0 

7.7 

6.4 

AMNH  212998  9 

Dominican  Republic 

46.6 

23.6 

21.5 

11.1 

4.5 

9.7 

7.5 

6.4 

ROM  45709  9 

Dominican  Republic 

47.1 

24.0 

22.1 

10.8 

4.5 

9.6 

7.6 

6.1 

TTU  22767  9 

Haiti 

46.8 

24.4 

22.5 

1 1.7 

4.5 

10.2 

8.1 

6.7 

ROM  45710  c? 

Dominican  Republic 

45.9 

24.3 

21.9 

11.5 

4.5 

10.0 

7.5 

6.7 

ROM  72710c? 

Dominican  Republic 

49.7 

24.5 

22.3 

11.2 

4.6 

10.1 

8.1 

6.4 

AMNH  39339c? 

Puerto  Rico 

48.8 

24.7 

22.4 

11.6 

4.6 

10.2 

7.8 

6.5 

AMNH  39340  c? 

Puerto  Rico 

48.8 

24.8 

22.5 

11.8 

4.5 

10.3 

7.9 

6.7 

Erophylla  sezekorni 


AS  5814 9 

Jamaica 

47.9 

24.5 

22.6 

11.5 

4.7 

9.7 

8.0 

6.6 

AS  5815  9 

Jamaica 

47.9 

24.7 

22.5 

11.0 

4.1 

9.7 

8.2 

6.3 

AS  5816 9 

Jamaica 

46.5 

24.0 

22.1 

11.0 

4.4 

9.5 

8.0 

6.5 

AS  5817  9 

Jamaica 

49.1 

24.8 

22.7 

11.0 

4.7 

9.7 

8.1 

6.6 

AMNH  45178  c? 

Jamaica 

45.4 

25.7 

23.3 

11.8 

4.7 

10.1 

8.2 

6.8 

AMNH  45179  c? 

Jamaica 

47.8 

25.5 

22.6 

11.3 

4.5 

10.0 

8.0 

6.6 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


105 


Appendix  1. — Continued. 


AMNH  45181  6 

Jamaica 

45.5 

25.3 

22.9 

11.4 

4.5 

10.0 

8.2 

6.6 

AMNH  45 1 82  d 

Jamaica 

45.4 

24.1 

22.9 

11.2 

4.5 

9.7 

7.8 

6.2 

Pliyllonycteris  aphylla 

TTU  21907  9 

Jamaica 

46.0 

24.6 

22.4 

4.9 

9.7 

7.5 

6.7 

TTU  21908  9 

Jamaica 

45.3 

24.9 

22.6 

5.0 

9.9 

7.8 

6.8 

TTU  21913  9 

Jamaica 

45.4 

24.4 

22.4 

5.1 

9.8 

7.8 

6.8 

TTU  21914  9 

Jamaica 

44.8 

23.9 

21.8 

5.0 

9.5 

7.6 

6.8 

TTU  21905  6 

Jamaica 

48.3 

25.8 

23.5 

4.7 

10.0 

8.0 

7.0 

TTU  2 1 906  6 

Jamaica 

44.3 

25.2 

22.8 

4.8 

10.2 

7.9 

6.9 

TTU  21909  d 

Jamaica 

47.6 

24.7 

22.7 

5.2 

9.9 

7.9 

7.0 

TTU  21915  <3 

Jamaica 

46.0 

25.1 

23.1 

5.2 

10.1 

7.9 

6.9 

Pliyllonycteris  major 

AMNH  40925  ? 

Puerto  Rico 

26.3 

24.6 

5.7 

11.4 

8.4 

8.1 

AMNH  40926  ? 

Puerto  Rico 

26.8 

25.1 

5.9 

1  1.1 

8.7 

7.9 

AMNH  40927  ? 

Puerto  Rico 

27.0 

25.2 

5.6 

11.0 

8.5 

7.8 

AMNH  40928  ? 

Puerto  Rico 

25.9 

5.8 

8.8 

8.3 

Pliyllonycteris  poeyi 

USNM  103548  9 

Cuba 

47.6 

24.5 

22.5 

5.4 

10.0 

7.7 

6.9 

USNM  103588  9 

Cuba 

46.5 

23.9 

21.5 

5.2 

10.5 

7.4 

6.8 

USNM  103589  9 

Cuba 

46.2 

23.7 

21.6 

5.3 

10.7 

7.0 

6.8 

USNM  103592  9 

Cuba 

46.6 

24.3 

22.2 

5.3 

10.0 

7.5 

6.9 

USNM  103537  d 

Cuba 

46.9 

25.3 

23.0 

5.7 

10.6 

7.8 

7.3 

USNM  103586  d 

Cuba 

46.5 

24.8 

22.5 

5.4 

10.8 

7.3 

6.8 

USNM  103597  d 

Cuba 

46.9 

25.7 

23.9 

5.3 

10.3 

7.9 

7.4 

USNM  103600.3 

Cuba 

47.1 

24.8 

22.6 

5.4 

10.4 

7.6 

7.2 

Pliyllonycteris  poey 

i  obtusa 

TTU  22783  9 

Haiti 

49.8 

24.2 

22.1 

5.5 

10.2 

7.1 

7.1 

TTU  22792  9 

Haiti 

46.4 

23.9 

21.6 

5,7 

10.9 

7.4 

6.9 

TTU  22793  9 

Haiti 

47.2 

24.0 

22.1 

5.5 

10.8 

7.2 

7.0 

TTU  22794  9 

Haiti 

47.7 

23.7 

22.0 

5.6 

10.0 

7.4 

7.2 

TTU  22772  3 

Haiti 

47.8 

25.2 

22.6 

5.5 

10.5 

7.5 

7.2 

TTU  22773  3 

Haiti 

47.5 

24.8 

22.4 

5.5 

10.4 

7.6 

6.7 

TTU  22782  3 

Haiti 

48.7 

25.4 

22.9 

5.4 

10.5 

7.4 

7.4 

TTU  22783  3 

Haiti 

48.7 

24.7 

22.4 

5.5 

10.2 

7.3 

6.9 

Desmodontinae 

Desmodus  rotund  us 

TTU  8228  9 

Tamaulipas 

60.2 

24.8 

21.4 

11.7 

5.1 

11.6 

3.5 

6.1 

TTU  8170  9 

San  Luis  Potosi 

60.1 

25.0 

21.5 

11.9 

5.2 

11.9 

3.3 

6.0 

KU  111209  9 

Nicaragua 

62.2 

25.2 

21.3 

12.4 

5.6 

12.3 

3.1 

6.1 

KU  111210  9 

Nicaragua 

60.1 

25.0 

21.0 

12.1 

5.4 

11.7 

3.4 

6.5 

TTU  9927  3 

San  Luis  Potosi 

56.9 

24.2 

20.8 

11.4 

5.5 

11.8 

3.4 

5.7 

KU  111204  3 

Nicaragua 

58.6 

24.2 

20.8 

11.8 

5.3 

12.0 

3.4 

6.2 

TTU  5426  3 

Trinidad 

55.3 

23.9 

20.3 

11.5 

5.2 

11.9 

3.5 

5.8 

TTU  5894  3 

Trinidad 

55.5 

23.5 

20.3 

11.7 

5.2 

11.8 

3.5 

5.7 

Diaemus  youngii 

USNM  409368  9 

Venezuela 

53.4 

25.3 

21.7 

13.8 

6.1 

13.2 

3.4 

6.0 

USNM  409374  9 

Venezuela 

54.5 

26.0 

21.7 

14.1 

6.1 

13.2 

3.4 

6.2 

USNM  409375  9 

Venezuela 

53.5 

24.8 

21.1 

14.1 

6.5 

12.6 

3.8 

6.4 

TTU  5232  9 

Trinidad 

51.0 

24.1 

20.4 

13.6 

6.2 

13.0 

3.2 

6.0 

USNM  405767  3 

Venezuela 

51.0 

24.3 

20.2 

13.5 

6.4 

12.9 

3.4 

5.8 

TTU  5233  3 

Trinidad 

51.3 

25.4 

21.5 

14.3 

6.1 

13.1 

3.3 

5.8 

TTU  5411  3 

Trinidad 

49.5 

24.7 

20.7 

13.4 

6.0 

13.0 

3.1 

5.9 

TTU  5428  3 

Trinidad 

50.1 

25.1 

21.2 

14.4 

6.0 

13.5 

3.3 

6.0 

Diphylla  ecaudata 

TTU  5658  9 

Texas 

53.7 

23.5 

20.5 

13.0 

7.6 

11.8 

3.3 

5.9 

TTU  10171  9 

Tamaulipas 

55.3 

23.5 

19.6 

13.0 

7.2 

11.7 

3.4 

6.2 

106 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Appendix  1. — Continued. 


TTU  10157  9 

Veracruz 

55.8 

24.0 

20.1 

12.9 

7.6 

11.6 

3.6 

6.2 

KU  115131  9 

Nicaragua 

56.1 

23.0 

20.0 

12.6 

7.0 

11.1 

3.6 

5.9 

TTU  lOOOOrf 

Veracruz 

54.0 

23.5 

19.9 

12.8 

7.3 

11.8 

3.5 

5.7 

KU  97854  d 

Nicaragua 

56.1 

23.8 

20.6 

13.0 

7.4 

11.6 

3.6 

6.1 

KU  1 15129  <3 

Nicaragua 

55.4 

23.1 

19.7 

12.7 

7.2 

11.2 

3.5 

5.8 

KU  115132  c? 

Nicaragua 

54.7 

23.1 

20.5 

13.1 

7.4 

11.4 

3.5 

6.1 

*  Measurements  as  given  by  Gardner  and  Patton  (1972). 
**  Measurements  as  given  by  Starrett  (1969). 


KARYOLOGY 


Robert  J.  Baker 


This  chapter  is  in  memory  of  Dr.  Claude  M.  Ward,  who  introduced  me  to 
the  world  of  bats  and  whose  premature  death  robbed  me  of  a  good  friend 
and  the  world  of  a  dedicated  educator. 

The  systematics  of  the  New  World  leaf-nosed  bats  are  based  primarily  on 
classical  morphological  features  such  as  shoulder  articulation,  dentition,  and 
other  cranial  features.  The  available  fossil  record  is  inadequate  and  probably 
will  always  be  too  poor  to  determine  much  about  the  evolutionary  relationships 
of  subfamilies  and  genera  (Smith,  1976).  As  an  adjunct  to  the  data  based  on 
classical  morphological  features,  data  from  chromosomal  and  electrophoretic 
studies  are  being  generated  (see  also  Straney  et  al.,  this  volume).  Hopefully,  a 
synthesis  of  the  data  from  these  and  other  works  will  result  in  a  reasonably 
complete  understanding  of  the  systematics  and  genetic  strategies  of  members  of 
the  family  Phyllostomatidae.  Data  derived  from  bat  chromosomes  also  serve  to 
verify,  refute  or  modify  the  proposed  models  of  chromosomal  evolution  (Wilson 
et  al.,  1975;  Bush,  1975). 

In  1966  when  I  first  began  working  with  the  chromosomes  of  this  family,  I 
assumed  that  chromosomal  divergence  in  the  standard  karyotypes  of  species, 
genera,  subfamilies,  and  the  like  generally  would  reflect  their  taxonomic  status 
and  the  evolutionary  time  that  any  two  lineages  had  been  separated.  However,  some 
taxa  (for  instance,  Glossophaga  and  Erophylla )  that  obviously  have  been  sepa¬ 
rated  long  enough  to  evolve  morphological  distinctness  deserving  of  generic  and 
subfamilial  status  had  indistinguishable  karyotypes,  whereas  other  species  (such 
as  Uroderma  bilobatum  and  Choeroniscus  intermedius,  see  also  Rhogeessa, 
Bickham  and  Baker,  1977)  contained  considerable  intraspecific  chromosomal 
divergence.  If  evolutionary  relationships  were  based  solely  on  standard  karyo¬ 
typic  data,  one  would  produce  a  considerably  different  classification  than  that 
currently  derived  from  classical  osteological  and  exomorphological  studies. 
Therefore,  I  began  to  question  the  value  of  chromosomal  divergence  as  a 
taxonomic  indicator.  1  presently  am  opposed  to  placing  too  much  emphasis  on 
degree  of  gross  karyotypic  divergence  as  a  justification  for  taxonomic  status 
(with  the  possible  exception  of  specific  distinctness).  Of  course,  the  longer  two 
lineages  have  been  separated,  the  more  probable  it  is  that  events  have  occurred 
that  result  in  karyotypic  divergence.  However,  karyotypic  changes  become 
established  in  a  species  at  such  irregular  intervals  that  one  cannot  depend  on  the 
rate  of  their  establishment  to  indicate  taxonomic  position. 

John  Bickham  and  I  are  preparing  a  manuscript  in  which  we  propose  that  the 
rate  and  magnitude  of  chromosomal  change  is  primarily  a  function  of  the  degree 
to  which  the  karyotype  is  adaptive  to  the  adaptive  zone  occupied  by  the  organism. 


107 


108 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


If  this  model  proves  accurate  then,  at  times,  organisms  would  undergo  relatively 
rapid  chromosomal  evolution  and  at  other  times  there  would  be  long  periods  of 
reduced  rates  of  chromosomal  change. 

The  fact  that  karyotypic  changes  do  not  evolve  at  a  constant  rate  is  not  too 
startling  if  one  realizes  it  is  a  well  documented  fact  that  morphological  features 
also  evolve  at  different  rates.  In  a  given  taxon,  some  features  can  become  highly 
derived  from  the  ancestral  condition,  whereas  others  remain  indistinguishable 
from  the  primitive  condition.  Meanwhile,  in  a  closely  related  taxon,  a  different 
suite  of  characters  can  become  derived  whereas  all  other  characters  remain  near 
the  primitive.  If  greater  emphasis  is  placed  on  the  derived  characters,  the  sys- 
tematics  would  result  in  greater  taxonomic  distance  than  if  the  classification  were 
based  only  on  the  characteristics  that  remained  in  the  primitive  condition.  A 
similar  case  might  be  made  for  the  degree  of  morphological  divergence — it  does 
not  necessarily  reflect  the  evolutionary  history.  Certainly,  parallelism  and  con¬ 
vergence  can  result  in  incorrect  “lumping,”  and  yet,  emphasis  on  most  rapidly 
evolving  features  may  result  in  oversplitting.  However,  the  fossil  record  reveals 
that  generally  there  is  agreement  between  total  morphological  divergence  and 
evolutionary  history.  In  light  of  data  from  the  fossil  record,  I  believe  that  in  the 
majority  of  cases  an  overview  of  classical  morphological  data  gives  a  more 
reasonable  and  accurate  reflection  of  the  evolutionary  history  than  does  degree 
of  chromosomal  divergence. 

On  the  other  hand,  there  are  cases  where  karyotypic  data  can  be  more  valuable 
than  general  morphology.  To  a  much  greater  extent  than  general  morphological 
information,  G-band  chromosomal  data  are  applicable  to  the  cladistic  methodol¬ 
ogies  of  Hennig  (1966).  The  likelihood  of  extensive  convergence  of  G-banding 
patterns  is  sufficiently  low  to  warrant  placing  considerable  confidence  in  the 
data.  The  typical  mammalian  genome  is  arranged  in  such  a  manner  that  there  are 
enough  chromosomal  arms  (linkage  groups)  to  provide  an  adequate  number  of 
data  points  to  determine  the  relationships  within  complex  taxa.  Additionally, 
G-band  chromosomal  characteristics  are  independent  of  exomorphological, 
cranial,  or  osteological  features  and,  therefore,  serve  as  an  independent  data 
source.  A  synthesis  of  findings  from  all  of  the  aforementioned,  plus  those  of  a 
biochemical  nature  (such  as  electrophoretic,  immunological,  and  DNA  hy¬ 
bridization),  should  give  the  most  accurate  interpretation  of  the  phylogeny  and 
systematics  of  a  taxon.  Also,  data  from  these  three  sources  (general  morphology, 
karyology,  and  biochemical)  will  be  necessary  to  understand  the  evolutionary 
strategy  of  major  taxa. 

Of  the  137  phyllostomatid  species  recognized  by  Jones  and  Carter  (1976), 
basic  karyotypic  data  are  available  for  105  (Table  1).  In  addition,  Gardner  (1977) 
reported  karyotypic  data  for  two  additional  taxa,  Artibeus  fuliginosus  and  A. 
planirostris ,  which  were  not  recognized  by  Jones  and  Carter  (1976).  Rep¬ 
resentative  standard  karyotypes  for  60  species  are  presented  in  Plates  1  through 
60,  which  follow  the  literature  cited.  I  have  attempted  to  illustrate  the  major 
chromosomal  complements  found  in  the  Phyllostomatidae.  Plates  are  arranged 
alphabetically  by  generic  and  species  names  within  subfamilies:  Phyllostomatinae, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


109 


Table  1. — Chromosomal  data  for  phyllostomatid  hats.  Subfamilies  are  arranged  in  the 
order  followed  by  Jones  and  Carter  (1976).  Genera  and  species  are  in  alphabetical  order. 
Symbols  are  2 n,  diploid  number;  FN,  Fundamental  Number;  M,  metacentric;  SM,  sub- 
metacentric;  ST,  subtelocentric;  A,  acrocentric.  Two  species  names  not  recognized  by  Jones 
and  Carter  (1976)  are  identified  by  an  asterisk.  Mesophylla  is  recognized  as  distinct  from 

Ectophylla. 


Number  of 


Taxon 

In 

FN 

X 

Y 

Y2  Authority 

specimens 

Phyllostomatinae 

Chrotopterus  auritus 

28 

52 

SM 

A 

Yonenaga,  1968; 

i 

28 

52 

SM 

A 

Yonenaga  et  al.,  1969 

i 

Lonchorhina  aurita 

32 

M 

A 

Baker  and  Hsu,  1970 

2 

32 

60 

M 

A 

Baker,  1973 

Lonchorhina  orinocensis 

No  information 

Macro phyll um  macro phyllum 

No  information 

Macrotus  californicus 

40 

Kniazeff  et  al.,  1967 

40 

60 

SM 

SM 

Nelson-Rees  et  al.,  1968 

10 

40 

60 

Davis  and  Baker,  1974 

155 

40 

60 

Greenbaum  and  Baker,  1976 

100 

Macrouts  waierhousii 

46 

60 

SM 

A 

Baker,  1967;  Hsu  et  al.,  1968 

5 

46 

60 

SM 

A 

Nelson-Rees  et  al.,  1968 

7 

46 

60 

SM 

A 

Davis  and  Baker,  1974 

44 

46 

60 

Nagorsen  and  Peterson,  1975 

4 

46 

60 

M 

A 

Greenbaum  and  Baker,  1976 

118 

46 

60 

SM 

A 

Patton,  1976 

2 

Micronycteris  behni 

No  information 

Micronycteris  brachyotis 

32 

60 

SM 

Patton,  1976 

i 

Micronycteris  daviesi 

No  information 

Micronycteris  hirsuta 

28 

32 

A 

A 

Baker,  1973 

30 

32 

A 

A 

Baker  et  al.,  1973 

7 

28 

32 

A 

A 

Baker  et  al.,  1973 

4 

Micronycteris  megalot  is 

40 

68 

ST 

A 

Baker,  1967;  Hsu  et  al.,  1968 

1 

40 

68 

SM 

A 

Patton,  1976 

1 

Micronycteris  minuta 

28 

50 

ST 

A 

Baker,  1973 

28 

50 

SM 

Patton,  1976 

1 

Micronycteris  nicefori 

28 

M 

A 

Baker  and  Hsu,  1970 

5 

28 

52 

SM 

Patton,  1976 

1 

Micronycteris  pits  ilia 

No  information 

Micronycteris  schmidtorum 

38 

66 

ST 

A 

Baker,  1973 

Micronycteris  sylvestris 

No  information 

Mimon  bennettii 

No  information 

Million  cozumelae 

34 

56 

Patton,  1976 

1 

Mimon  crenulatum 

32 

Baker  and  Hsu,  1970 

2 

32 

60 

M 

M 

Baker  et  al.,  1 972 /z 

20 

32 

60 

SM 

A 

Hsu  and  Benirschke,  1974 

32 

60 

SM 

A 

Gardner,  1977 

32 

60 

SM 

M 

Patton,  1976 

1 

Mimon  koepckeae 

32 

60 

SM 

A 

Gardner,  1977 

Pbylloderma  stenops 

32 

58 

Baker  and  Hsu,  1970 

1 

32 

58 

M 

A 

Baker,  1973 

Phyllostom us  d i scalar 

32 

60 

SM 

A 

Baker,  1967;  Hsu  et  al.,  1968 

4 

32 

9 

9 

Yonenaga,  1968 

1 

32 

60 

M 

A 

Kiblisky,  1969 

4 

32 

9 

9 

Yonenaga  et  al.,  1969 

1 

32 

SM 

A 

Baker  and  Hsu,  1970 

2 

32 

60 

SM 

A 

Baker,  1970 

1 

32 

60 

SM 

Patton,  1976 

1 

Phyllostomus  elongatus 

32 

58 

SM 

A 

Baker,  1973 

Phyllostomus  hastatus 

32 

58 

SM 

A 

Yonenaga,  1968 

5 

32 

58 

SM 

A 

Yonenaga  et  al.,  1969 

5 

32 

58 

M 

A 

Kiblisky,  1969 

7 

32 

SM 

A 

Baker  and  Hsu,  1970 

2 

32 

58 

SM 

A 

Patton,  1976 

2 

110 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1. —  Continued. 


Phyllostomus  lutifolius 

No  information 

Tonatia  bidens 

16 

20 

M 

A 

Baker  and  Hsu,  1970 

3 

16 

20 

SM 

Patton,  1976 

2 

Tonatia  brasiliensis 

30 

56 

ST 

A 

Gardner,  1977 

Tonatia  carrikeri 

26 

46 

Gardner,  1977 

Tonatia  minuta 

30 

SM 

A 

Baker  and  Hsu,  1970 

3 

30 

56 

SM 

A 

Baker,  1973 

30 

56 

SM 

Patton,  1976 

1 

Tonatia  silvicola 

34 

60 

SM 

A 

Gardner,  1977 

Tonatia  venezuelae 

30 

56 

This  paper 

1 

Trachops  cirrhosus 

30 

56 

ST 

A 

Baker,  1967;  Hsu  et  al.,  1968 

4 

Vampyrum  spectrum 

30 

56 

Baker  and  Hsu,  1970 

1 

30 

56 

SM 

A 

Baker,  1973 

Glossophaginae 

Anoura  brevirostrum 

No  information 

Anoura  caudifer 

30 

Yonenaga,  1968 

30 

56 

SM 

A 

Baker,  1973 

Anoura  cultrata 

30 

56 

SM 

A 

This  paper 

1 

Anoura  geoffroyi 

30 

56 

SM 

A 

Baker,  1967;  Hsu  et  al.,  1968 

3 

30 

SM 

A 

Baker  and  Hsu,  1970 

3 

SM 

Pathak  and  Stock,  1974 

Anoura  werckleae 

No  information 

Choeroniscus  godmani 

19 

32 

SM 

ST 

A  Baker,  1967 

5 

19 

Hsu  et  al.,  1968 

5 

19 

32 

SM 

A 

A  Baker,  1970a 

1 

20 

36 

SM 

Patton  and  Gardner,  1971 

1 

20 

36 

This  paper 

2 

Choeroniscus  inca 

No  information 

Choeroniscus  intermedius 

20 

36 

Baker,  1970a 

20 

Baker,  1973 

SM 

Pathak  and  Stock,  1974 

1 

20 

36 

SM 

A 

Stock,  1975 

1 

Choeroniscus  minor 

No  information 

Choeroniscus  periosus 

No  information 

Choeronycteris  mexicana 

16 

24-26 

Baker,  1967;  Hsu  et  al.,  1968 

1 

16 

24 

SM 

SM 

Baker,  1973 

Glossophaga  ulticola 

32 

60 

M 

A 

Baker,  1967 

4 

Glossophaga  commissarisi 

32 

60 

M 

A 

Baker,  1967;  Hsu  et  al.,  1968 

5 

Glossophaga  longirostris 

32 

60 

M 

A 

This  paper 

Glossophaga  soricina 

32 

60 

M 

A 

Baker,  1967;  Hsu  et  al.,  1968 

14 

32 

M 

A 

Baker  and  Hsu,  1970 

4 

32 

60 

SM 

A 

Baker,  1970a 

1 

Hylonycteris  underwoodi 

16 

24 

Baker,  1973 

Leptonycteris  curasoae 

No 

information 

Leptonycteris  sanborni 

32 

60 

M 

A 

Baker,  1967;  Hsu  et  al.,  1968 

5 

Leptonycteris  nivalis 

32 

60 

Baker,  1973 

Lichonycteris  degener 

No  information 

Lichonycteris  obscura 

28 

50 

SM 

A 

Baker,  1973  (data  incorrect) 

1 

24 

44 

This  paper 

2 

Lionycteris  spurred i 

28 

50 

SM 

A 

This  paper 

1 

Lonchophylla  concava 

No  information 

Lonchophylla  hesperia 

No  information 

Lonchophylla  mordax 

No  information 

Lonchophylla  robusta 

28 

50 

SM 

A 

Baker,  1973 

Lonchophylla  thomasi 

30 

34 

Baker,  1973 

32 

38 

Gardner,  1977 

Monophyllus  plethodon 

32 

60 

SM 

A 

This  paper 

3 

Monophyllus  redmani 

32 

60 

SM 

A 

Baker  and  Lopez,  19706 

7 

Musonycteris  harrisoni 

No  information 

Platalina  genovensium 

No  information 

Scleronycteris  ega 

No  information 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


Ill 


Table  1. —  Continued. 


Carolliinae 


CaroIIia  brevicauda 

20-21 

36 

ST 

A 

A 

Patton  and  Gardner,  1971 

4 

20-21 

36 

Stock,  1975 

1 

CaroIIia  casianea 

20-21 

36 

ST 

A 

A 

Baker  and  Bleier,  1971 

4 

22 

38 

SM 

A 

Patton  and  Gardner,  1971  (Peru) 

5 

20-21 

36 

ST 

Patton  and  Gardner,  1971 

1 

(Costa  Rica) 

22 

38 

SM 

A 

Hsu  and  Bernischke,  1973 

20-21 

36 

SM 

Pathak  and  Stock,  1974 

22 

38 

SM 

A 

Stock,  1975 

1 

22 

38 

SM 

A 

Hsu  et  al.,  1975 

1 

CaroIIia  perspicillata 

20-21 

36 

ST 

A 

A 

Baker,  1967 

2 

21 

Hsu  et  al.,  1968 

2 

20-21 

36 

SM 

A 

A 

Yonenaga,  1968 

4 

20-21 

36 

ST 

A 

A 

Kiblisky,  1969 

3 

20-21 

36 

SM 

A 

A 

Yonenaga  et  al.,  1969 

4 

20 

36 

ST 

A 

A 

Baker,  1970  a,  19706 

1 

20-21 

36 

ST 

A 

A 

Baker  and  Hsu,  1970 

4 

20-21 

36 

SM 

A 

A 

Baker  and  Bleier,  1971 

5 

20-21 

36 

ST 

A 

A 

Patton  and  Gardner,  1971 

7 

20-21 

36 

SM 

A 

A 

Pathak  and  Stock,  1974 

20-21 

36 

SM 

A 

A 

Hsu  et  al.,  1975 

2 

20-21 

36 

ST 

A 

A 

Stock,  1975 

1 

CaroIIia  subrufa 

20-21 

36 

ST 

A 

A 

Baker,  1967 

12 

20-21 

Hsu  et  al.,  1968 

11 

20 

36 

ST 

Baker,  1970a,  19706 

1 

20-21 

36 

ST 

A 

A 

Baker  and  Bleier,  1971 

2 

Rhinophylla  alethina 

No  information 

RJi  inophylla  fischerae 

34 

56 

SM 

A 

Baker  and  Bleier,  1971 

1 

RJi  inophylla  pumilio 

36 

62 

M 

A 

Baker  and  Bleier,  1971 

6 

36 

62 

M 

SM 

Hsu  and  Benirschke,  1973 

Stenoderminae 

Ametrida  centurio 

30-31 

ST 

SM 

M 

Baker  and  Hsu,  1970 

5 

Ardops  nichollsi 

30-31 

56 

SM 

ST 

A 

Greenbaum  et  al.,  1975 

10 

Ariteus  flavescens 

30-31 

56 

ST 

ST 

A 

Greenbaum  et  al.,  1975 

12 

Artibeus  aztecus 

30-31 

56 

ST 

A 

A 

Baker,  1973 

Artibeus  cinereus 

30-31 

56 

ST 

SM 

M 

Baker  and  Hsu,  1970 

4 

Arlibeus  concolor 

No  information 

Arlibeus  ful  ig  ino  su  s* 

30-31 

56 

ST 

A 

A 

Gardner,  1977 

Artibeus  glaucus 

30-31 

56 

ST 

A 

A 

Gardner,  1977 

Artibeus  hirsutus 

30-31 

56 

ST 

ST 

A 

Baker,  1973 

Artibeus  inopinatus 

30-31 

56 

ST 

ST 

A 

This  paper 

5 

Artibeus  jamaicensis 

30-31 

56 

ST 

A 

A 

Baker,  1967 

15 

30-31 

56 

Hsu  et  al.,  1968 

9 

30-31 

56 

ST 

A 

A 

Kiblisky,  1969 

2 

30-31 

ST 

A 

A 

Baker  and  Hsu,  1970 

3 

30-31 

56 

ST 

A 

A 

Baker  and  Lopez,  19706 

5 

Artibeus  lituratus 

30-31 

56 

ST 

A 

A 

Baker,  1967 

8 

30-31 

Hsu  et  al.,  1968 

8 

30-31 

56 

SM 

A 

A 

Yonenaga,  1968 

2 

30-31 

56 

SM 

A 

SM 

Becak  et  al.,  1969 

4 

30-31 

56 

ST 

A 

A 

Kiblisky,  1969 

3 

30-31 

56 

SM 

A 

A 

Yonenaga  et  al.,  1969 

2 

30-31 

ST 

ST 

A 

Baker  and  Hsu,  1970 

2 

SM 

Pathak  and  Stock,  1974 

Artibeus  phaeotis 

30 

56 

ST 

SM 

Baker,  1967 

4 

30 

56 

ST 

SM 

Hsu  et  al.,  1968 

2 

Artibeus  planirostris* 

30-31 

56 

ST 

A 

A 

Gardner,  1977 

Artibeus  toltecus 

30-31 

56 

ST 

A 

A 

Baker,  1967 

4 

30-31 

56 

ST 

A 

A 

Hsu  et  al.,  1968 

4 

Artibeus  watsoni 

30 

56 

ST 

SM 

Baker,  1973 

112 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1. —  Continued. 


Centurio  sen  ex 

28 

52 

Baker,  1967;  Hsu  et  al.,  1968 

1 

28 

ST 

SM 

Baker  and  Hsu,  1970 

1 

Chiroderma  doriae 

No  information 

Chiroderma  im provision 

26 

48 

ST 

ST 

Baker  and  Genoways,  1976 

1 

Chiroderma  salvini 

26 

48 

ST 

SM 

Baker,  1973 

Chiroderma  trinit  atom 

26 

ST 

SM 

Baker  and  Hsu,  1970 

3 

26 

48 

ST 

ST 

Baker  and  Genoways,  1976 

26 

48 

Gardner,  1977 

Chiroderma  villosum 

26 

48 

ST 

SM 

Baker,  1967;  Hsu  et  al.,  1968 

3 

26 

ST 

SM 

Baker  and  Hsu,  1970 

4 

SM 

Pathak  and  Stock,  1974 

26 

48 

Gardner,  1977 

Ectophylla  alba 

30 

56 

SM 

A 

Greenbaum  et  al.,  1975 

1 

30 

56 

This  paper 

Enchisthenes  hartii 

30 

56 

Baker,  1967;  Hsu  et  al.,  1968 

2 

30-31 

ST 

SM 

A 

Baker  and  Hsu,  1970 

2 

Me sophy lla  macconnelli 

21-22 

A 

Baker  and  Hsu,  1970 

16 

21 

Baker  and  Hsu,  1970 

1 

21-22 

20 

A 

Hsu  and  Benirschke,  1971 

21-22 

20 

A 

Baker  et  al.,  1973 

27 

Phyllops  falcatus 

No  information 

Phyllops  haitiensis 

30-31 

56 

ST 

ST 

A 

Greenbaum  et  al.,  1975 

8 

30 

56 

Nagorsen  and  Peterson,  1975 

3 

Pyyoderma  bilab  iatum 

No  information 

Sphaeronvcteris  toxophyllum 

28 

52 

ST 

SM 

Baker,  1973 

Stenoderma  rufum 

30-31 

56 

ST 

A 

A 

Baker  and  Lopez,  1 970/z 

16 

30-31 

56 

ST 

A 

A 

Genoways  and  Baker,  1972 

16 

Sturnira  aratathomasi 

No  information 

Sturnira  bidens 

30 

56 

ST 

A 

Gardner  and  O’Neill,  1969 

2 

Sturnira  erythromos 

30 

56 

ST 

A 

Gardner  and  O’Neill,  1969 

6 

Sturnira  lilium 

30 

56 

ST 

SM 

Baker,  1967;  Hsu  et  al.,  1968 

15 

30 

56 

ST 

SM 

Kiblisky,  1969 

3 

30 

56 

ST 

SM 

Baker  and  Hsu,  1970 

4 

Sturnira  ludovici 

30 

56 

ST 

SM 

Baker,  1967;  Hsu  et  al.,  1968 

2 

30 

56 

Kiblisky,  1969 

1 

Sturnira  mayna 

30 

56 

ST 

A 

Gardner,  1977 

Sturnira  mordax 

30 

56 

Baker,  1973 

Sturnira  nana 

30 

56 

ST 

A 

Gardner,  1977 

Sturnira  thomasi 

30 

56 

This  paper 

Sturnira  tildae 

30 

ST 

SM 

Baker  and  Hsu,  1970 

3 

Uroderma  b  Hob  atom 

44 

48 

ST 

SM 

Baker,  1967;  Hsu  et  al.,  1968 

4 

42 

ST 

SM 

Baker  and  Hsu,  1970 

3 

38 

44 

ST 

SM 

Baker  and  Lopez,  1970a 

5 

42 

50 

ST 

SM 

Baker  and  Lopez,  1970a 

13 

42 

50 

SM 

SM 

Hsu  and  Benirschke,  1971 

44  or  43 

48 

ST 

SM 

O 

Baker  and  McDaniel,  1972 

122 

38 

44 

SM 

M 

Baker  et  al.,  1972 

39 

45 

Baker  et  al.,  1972 

total  of  144 

44  or  43 

48 

SM 

M 

Baker  et  al.,  1972 

38 

Baker  et  al.,  1975 

88 

39 

Baker  et  al.,  1975 

4 

40 

Baker  et  al.,  1975 

1 

41 

Baker  et  al.,  1975 

1 

42 

Baker  et  al.,  1975 

1 

43 

Baker  et  al.,  1975 

14 

44 

Baker  et  al.,  1975 

82 

Uroderma  magnirostrum 

36 

62 

ST 

SM 

Baker  and  Lopez,  1970a 

II 

35 

62 

ST 

SM 

Baker  and  Lopez,  1970a 

2 

36 

60 

SM 

M 

Hsu  and  Benirschke,  1971 

Vampyressa  bidens 

26 

48 

Gardner,  1977 

Vampyressa  brocki 

24 

44 

Baker  and  Genoways,  1972 

3 

24 

44 

Baker  et  al.,  1973 

3 

24 

44 

ST 

Gardner,  1977 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


113 


Table  1. —  Continued. 


Vampyressa  melissa 

14 

24 

ST 

Gardner,  1977 

Vampyressa  nymphuea 

26 

48 

ST 

SM 

Baker,  1973 

26 

48 

ST 

SM 

Baker  et  al. ,  1973 

26 

48 

ST 

A 

Gardner,  1977 

5 

Vampyressa  pusilla 

23-24 

22 

ST 

Baker,  1973 

18 

20 

ST 

SM 

Baker,  1973 

18 

20 

ST 

ST 

Baker  et  al.,  1973 

13 

24-23 

22 

Baker  et  al.,  1973 

9 

22-23 

22 

ST-A 

SM 

Gardner,  1977 

Vam pyrod es  caraceiol  i 

30 

ST 

SM 

Baker  and  Hsu,  1970 

4 

30 

56 

ST 

SM 

Baker,  1973 

Vampyrops  aurarius 

No 

information 

Vampyrops  brachycephalus 

30 

56 

ST 

SM 

Baker,  1973 

Vampyrops  dorsalis 

30 

56 

ST 

SM 

Baker,  1973 

Vampyrops  helleri 

30 

56 

ST 

SM 

Baker,  1967;  Hsu  et  al.,  1968 

2 

30 

ST 

SM 

Baker  and  Hsu,  1970 

4 

Vampyrops  infuscus 

30 

56 

ST 

A 

Gardner,  1977 

Vampyrops  lineatus 

No 

information 

Vampyrops  nigellus 

30 

56 

ST 

A 

Gardner,  1977 

Vampyrops  recifinus 

No 

information 

Vampyrops  vittatus 

30 

56 

ST 

A 

Baker,  1973 

Brachyphyllinae 

Brachyphylla  cavernarum 

32 

60 

SM 

A 

Baker  and  Lopez,  19706 

11 

Brachyphylla  nana 

32 

60 

SM 

A 

This  paper 

3 

Brachyphylla  pumila 

32 

60 

SM 

A 

Nagorsen  and  Peterson,  1975 

4 

Erophylla  sezekorni 

32 

60 

Baker  and  Lopez,  19706 

11 

32 

60 

SM 

A 

Nagorsen  and  Peterson,  1975 

4 

Phyllonycteris  aphylla 

32 

60 

SM 

A 

This  paper 

Phyllonycteris  major 

No 

information 

Phyllonycteris  ohtusa 

32 

60 

SM 

A 

Nagorsen  and  Peterson,  1975 

1 

32 

60 

SM 

A 

This  paper 

Phyllonycteris  poeyi 

No 

information 

Desmodontinae 

Desmodus  rotundas 

28 

52 

SM  A 

&  ST 

Forman  et  al.,  1968 

13 

28 

52 

Yonenaga  et  al.,  1969 

6 

28 

52 

SM 

ST 

Cadena  and  Baker,  1976 

Diaemus  younyii 

32 

60 

SM 

A 

Forman  et  al.,  1968 

4 

32 

60 

Cadena  and  Baker,  1976 

1 

Diphylla  ecaudata 

28 

52 

SM 

A 

Baker,  1973  (data  incorrect) 

32 

60 

Cadena  and  Baker,  1976 

2 

32 

60 

Gardner,  1977 

Plates  1  to  17;  Glossophaginae,  18  to  29;  Carolliinae,  30  to  32;  Stenoderminae, 
33  to  52;  Phyllonycterinae,  53  to  57;  Desmodontinae,  58  to  60. 

Determination  of  Primitive  Karyotype 

One  very  important  point  of  information  relative  to  determining  evolutionary 
events  and  their  systematic  implications  is  an  understanding  of  the  primitive 
versus  the  derived  condition.  Because  there  is  no  fossil  record  for  karyotypes, 
primitive  cytogenetic  aspects  are  difficult  to  ascertain. 

Prior  to  the  availability  of  G-band  data,  two  theories  were  developed  as  to  the 
diploid  and  fundamental  characteristics  of  the  primitive  karyotype  for  the 
family  Phyllostomatidae.  Baker  (1967,  1973)  proposed  that  the  primitive  karyo¬ 
type  for  the  Phyllostomatidae  consisted  of  a  diploid  number  (2 n)  of  30  or  32, 
with  a  fundamental  number  (FN)  of  56  to  60.  This  theory  was  based  on  the 


114 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


widespread  occurrence  of  the  2/7  =  30  or  32,  FN  =  56-60  karyotype  among  species 
from  the  different  subfamilies;  the  alternative  explanation  was  to  assume  the 
condition  arose  through  convergent  evolution.  Gardner  (1977),  on  the  other  hand, 
proposed  that  the  primitive  karyotype  was  2/7  =  36  to  40  with  an  FN  near  the 
minimum  for  this  diploid  number,  38  or  slightly  higher.  The  significant  difference 
between  the  two  theories  centers  around  the  types  of  chromosomal  rearrange¬ 
ments  required  to  derive  the  karyotypes  found  in  extant  species.  The  2/7  =  30  or  32, 
FN  =  60,  would  require  terminalization  of  centromeres  by  pericentric  inversion  or 
centric  transpositions  in  addition  to  translocations  (especially  centric  fusions)  as 
theprimary  rearrangements,  whereasthe2/7  =  36-40,  FN  =  38,wouldrequirecentral- 
ization  of  centromeres  (by  pericentric  inversion  or  centric  transpositions)  in  addi¬ 
tion  to  some  fusions. 

It  is  of  interest  to  note  that  when  Gardner  (1977:314-315)  interpreted  phy¬ 
logenetic  relationships  within  the  family  based  on  chromosomal  evolution  from 
a  primitive  karyotypic  condition  of  a  higher  diploid  number  (about  40)  and  a 
lower  fundamental  number  (about  38),  his  three  “major  deviations  from  more 
classical  portrayals”  were  essentially  those  proposed  earlier  based  on  a  primitive 
2/j  =  32,  FN  =  60,  karyotype.  Relative  to  Gardner’s  case  1,  Baker  and  Lopez 
(19706:471)  pointed  out  the  “possibility  of  a  close  phylogenetic  relationship” 
of  the  phyllonycterine  genera  to  Monophyllus.  In  Gardner’s  case  2,  Baker  (1967: 
423),  basing  his  remarks  on  karyotypes,  not  only  suggested  that  Sturnira  “must 
have  evolved  from  the  Stenoderminae  complex,”  he  also  regarded  the  two  sub¬ 
families  as  synonymous,  which  is  the  systematic  relationship  followed  by  Jones 
and  Carter  (1976:20).  In  case  3,  Greenbaum  etal.  (1975)  suggested  the  recognition 
of  Mesophylla  as  generically  distinct  from  Ectophylla.  The  point  is  that  even 
though  a  2/7  =  40,  FN  =  38,  primitive  karyotype  theory  might  be  a  viable  alternative 
to  the  2/7  =  32,  FN  =  60,  theory  in  several  examples,  the  systematic  implications 
of  the  chromosomal  data  are  the  same. 

With  data  from  G-bands,  it  became  possible  to  identify  homologous  segments 
between  variant  karyotypes  even  at  the  subfamilial  level  (Mascarello  et  al.,  1974), 
and  G-band  studies  became  the  means  for  testing  these  two  theories.  It  could  be 
predicted  that  if  the  theory  of  2/7  =  30  or  32,  FN  =  56-60,  were  true,  there  should  be 
considerable  homology  of  banding  patterns  between  the  two  arms  of  the  supposed 
homologous  elements  of  the  2/7  =  30  or  32,  FN  =56  or  60  karyotypes  within  the 
family,  and  although  some  elements  in  each  karyotype  may  have  been  rear¬ 
ranged,  the  same  pairs  should  not  always  be  affected.  On  the  other  hand,  if  the 
2/7  =  40,  FN  =  38  (Gardner,  1977)  karyotype  proved  primitive,  G-banding  pat¬ 
terns  of  biarmed  elements  of  the  2/7=32  karyotypes  from  separate  subfamilies 
should  show  little  homology  between  the  subfamilies.  Therefore,  G-banding 
homology  among  these  karyotypes  with  lower  fundamental  numbers  from 
different  subfamilies  would  be  strong  proof  in  favor  of  Gardner’s  theory. 

Patton  (1976)  examined  G-banded  chromosomes  of  five  genera  (involving 
10  species)  of  the  subfamily  Phyllostomatinae  as  well  as  one  species  from  the 
families  Mormoopidae  ( Pteronotus  parnellii)  and  Noctilionidae  ( Noctilio 
albiventris).  His  results  indicated  that  the  FN  =  60  was  primitive  for  the  Phyl- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


115 


lostomatinae  as  well  as  for  the  Mormoopidae  and  Noctilionidae.  Macrotus  (as 
well  as  several  other  species  of  the  Phyllostomatinae),  Pteronotus,  and  Noctilio 
all  have  30  pairs  of  autosomal  arms.  When  the  G-banded  karyotypes  of  these  three 
genera  are  compared,  the  thirty  homologous  arms  found  in  the  karyotype  of  each 
genus  have  a  distinguishable  counterpart  in  the  karyotypes  of  the  other  two  genera. 
The  most  logical  interpretation  of  these  data  is  that  the  number  of  autosomal 
arms  in  the  karyotype  of  the  common  ancestor  of  Macrotus,  Pteronotus,  and 
Noctilio  was  30  pairs  (FN  =  60),  which  have  retained  their  respective  G-banding 
patterns  since  their  separation  from  a  common  ancestor.  The  alternative  ex¬ 
planation,  that  the  G-band  similarity  between  the  representatives  of  these  three 
families  is  the  result  of  the  evolution  of  convergent  G-banding  patterns  in  the  exact 
same  number  of  pairs  (30)  of  autosomal  arms,  is  less  plausible  (Patton,  1976). 
Additionally,  data  from  the  G-banded  karyotypes  of  other  taxa  thus  far  studied 
(by  Patton,  1976,  and  unpublished  data  including  representatives  of  the  Des- 
modontinae,  Glossophaginae,  and  Stenoderminae)  support  the  conclusion  that 
the  FN  =  60  was  primitive  for  the  Phyllostomatidae.  Derivation  of  the  various 
karyotypes  of  the  taxa  studied  from  any  of  the  karyotypes  with  the  more  aberrant 
fundamental  numbers  (such  as  Tonatia  bidens  FN  =  20  or  Micronycteris  megalotis 
FN  =  68)  would  require  many  convergent  chromosomal  rearrangements  in  order 
to  avoid  concluding  that  Macrotus  was  more  closely  related  to  the  mormoopids 
and  noctilionids  than  to  the  other  phyllostomatids. 

The  primitive  diploid  number  for  the  Phyllostomatidae  was  believed  to  be 
2u  =  46  (Patton,  1976).  The  following  discussion,  modified  from  Patton’s  thesis, 
points  out  the  reasons  for  this  conclusion. 

A  diploid  number  of  46  (with  16  biarmed  autosomes,  28  acrocentric  autosomes, 
plus  two  sex  elements)  is  most  probably  like  the  primitive  condition  (Patton, 
1976).  Essentially,  this  is  the  karyotype  of  Macrotus  waterhousii  (Fig.  1).  Data 
supporting  this  conclusion  are  the  eight  pairs  of  biarmed  elements  found  in  the 
karyotype  of  Macrotus  that  have  corresponding  biarmed  elements  in  the  karyo¬ 
type  of  Noctilio.  Seven  of  these  eight  pairs  are  present  also  in  Pteronotus,  Tonatia 
minuta,  Mimon  crenulatum,  Phyllostomus  discolor,  and  Phyllostomus  hastatus. 
The  majority  of  these  eight  pairs  are  identifiable  in  most  of  the  karyotypes  of  other 
phyllostomatine  species  studied.  Therefore,  it  is  likely  that  these  eight  biarmed 
pairs  were  primitive  for  the  phyllostomatoid  karyotype.  In  addition  to  the  eight 
biarmed  pairs  described  as  common  for  Noctilio,  Pteronotus,  and  Macrotus, 
the  karyotypes  of  most  species  examined  include  several  other  biarmed  elements, 
the  banding  patterns  of  which  suggest  independent  fusions  of  acrocentric  ele¬ 
ments. 

An  alternative  hypothesis  would  be  to  propose  a  noctilionid-mormoopid- 
like  karyotype  as  primitive.  Such  a  primitive  karyotype  would,  however,  require 
additional  events — fission  would  have  to  precede  several  independent  fusions. 
As  demonstrated  by  Mascarello  et  al.  (1974)  for  rodents,  the  establishment  of 
fission  rearrangements  is  quite  rare,  whereas  Robertsonian  fusion  products  are 
the  most  common  type  of  euchromatic  variation  observed  between  closely  related 
taxa.  Therefore,  a  fission-fusion  mode  not  only  would  require  additional  events, 
it  would  also  be  less  probable  from  a  cytogenetic  standpoint. 


116 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


117 


In  the  following  paragraphs  on  the  evolutionary  relationships  indicated  by 
karyotypic  data,  it  is  assumed  that  the  primitive  karyotype  for  the  Phyllosto- 
matoidea  was  2 n  =  46,  FN  =  60  and  with  a  morphology  similar  to  that  of  Macrotus 
waterhousii.  The  discussion  is  essentially  limited  to  G-band  data  because  all 
other  would  be  too  speculative  and  G-band  studies  of  most  subfamilies  will  un¬ 
doubtedly  appear  shortly.  Proposed  karyotypic  relationships  for  some  phyl- 
lostomatid  taxa,  based  on  standard  karyotypes,  are  presented  in  Baker  (1973), 
Greenbaum  et  al.  (1975),  and  Gardner  (1977). 

Systematic  Affinities 
Familial  Affinities 

The  first  instance  where  the  members  of  the  Mormoopidae,  Noctilionidae, 
and  Phyllostomatidae  were  classified  together,  but  distinct  from  all  other  bats,  was 
Winge  (1941).  Smith  (1972)  drew  similar  conclusions — the  Phyllostomatoidea 
consisted  of  the  families  Mormoopidae,  Noctilionidae,  and  Phyllostomatidae. 
G-band  chromosomal  data  strongly  support  this  classification  and  suggest  that 
Pteronotus  and  Noctilio  shared  a  common  evolutionary  ancestor  in  which  five 
Robertsonian  fusions  became  established  (Patton,  1976).  These  data  indicate  that 
the  Noctilionidae  and  Mormoopidae  are  more  closely  related  to  each  other  than 
either  is  to  the  Phyllostomatidae.  Smith  (1972)  came  to  the  same  conclusions 
based  on  morphological  data.  The  most  recent  common  ancestor  of  Pteronotus 
and  Noctilio  probably  had  a  2n  =  36  condition. 

The  degree  of  chromosomal  divergence  distinguishing  Noctilio  from  Pteronotus 
is  the  least  known  to  separate  two  mammalian  families.  Before  someone 
jumps  to  the  conclusion  that  the  families  Mormoopidae  and  Noctilionidae  are 
confamilial,  I  would  point  out  that  prior  to  the  study  by  Patton  (1976),  there  had 
been  considerable  disagreement  as  to  the  evolutionary  affinities  of  both  families 
(Smith,  1972).  In  fact,  there  would  be  little  agreement  as  to  what  family 
Noctilio  should  be  placed  in  if  it  were  not  awarded  familial  status.  Some  clas¬ 
sifications  have  included  the  mormoopids  as  a  subfamily  of  the  Phyllostomatidae 
(Miller,  1907;  see  also  the  review  by  Smith,  1972),  and  the  chromosomal  data 
merely  indicate  that  if  all  lineages  evolved  from  the  most  recent  ancestor  of  the 
mormoopid-phyllostomatid  line  are  to  be  included  in  the  family  Phyllostomatidae, 
then  the  Noctilionidae  should  also  be  reduced  to  a  subfamily. 

Chromosomal  data  from  Noctilio  and  Mormoops  further  document  the  fact 
that  karyotypic  change  is  not  a  requirement  for  the  evolution  of  a  magnitude  of 
morphological  difference  worth  of  recognition  of  a  higher  taxonomic  category 
(Patton,  1976).  It  has  been  suggested  by  Wilson  et  al.  (1975)  that  the  large  degree 
of  morphological  evolution  in  mammals  is  due  to  regulator  gene  alterations  by 


Fig.  1. — A  composite  of  two  G-banded  karyotypes  of  Macrotus  waterhousii  prepared  for 
use  as  standard  reference  in  describing  chromosomal  events  in  the  family  Phyllostomatidae 
as  proposed  by  Patton  (1976).  Both  homologs  from  the  two  spreads  are  presented  in  order 
that  minor  variation  can  be  observed.  Figure  courtesy  of  Rebecca  A.  Bass. 


118 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


chromosomal  mutations.  However,  few  changes  in  primitive  linkage  groups  are 
often  characteristic  of  rather  divergent  taxa  of  bats  and  rodents  (see  also  Mascarello 
et  al.,  1974;  Stock  and  Hsu,  1973),  which  leads  one  to  conclude  that  at  least  some 
mammalian  taxa  have  evolved  primarily  via  point  mutations  and  have  conserved 
the  primitive  gene  arrangements.  A  similar  conclusion  can  be  drawn  for  reptiles, 
based  on  chromosomal  banding  analysis  of  turtles  (Stock,  1972;  Bickham  and 
Baker,  1976),  and  for  birds  (Stock  et  al.,  1974). 

Subfamilial  Affinities 

There  has  been  only  one  paper  in  which  G  and  C-band  data  have  been  used  to 
relate  species  from  different  subfamilies  (Stock,  1975)  and  this  work  found 
essentially  no  G-band  autosomal  homologies  between  Carollia  (subfamily 
Carolliinae)  and  Choeroniscus  (subfamily  Glossophaginae).  From  standard 
karyotypes,  a  close  relationship  between  these  two  genera  had  been  proposed 
(Baker,  1967).  Stock  noted  that  the  X  elements  were  essentially  the  same  between 
the  two  genera  but  concluded  that  there  were  no  data  supporting  a  close  common 
ancestor  for  Carollia  and  Choeroniscus  and  suggested  that  these  two  genera 
be  placed  in  separate  subfamilies.  I  have  little  doubt  that  a  complete  G-band  study 
of  the  genera  within  all  subfamilies  will  reveal  the  evolutionary  relationships  of 
most  subfamilies.  G  and  C-band  studies  on  the  Brachyphyllinae  and  Des- 
modontinae  (by  Rebecca  A.  Bass)  and  Stenoderminae  (by  Anette  Johnson)  are 
presently  being  conducted  in  my  laboratory. 

Relationships  Within  Subfamilies 

Phyllostomatinae. — Relationships  within  the  subfamily  Phyllostomatinae 
were  studied  by  Patton  (1976),  but  his  results  were  somewhat  incomplete  because 
only  five  of  11  genera  (involving  10  of  33  species)  were  studied;  these  were 
arranged  into  three  groups:  1)  Micronycteris,  2)  Tonatia,  Mimon ,  and  Phyl- 
lostomus,  and  3)  Macrotus. 

The  Macrotus  group  could  have  evolved  from  any  lineage  just  as  long  as  it 
became  separated  from  the  other  stocks  prior  to  the  establishment  of  any  chro¬ 
mosomal  rearrangements.  The  karyotype  of  Macrotus  waterhousii  has  been 
proposed  as  like  that  which  was  primitive  for  the  family  (see  above).  The  karyo¬ 
type  of  M.  californicus  (2n  —  40,  FN  =  60)  would  then  be  derived  by  three  centric 
fusions  (Davis  and  Baker,  1974),  which  would  have  been  independent  events 
from  fusions  established  in  the  other  two  lines  discussed  below. 

Patton’s  (1976)  Micronycteris  group  is  characterized  by  the  sharing  of  two 
derived  arrangements.  One  is  a  terminal  translocation  of  chromosome  13  onto 
pair  26/25  and  the  other  is  a  Robertsonian  fusion  between  acrocentric  pairs 
1 8  and  2 1 .  All  other  rearrangements  within  the  Micronycteris  cluster  appear  to 
have  been  achieved  through  independent  events  within  the  three  subgenera 
( Trinycteris,  Micronycteris,  and  Lampronycteris)  studied  by  Patton.  The 
hypothesized  primitive  karyotype  for  the  subgenera  Trinycteris  and  Micronycteris 
would  be  2n  =  42,  FN  =  58.  The  fact  that  these  species  ( minuta ,  nicefori,  and 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


119 


brachyotis)  representing  three  subgenera,  can  be  chromosomally  related,  strongly 
reinforces  the  natural  status  of  at  least  portions  of  the  genus.  I  have  heard  several 
people  propose  that  this  genus  is  a  catchall  with  several  species  of  questionable 
generic  affinity.  One  species  that  cannot  be  related  chromosomally  to  the  other 
representatives  of  the  genus  thus  far  studied  is  M.  megalotis,  the  type  species  of 
the  genus. 

The  Tonatia- Mimon- Phyllostomus  group  is  identified  by  five  shared  derived 
chromosomal  events:  four  Robertsonian  fusions  (22/3,  8/9,  17/12,  29/27)  and 
one  inversion  (4/5).  These  chromosomal  characteristics  are  shared  by  Tonatia 
minuta,  Phyllostomus  discolor,  P.  hastatus,  and  Mimon  crenulatum.  The  ancestral 
karyotype  for  the  common  ancestor  probably  had  a  2 n  =  38,  FN  =  60.  A  common 
ancestor  for  Phyllostomus  hastatus,  P.  discolor,  and  Mimon  crenulatum  is 
suggested  by  three  shared  fusion  events  (18/13,  1 4a/2 1 ,  30/28).  This  would 
mean  that  the  common  ancestor  for  these  three  species  had  a  karyotype  with  a 
2n  =  32  or  34.  As  Robertsonian  fusion  products  occurring  independently  in  forms 
containing  only  two  acrocentric  linkage  groups  could  only  lead  to  the  same 
fusion  product,  a  2/?  =  34  divergence  cannot  be  totally  discounted  (Patton,  1976). 
The  possibility  of  a  2/7=34  divergence  is  strengthened  by  Mimon  cozumelae 
having  a  2n  =  34,  FN  =  60  karyotype. 

The  karyotype  of  Tonatia  bidens  (2/i  =  16)  is  so  derived  from  the  Macrotus  and 
Tonatia  minuta  karyotypes  that  it  could  not  be  related  to  those  of  other  members 
of  the  subfamily.  Again,  this  points  out  a  case  where  most  chromosome  divergence 
has  been  limited  to  changes  that  can  be  traced  by  homology  of  G-bands,  but 
during  the  evolution  of  T.  bidens  numerous  chromosomal  changes  became 
established.  If  systematic  position  were  based  solely  on  chromosomal  divergence, 
one  would  have  to  recognize  T.  bidens  as  generically  distinct  from  other  phyl- 
lostomatines  possibly  with  subfamilial  status,  a  ridiculous  conclusion  in  my 
opinion. 

Glossophaginae. — There  are  no  G-band  studies  on  the  generic  relationships 
within  the  Glossophaginae.  The  only  published  G-banded  karyotype  is  of 
Choeroniscus  intermedius  (Stock,  1975),  which  is  discussed  above  under  sub- 
familial  relationships. 

Gardner  (1977)  presented  a  phylogeny  of  the  Glossophaginae  based  on 
standard  karyotypes  and  in  most  cases  has  followed  the  most  parsimonious  routes. 
However,  I  cannot  accept  that  the  similar  karyotypes  of  Choeronycteris  and 
Hylonycteris  are  the  result  of  parallelism.  This  2/7=16  karyotype  is  undoubtedly 
derived,  and  I  feel  that  it  is  explained  best  as  being  due  to  their  common  ancestor 
having  a  diploid  number  of  16.  G-banding  should  be  valuable  in  settling  this 
difference  in  interpretation. 

Carolliinae. — G-band  data  (Stock,  1975)  have  been  published  for  one  (Carollia, 
three  species  studied)  of  the  two  genera  of  the  Carolliinae.  Carollia  brevicauda 
and  C.  perspicillata  share  two  chromosomal  features  (an  X-autosomal  trans¬ 
location  and  similar  heterochromatin  patterns)  that  distinguish  these  two  species 
from  at  least  some  individuals  of  C.  castanea.  Pine  (1972),  in  a  study  based  on 
classical  morphological  features,  concluded  that  C.  brevicauda  and  C.  perspicillata 


120 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  2. — G-banded  karyotype  of  Artibeus  jamaicensis.  Figure  courtesy  of  M.  Anette 
Johnson. 


were  more  closely  related  to  each  other  than  either  is  to  C.  castanea.  Carollia 
castanea  has  two  chromosomal  races  that  are  based  on  the  presence  of  the  X- 
autosomal  translocation  in  specimens  from  Central  America  (Patton  and  Gardner, 
1971)  and  Colombia  (Baker  and  Bleier,  1 970)  and  the  absence  of  this  translocation 
in  Peruvian  specimens  (Patton  and  Gardner,  1971). 

Patton  and  Gardner  (1971)  argued  that  the  absence  of  the  X-autosomal  trans¬ 
location  in  some  populations  of  C.  castanea  is  the  result  of  the  primitive  condition 
being  maintained.  This  would  best  explain  the  current  taxonomic  distribution 
of  the  X-autosomal  translocation  if  the  ancestor  of  all  Carollia  species  was 
polymorphic  for  this  translocation.  In  C.  perspicillata ,  C.  subrufa,  and  C.  brevi- 
cauda,  this  translocation  became  fixed  and  characteristic  of  the  species,  whereas 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


121 


Chromosome  groups 
B  C  D  E  F 


*  % 

n»  *■* 

00 

«•« 

*  M 

m 

m, 

<u>  £& 

2N=44 

c  a 

43 

£  J 

A  a 

«• 

fittt 

42 

X 

-  _■ 

»u 

40 

nn 

»er 

wz 

39 

*> 


ii  ii  m  i«  si 

sa 

tZ  s; 

r» 

***  ** 

for  each  karyotype:  2  n  of  44  is  A  A  BB 
40  is  Aa  Bb  cc ;  and  2// =  39  is  aa  Bh  cc. 


Fig.  3. — G-banded  karyotype  and  partial  karyo¬ 
types  of  specimens  of  Uroderma  bilobatum  with 
various  diploid  numbers.  Top  row.  Complete 
karyotype  of  a  specimen  with  2/7  =  44.  Rows  2-4 
are  group  A,  B,  and  C  chromosomes  of  specimens 
with  a  2/7  of  43,  42,  40,  and  39,  respectively.  Chro¬ 
mosomes  in  groups  A,  B,  and  C  account  for  all  the 
variation  in  diploid  number.  D  chromosomes  are 
the  two  sex  elements.  E  chromosomes  are  the  large 
biarmed  elements  and  F  chromosomes  are  the 
acrocentric  elements  that  are  not  involved  in  the 
variation.  Group  A  chromosomes  appear  as  either 
a  small  pair  of  biarmed  elements  plus  a  pair  of 
acrocentrics  (designated  as  the  A  A  morph),  a 
small  biarmed  element  plus  an  acrocentric  and  a 
subtelocentric  element  (Aa  morph)  or  as  two  sub- 
telocentric  elements  (aa).  Group  B  chromosomes 
appear  as  two  pairs  of  acrocentric  elements  (BB), 
a  pair  of  acrocentrics  unequal  in  size  and  a  sub- 
metacentric  element  (Bb)  or  two  submetacentric 
elements  (bb).  Group  C  appears  as  four  acrocentric 
elements  (CC),  three  acrocentric  elements  (Cc)  or 
two  acrocentric  elements  (cc).  Genetic  designation 
CC;  2/7  =  43  is  AA  Bb  CC:  2/7  =  42  is  Aa  BB  Cc;2n  — 


in  C.  castanea  some  populations  became  fixed  for  both  conditions.  This  explana¬ 
tion  might  be  correct  and  I  agree  that  it  is  the  first  choice;  however,  based  on  the 
limited  data  now  available,  an  alternative  explanation  cannot  be  ruled  out.  It 
is  possible  that  the  absence  of  the  X-autosomal  translocation  in  some  C.  castanea 
is  due  to  a  fission  of  these  elements  and  represents  a  condition  more  derived  than 
that  characteristic  of  C.  perspicillata,  C.  subrufa,  and  C.  brevicauda. 

Stenoderminae. — G  and  C-band  chromosomal  data  for  Sturnira  lilium, 
Artibeus  jamaicensis  (Fig.  2),  Enchisthenes  harti,  and  Uroderma  bilobatum 
(2n  =  44  cytotype,  Fig.  3)  are  described  by  Baker  et  al.  (1979).  The  G-band- 
ing  pattern  for  Artibeus  and  Sturnira  revealed  that  the  similarity  in  the  gross 
karyotypes  reflected  homology  with  only  one  autosomal  change  (a  pericentric 


122 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


inversion)  distinguishing  their  respective  karyotypes.  The  karyotype  of  Enchist- 
henes  harti  could  be  derived  from  the  Artibeus  karyotype  by  a  reciprocal  trans¬ 
location  involving  two  autosomes.  This  translocation  changes  two  submetacentric 
chromosomes  in  Artibeus  to  two  subtelocentric  chromosomes  in  Enchisthenes. 

It  was  more  difficult  to  show  homology  between  Artibeus  and  the  Uroderma 
2/i  =  44  karyotype.  Two  pairs  of  Artibeus  autosomes  were  homologous  with 
elements  in  Uroderma ;  the  other  1 2  pairs  of  Artibeus  (85  per  cent  of  the  autosomal 
pairs)  autosomes  required  rearrangement  to  derive  the  Uroderma  karyotype. 
For  some  chromosomal  segments,  homologous  elements  could  not  be  determined 
between  the  two  karyotypes. 

Artibeus  has  five  (Baker  et  al.,  1979)  pairs  of  biarmed  chromosomes  that  are 
homologous  with  pairs  found  in  Macrotus.  The  biarmed  pairs  homologous  between 
Artibeus  and  Macrotus  are  thought  to  be  primitive  for  the  family  (Patton,  1976). 
Uroderma  and  Macrotus  have  no  homologous  biarmed  chromosomes;  however, 
they  share  acrocentric  chromosomal  homologies. 

Only  two  pairs  of  chromosomes  (both  biarmed)  were  shared  by  all  four 
stenodermine  genera  studied.  These  two  pairs  are  not  found  in  any  of  the  other 
subfamilies  studied  (Phyllostomatinae,  Patton,  1976;  Glossophaginae  and 
Carolliinae,  Stock,  1975)  and  are,  therefore,  potentially  valuable  indicators  of 
a  common  ancestry  for  these  and  other  stenodermine  genera.  Such  marker  elements 
should  prove  valuable  in  determing  if  Brachyphylla  has  evolutionary  affinities 
with  the  Stenoderminae.  The  G-band  data  for  Sturnira  are  interpreted  as 
additional  documentation  that  the  genus  Sturnira  has  a  common  ancestry  with 
the  Stenoderminae  and  should  be  recognized  as  a  member  of  that  subfamily 
(Baker  et  al.,  1978).  G-banded  karyotypes  for  Uroderma  bilobatum  are  shown  in 
Fig.  3  and  are  discussed  below  in  the  following  section. 

Desmodontinae  and  Brachyphyllinae. — No  G-banded  karyotypes  have  been 
published  for  the  subfamilies  Desmodontinae  and  Phyllonycteririae. 

[Note  added  in  galley. — G-band  data  are  now  available  for  several  additional 
species  so  that  the  following  important  conclusions  can  be  drawn.  The  glos- 
sophagine  genera  Glossophaga  and  Monophyllus  have  identical  G-band  chromo¬ 
somal  homologies  with  species  of  Phyllonycteris,  Erophylla,  and  Brachyphylla. 
These  data  indicate  that  these  five  genera  shared  a  common  ancestor  after  sep¬ 
arating  from  the  other  subfamilial  lineages  (with  the  possible  exception  of  the 
Carolliinae)  and  that  Brachyphylla  is  properly  associated  with  the  genera 
Phyllonycteris  and  Erophylla  (Baker  and  Bass,  1979),  as  was  suggested  by  Silva 
Taboada  and  Pine  (1969).  However,  when  the  genus  Brachyphylla  is  placed  in 
this  subfamily,  Brachyphyllina  Gray,  1866,  becomes  the  oldest  available 
family-group  name  for  the  subfamily  (Phyllonycterinae  was  first  proposed  by 
Miller,  1907).  The  proper  name  of  the  subfamily  then  would  be  Brachyphyllinae. 

In  a  manuscript  recently  submitted  for  publication  by  Rebecca  A.  Bass  and  the 
author,  it  was  shown  that  the  vampire  bats  (Desmodontinae)  shared  a  common 
ancestry  with  the  glossophagines  and  brachyphyllines,  after  this  lineage  separated 
from  the  remainder  of  the  family.  ] 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


123 


Variation  within  Species 

From  the  standpoint  of  population  biology,  this  is  the  level  where  chromosomal 
variation  can  be  used  to  make  the  most  significant  studies.  The  role  of  various 
types  of  mechanisms  of  chromosomal  evolution  can  be  studied  as  an  isolating 
mechanism,  effective  means  of  producing  heterosis,  supergenes,  etc.  Variation  at 
this  level  can  be  due  to  populational  polymorphisms  or  chromosomal  races. 

Polymorph  isms 

A  widely  distributed  polymorphism  has  been  described  (Baker  et  al.,  1912b) 
for  Mimon  crenulatum.  The  polymorphism  was  found  in  samples  from  Peru, 
Trinidad,  and  Colombia  and  is  believed  to  be  restricted  to  the  fifth  largest  pair  of 
autosomes.  Three  morphs  of  this  chromosome  were  identified  from  each  of  the 
three  localities.  For  polymorphism  to  be  maintained  over  such  a  wide  geographic 
range,  it  must  offer  a  selective  advantage  to  the  species  greater  than  the  expense 
of  its  maintenance. 

Baker  and  Lopez,  (1970a)  demonstrated  a  polymorphism  also  for  Uroderma 
magnirostrum.  Eleven  of  thirteen  specimens  examined  from  Colombia  had  a 
diploid  number  of  36,  whereas  two  had  a  diploid  number  of  35.  Because  the  size 
of  the  additional  biarmed  element  was  greater  than  a  fusion  between  any  two 
acrocentrics,  the  polymorphic  system  may  not  be  the  result  of  a  simple  centric 
fusion. 

Other  cases  of  chromosomal  variation  at  a  single  locality  are  based  on  the 
discovery  of  a  single  aberrant  individual,  which  may  represent  a  balanced  poly¬ 
morphic  system  or  variation  that  originated  within  that  individual. 

A  centric  fusion  was  reported  in  a  female  Mesophylla  macconnelli  from 
Trinidad;  nine  other  specimens  from  this  locality  did  not  possess  the  chromosome. 
An  Artibeus  toltecus  from  San  Luis  Potosi,  Mexico,  had  a  2n  =  32  with  what 
appeared  to  be  a  trisomy  for  a  small  autosome  and  one  other  male  from  this  locality 
had  a  2n  =  31,  which  is  normal  for  the  species. 

In  a  sample  of  78  Uroderma  bilobatum  from  near  Choluteca,  Honduras,  one 
individual  had  a  2n  —  31,  which  resulted  from  a  fusion  of  two  acrocentrics 
into  a  metacentric  of  the  same  general  size  range  as  the  subtelocentric  autosomes. 
Chromosomal  variation  at  this  locality  is  common  as  a  result  of  hybridization 
between  two  cytotypes  (see  the  discussion  on  Uroderma  below);  however,  this 
centric  fusion  is  easily  identifiable  from  those  events  that  separate  the  two 
cytotypes  because  the  fusion  product  is  a  metacentric,  and  such  an  element  has 
not  been  observed  in  332  other  specimens  of  Uroderma  bilobatum  from  Central 
America. 


Chromosomal  Races 

Chromosomal  races  are  known  for  three  species  of  phyllostomatid  bats.  What 
originally  was  reported  as  chromosomal  races  in  Macrotus  waterhousii  proved 
to  be  specific  differences  characteristic  of  two  species:  M.  waterhousii ,  with 


124 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


2n  —  46,  and  M.  californicus,  with  2n  —  40  (Davis  and  Baker,  1974;  Greenbaum 
and  Baker,  1976).  Two  races  are  known  for  Micronycteris  hirsuta  (Baker  et  al., 
1973).  One  is  a  2n=  30  cytotype  from  Middle  America  characterized  by  a  single 
pair  of  submetacentric  autosomes.  Specimens  from  Trinidad,  on  the  other  hand, 
have  a  karyotype  with  a  2n  =  28,  FN  =  32  and  show  two  pairs  of  submetacentric 
autosomes  and  two  less  pairs  of  acrocentrics.  The  degree  of  divergence  in  cranial 
and  forearm  measurements  in  the  specimens  karyotyped  is  too  low  to  suggest 
that  the  two  chromosomal  races  represent  distinct  species  (Baker  et  al.,  1973). 

Two  races  of  Vampyressa  pusilla  were  described  by  Baker  et  al.  (1973).  One 
race  has  a  2n=18,  FN  =  20  with  two  pairs  of  submetacentric  autosomes  and 
six  acrocentric  pairs.  The  X  is  a  subtelocentric,  and  the  Y  is  a  small  distinctly 
biarmed  element.  This  race  is  known  from  Honduras,  Nicaragua,  and  Costa 
Rica.  The  second  race,  found  in  Colombia,  has  a  karyotype  that  consists  of  a 
2n  —  24  in  females  and  a  2n  =  23  in  males,  with  an  FN  of  22.  There  are  no  sub¬ 
metacentric  autosomes.  To  explain  divergence  between  the  two  races  requires  at 
least  three  events.  Even  though  the  magnitude  of  variation  is  greater  than  that 
characteristic  of  most  congeneric  species  of  phyllostomatids,  no  exomorphological 
or  cranial  differences  were  found  that  could  distinguish  the  races  (Baker  et  al., 
1973).  Data  from  V.  pusilla  documents  another  case  of  discordant  rates  of 
evolution  between  classical  and  karyotypic  morphology. 

Uroderma  bilobatum,  Peters’  tent-making  bat,  is  the  third  species  of  phyl- 
lostomatid  bat  known  to  have  chromosomal  races.  The  three  chromosomal  races 
reported  for  this  species  have  been  the  object  of  considerable  study  (Baker  and 
Lopez,  1970a;  Baker  et  al.,  1972a;  Baker  and  McDaniel,  1972;  Baker  et  al., 
1975);  one  zone  of  contact  between  two  races  has  been  located.  Information  on 
the  nature  and  dynamics  of  this  zone  could  be  valuable  in  understanding  some 
aspects  of  the  speciation  process. 

Elucidation  of  the  processes  by  which  one  species  becomes  transformed  into 
two  or  more  is  the  key  to  understanding  evolution.  The  genetic  interactions 
involved  between  two  diverging  populations  within  a  species  dictate  the  evolu¬ 
tionary  future  of  these  populations.  Although  several  theories  have  been 
postulated  for  such  genetic  interactions  and  their  relationships  to  the  process  of 
speciation,  actual  measurements  of  the  interaction  are  difficult  to  make  and 
definitive  data  are  lacking. 

An  important  aspect  involved  in  speciation  is  the  chromosomal  com¬ 
patibility  between  diverging  populations.  One  proposed  model  of  speciation 
(stasipatric  speciation  by  White,  1968)  is  based  entirely  on  chromosomal  di¬ 
vergence.  The  situation  with  JJroderma  bilobatum  (see  details  below)  does  not 
exactly  fit  the  stasipatric  model  put  forth  by  White;  however,  Uroderma  offers 
a  unique  opportunity  to  examine  the  role  of  karyotypic  diversity  and  the  resulting 
interaction  between  two  interbreeding  populations.  A  detailed  understanding  of 
the  mechanisms  and  events  occurring  at  the  contact  zone  between  two  chro- 
mosomally  characterized  populations  of  Uroderma  bilobatum  is  important 
because  we  will  be  able  to  observe  a  stage  of  evolution  that  could  result  in  the 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


125 


formation  of  two  species.  It  could  provide  insight  into  how  chromosomal  changes 
become  fixed  within  a  population. 

The  paucity  of  measurements  on  the  genetic  interactions  resulting  in  speciation 
(especially  in  mammals)  can  be  attributed  to  both  the  difficulties  in  obtaining  such 
measurements  and  the  inability  of  available  techniques  to  identify  appropriate 
biological  situations  for  study.  In  order  to  attempt  to  measure  the  genic  interactions 
that  might  produce  speciation,  it  is  first  essential  to  locate  a  situation  where  popu¬ 
lations  have  diverged.  In  addition,  it  is  necessary  to  be  able  to  identify  within 
the  population  first  generation  crosses  between  the  types  (referred  to  as  Fis, 
although  this  does  not  imply  specific  status  of  the  types)  and  backcross  individ¬ 
uals. 

Measurements  of  degree  of  exomorphological  and  cranial  divergence  have 
proven  inadequate  for  such  studies.  By  the  point  in  time  when  organisms  are 
sufficiently  diverged  to  enable  the  recognition  of  Fi  and  backcross  individuals  by 
these  techniques,  the  stage  at  which  the  most  significant  interactions  occur  has 
passed.  Numerous  studies  can  be  cited  to  document  this  problem  (see  Lidicker, 
1962,  for  a  review  of  the  problems  of  subspecific  evolution  in  mammals).  Eveji 
when  interpreted  with  the  use  of  the  most  sophisticated  multivariate  techniques, 
measurements  of  exomorphological  and  cranial  features  cannot  identify  with  any 
certainty  the  Fi  and  backcross  individuals  or  measure  genetic  interaction  (see 
Baker  et  al. ,  1 975).  The  extent  to  which  gene  flow  has  been  reduced  when  alloptric 
populations  reestablish  contact  simply  cannot  be  ascertained  with  any  degree  of 
accuracy  from  measurements  of  exomorphological  and  cranial  features. 

In  cases  where  two  interacting  populations  are  characterized  by  chromosomal 
differences,  Ft  individuals  will  have  a  predictable  karyotype  unique  from  that  of 
both  parental  types.  If  the  chromosomal  differences  are  of  sufficient  magnitude, 
the  first  generation  backcross  individuals  will  have  karyotypes  distinguishable 
from  the  Fi  and  parental  karyotypes.  Such  biological  situations  provide  an  ex¬ 
cellent  case  for  detailed  investigations  into  the  genetic  interactions  of  divergent 
populations  and  the  process  of  speciation. 

It  should  be  pointed  out,  however,  that  anytime  karyotypes  are  used  to  identify 
diverging  populations,  one  is  studying  a  special  case  because  chromosomes  are 
involved  and  chromosomes  could  be  the  primary  isolating  mechanism.  There  are 
many  isolating  mechanisms  known,  and  it  is  possible  that  each  represents  a 
special  case.  It  is  also  probable  that  no  single  isolating  mechanism  is  involved  in 
all  cases  of  speciation.  The  aim  of  the  detailed  study  of  Uroderma  in  my  laboratory 
is  to  investigate  the  role  of  chromosomal  divergence  in  the  evolutionary  process 
as  exemplified  by  these  bats. 

The  classical  systematics  and  distribution  of  U.  bilobatum  are  as  follows: 
Uroderma  bilobatum  occurs  at  lower  elevations  from  southern  Mexico  southward 
through  parts  of  tropical  South  America.  Based  on  variations  in  external  and 
cranial  measurements,  karyology,  and  pelage  color,  six  subspecies  ( bilobatum , 
molaris,  convexum,  trinitatum,  davisi,  and  thomasi )  are  recognized  (Davis, 
1968;  Baker  and  McDaniel,  1972).  Extensive  chromosomal  investigations  of  the 


126 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  4. — Geographic  distribution  of  samples  of  Uroderma  bilobatum  within  contact  zone 
(see  Table  2).  Specific  localities  are  1)  El  Salvador:  La  Paz:  3.0  mi.  NW  La  Herradura;  2) 
Usulatan:  3.0  mi.  E  Usulatan;  3)  Honduras:  Valle:  9  mi.  SE  Nacaome;  4)  Valle:  10.0  mi. 
SSW  Nacaome,  5)  Choluteca:  10.2  mi.  NW  Choluteca,  6)  Choluteca:  11.5  mi.  SW  Cholu- 
teca;  7)  Nicaragua:  Chinandega:  3.5  mi.  NW  and  1.5  mi.  S  Chinandega.  The  2/i  =  44  parental 
cytotype  occurs  at  localities  1-3,  both  parental  cytotypes  are  present  at  locality  4,  and  the 
2/;  =  38  parental  cytotype  occurs  at  localities  5-7. 

Uroderma  bilobatum  complex  have  revealed  chromosomal  divergence  greater 
than  that  reported  for  any  other  species  of  bat  (Baker,  1967,  1970a,  1970ft; 
Hsu  et  al.,  1968;  Baker  and  Hsu,  1970;  Capanna  and  Cibitelli,  1970). 

Karyotypically,  the  U.  bilobatum  complex  can  be  divided  into  the  following 
groups:  2n  =  44,  davisi  (central  Honduras  north  to  southern  Mexico;  (Baker  and 
McDaniel,  1972);  2«  =  38,  including  convexum  (central  Honduras  south  to 
northern  South  America  on  the  Pacific  versant),  and  molar  is  (Mexico  to  Nicaragua 
on  Atlantic  versant;  as  suggested  by  Davis,  1968);  and  2n  =  42,  consisting  of  the 
nominal  subspecies  trinitatum  and  bilobatum  (South  American  mainland). 
Uroderma  ft.  thomasi,  which  has  not  been  karyotyped,  is  known  from  western 
South  America.  Uroderma  ft.  convexum  (2 n  =  38)  and  U.  ft.  davisi  (2n  =  44)  have 
been  found  to  form  a  contact  zone  over  200  kilometers  in  length  (Fig.  4)  that 
extends  from  southern  El  Salvador,  across  the  Pacific  coast  of  Honduras  and 
northwestern  Nicaragua  (Baker  et  al.,  1975). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


127 


Conclusions  concerning  the  nature  of  chromosomal  variation  in  Uroderma 
between  the  2/i  =  38  and  2/7  =  44  forms  are  based  on  G-band  data  (Fig.  3).  The 
diploid  number  at  the  contact  zone  in  Central  America  ranges  from  38  to  44, 
with  individuals  of  all  intermediate  diploid  numbers  being  represented.  North¬ 
west  of  the  contact  zone  the  diploid  number  is  44  and  to  the  southeast  is  38 
(Fig.  4).  Intermediate  individuals  are  not  known  from  outside  of  the  zone  of 
contact.  The  differences  between  the  two  parental  types  (38  to  44)  result  from 
three  separate  events,  each  involving  a  translocation  or  fission,  depending  on 
direction  of  the  event.  The  first  change  to  be  discussed  (designated  the  A  chro¬ 
mosomes)  is  shown  in  column  A  of  Fig.  3.  One  morph  (represented  by  a  capital 
A)  has  a  small  biarmed  element  and  an  acrocentric  element;  the  other  morph 
(represented  by  a  lower  case  a)  has  these  two  elements  fused  to  form  a  subtelo- 
centric  chromosome.  Where  only  a  standard  karyotype  is  available,  the  number 
of  large  A' s  in  the  karyotype  will  be  reflected  by  the  number  of  small  biarmed 
autosomes  present  in  the  complement. 

The  second  event  (designated  the  B  chromosomes)  involves  a  centric  fusion- 
fission  in  which  morph  B  (column  B,  Fig.  3)  appears  as  two  acrocentrics  (the 
smallest  acrocentric  in  the  2/7  =  44  karyotype  and  one  of  the  medium-sized 
acrocentric  elements).  Morph  b  is  a  subtelocentric  element  representing  a  fusion 
of  the  two  acrocentric  elements  in  B.  This  variation  can  be  recognized  in  a  standard 
karyotype  because  each  b  is  reflected  by  a  decrease  of  one  in  the  diploid  number 
and  an  increase  of  one  in  the  number  of  large  biarmed  elements,  without  effecting 
a  decrease  in  the  number  of  small  biarmed  autosomes. 

The  third  change  (designated  the  C  chromosomes),  as  shown  in  column  C  of 
Fig.  3,  is  a  terminal  translocation  in  which  a  small  acrocentric  element  is  trans¬ 
located  to  the  end  of  the  long  arm  of  the  longest  acrocentric  element  in  the  karyo¬ 
type.  For  each  morph  C,  there  will  be  two  acrocentrics  in  the  karyotype,  whereas 
each  morph  c  is  a  single  large  acrocentric  in  which  the  segments  homologous  to 
the  two  C  acrocentrics  are  fused.  Production  of  the  c  morph  reduces  the  diploid 
number  by  one  and  reduces  the  number  of  acrocentrics  by  one  but  does  not  alter 
the  number  of  biarmed  elements  (either  small  or  large)  in  the  karyotype. 

Although  the  exact  nature  of  these  changes  can  be  identified  only  by  the  G-band 
patterns,  the  three  changes  produce  distinct  morphological  differences  in  the 
chromosomes  that  allows  one  to  determine  the  chromosomal  phenotype  from  a 
standard  karyotype  for  the  A,  B,  and  C  chromosomes  of  any  individual.  Using 
the  ABC  designation  for  the  chromosomal  variation  enables  the  characterization 
of  all  of  the  individuals  involved  in  the  contact  zone.  An  animal  with  AABBCC 
would  be  a  2n  =  44  parental  type  and  an  animal  with  aabbcc  would  be  a  2n  —  38 
parental  type.  Each  capital  letter  in  the  phenotype  will  raise  the  diploid  number 
above  38  by  one.  For  example,  an  animal  with  a  phenotype  of  aaBbCCo r  AaBbCc 
would  have  a  diploid  number  of  41  and  an  animal  with  AABbCC  would  have  a 
diploid  number  of  43.  I  have  determined  the  chromosomal  phenotype  for  333 
specimens  from  the  zone  of  contact. 

C-banding  patterns  are  important  because  they  identify  segments  of  the 
chromosomes  that  are  believed  to  be  heterochromatic  in  nature.  Variation  in 


128 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


the  amount  of  C-band  material  between  karyotypes  is  not  thought  to  interfere 
with  meiosis  as  does  variation  in  euchromatin.  It  is  important,  therefore,  to  know 
the  amount  and  placement  of  C-band  positive  material  within  the  three  chro¬ 
mosomal  variants.  The  karyotype  of  Uroderma  bilobatum  contains  very  little 
C-band  positive  material.  Most  biarmed  elements  have  a  small  amount  near  the 
centromere  and  one  medium-sized  acrocentric  (not  one  of  the  A,  B,  or  C  chro¬ 
mosomes)  has  a  C-band  proximal  to  the  centromere.  Although  all  of  the  A,  B,  and 
C  chromosomes  have  a  small  amount  of  centromeric  C-band  material,  none  of  the 
major  segments  involved  in  the  variation  is  C-band  positive.  The  small  biarmed 
pair  involved  in  the  Aa  variation,  however,  has  heterochromatin  incorporated 
into  part  of  one  arm.  All  of  the  C-bands  identified  in  this  small  biarmed  element 
(of  the  A  morph)  are  present  in  the  subtelocentric  a  morph  fusion  product  and 
the  a  morph  subtelocentric  has  about  as  much  C-band  material  as  do  the  two 
elements  of  the  A  morph. 

Although  the  break  and  alteration  may  have  occurred  in  this  C-band  positive 
area,  no  major  addition  or  deletion  of  C-band  material  seems  to  have  occurred. 
Variations  in  the  C-banding  patterns  do  not  seem  to  be  involved  in  the  genetic 
strategy  of  Uroderma.  This  constitutes  a  major  difference  between  chromosomal 
evolution  in  this  species  and  that  seen  in  some  rodents,  for  example,  Peromyscus 
(Duffey,  1972;  Pathak  et  al.,  1973). 

The  zone  of  contact  between  the  cytotypes  of  Uroderma  is  about  200  kilometers 
in  length  but  its  width  is  not  known.  Because  Uroderma  is  ecologically  restricted 
to  the  relatively  low  lands  of  the  coast,  the  zone  cannot  be  over  100  kilometers 
wide  at  many  places  and  must  be  considerably  narrower  at  some.  The  two  parental 
cytotypes  occur  sympatrically  at  a  single  locality  in  my  sample  (Fig.  4),  and  the 
area  of  overlap  of  parental  cytotypes  is  probably  not  much  longer  than  30  kilo¬ 
meters.  At  the  locality  where  the  two  parental  cytotypes  occur  sympatrically,  most 
individuals  have  a  hybrid  karyotype  (for  instance,  within  a  sample  size  of  15, 
one  bat  had  2n  —  38,  one  had  2n=39,  two  had  2/7  =  40,  five  had  2n  =  4l,  one  had 
2u  =  41,  one  had  2/7  =  42,  three  had  2/7  =  43,  and  two  had  2/7  =  44).  Intensive 
hybridization  occurs  in  the  central  part  of  the  zone  between  Nacome  and  Choluteca, 
Honduras.  Away  from  this  area,  parental  cytotypes  probably  do  not  come  into 
direct  contact,  and  hybrid  karyotypes  are  found  much  less  frequently;  I  suggest 
that  these  are  primarily  the  result  of  the  survival  and  successful  reproduction  of 
backcross  individuals. 

Different  types  of  chromosomal  rearrangements  produce  different  meiotic 
aberrations  and,  therefore,  the  percentage  of  sterile  gametes  in  a  heterozygote 
will  be  a  function  of  the  nature  of  the  rearrangement.  If  the  rate  of  production  of 
sterile  gametes  is  the  only  factor  regulating  the  penetration  of  a  chromosomal 
morph  of  one  parental  type  into  a  population  of  the  other  parental  form,  an  increase 
of  sterile  gametes  should  result  in  a  decrease  in  successful  penetration  into  the 
other  cytotype.  Furthermore,  across  the  zone  of  contact  the  frequency  of  the 
penetrating  chromosomal  morphs  should  produce  a  symmetrical  bell-shaped 
curve  reflecting  the  greater  number  of  Fi  backcross  individuals  near  the  zone  and 
the  decrease  in  such  individuals  with  distance  away  from  the  area  of  primary 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


129 


contact.  The  width  of  this  symmetrical  curve  for  a  given  chromosomal  aberration 
would  be  a  function  of  the  severity  of  meiotic  selection  against  heterozygotes  of 
that  type  of  aberration. 

If  factors  other  than  meiotic  mechanisms  play  a  role  in  the  penetration  of  one 
chromosomal  morph  into  populations  of  the  other,  there  is  no  reason  to  assume 
that  selection  on  both  sides  of  the  zone  should  be  the  same  and  the  frequency  of  Fi 
and  heterozygous  individuals  across  the  zone  would  not  be  symmetrical. 

The  frequencies  ( p  and  ^-values)  of  the  various  chromosomal  morphs  from 
333  specimens  of  Uroderma  bilobatum  from  the  contact  zone  are  shown  in  Table 
2.  The  two  northernmost  localities  (La  Herradura  and  Usulatan)  have  similar 
chromosomal  frequencies.  Notably,  the  b  morph  of  the  B  chromosomes  has  been 
the  most  successful  in  surviving  in  these  populations,  whereas  the  c  morph  was 
not  found  to  be  present  in  any  of  the  133  specimens  from  these  two  localities. 
This  might  be  predicted  based  on  the  type  of  segregation  that  would  result  in  a 
heterozygote  for  the  respective  B  and  C  chromosomes.  Centric  fusions  and 
fissions  (origin  of  the  B  chromosomal  system)  are  not  believed  to  interfere  greatly 
with  the  meiotic  process,  especially  if  preferential  segregation  occurs.  Proper 
segregation  probably  would  not  be  affected,  and  therefore  natural  selection  at  the 
meiotic  level  would  be  ineffective  in  eliminating  such  variation  from  the  popu¬ 
lation.  On  the  other  hand,  such  translocations  as  might  have  given  rise  to  the 
C  chromosomal  variation  should  result  in  loss  of  about  25  per  cent  of  the  gametes 
in  the  heterozygote  if  there  has  been  crossing  over  in  the  portion  homologous  to 
the  large  acrocentric.  It  would  appear  that,  in  the  absence  of  other  factors,  the 
variation  in  the  B  chromosomes  would  be  more  common  in  all  populations  than 
would  variation  in  the  C  chromosome.  In  samples  from  the  southeastern  part 
of  the  contact  zone,  survival  of  the  B  chromosome  is  less  frequent  than  C;  C 
actually  accounts  for  about  4.5  per  cent  of  the  C  chromosomes  at  the  Choluteca 
locality  (Table  2,  Fig.  4).  Two  of  86  individuals  were  heterozygous  (Cc)  at  the 
Chinandega  locality.  The  per  cent  variation  resulting  from  each  chromosomal 
change  is  not  the  same  northwest  and  southeast  of  the  central  part  of  the  zone  (Table 
2),  which  suggests  that  successful  reproduction  of  hybrid  and  backcross  individuals 
is  not  explained  totally  by  meiotic  problems,  but  that  possibly  fitness  of  the  adult 
varies  as  well. 

It  also  should  be  noted  that  although  Chinandega  is  closer  to  the  central  part 
of  the  contact  zone  than  is  La  Herradura,  less  total  chromosomal  variation  is 
found  at  Chinandega  (4.6  per  cent  of  the  individuals  had  hybrid  karyotypes)  than 
at  La  Herradura  (14  per  cent  had  hybrid  karyotypes). 

Baker  et  al.  (1975)  concluded  that  the  chromosomal  data  pointed  to  con¬ 
siderable  chromosomal  flow  between  the  cytotypes.  At  that  time  it  was  not 
possible  to  identify  patterns  in  exchange  and  survival  of  the  different  morphs. 
From  the  above  data  (Table  2),  there  is  clearly  a  pattern  of  selective  chromosomal 
flow  between  cytotypes.  If  the  variation  in  the  C  chromosomes  is  used  to  estimate 
chromosomal  flow  (and  implied  gene  exchange)  of  the  2n  =  38  chromosomes  into 
the  2n  =  44  populations,  the  data  strongly  suggest  no  exchange  (the  one  individual 
at  Nacome  that  was  heterozygous,  Cc,  was  a  presumed  Fi).  On  the  other  hand,  if 


130 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  2. — Frequency  of  the  chromosomal  morphs  at  localities  in  the  zone  of  hybridization. 
Numbers  preceding  localities  identify  geographic  samples  in  Fig.  4,  where  exact  localities 
are  given.  The  2n  =  44  parental  type  occurs  at  localities  1-3.  Both  parental  cytotypes  occur  at 
locality  4.  At  localities  5-7,  the  2n  =  38  parental  type  is  present. 


1 .  La  Herradura 

2.  Usulatan 

3.  Nacaome 

4.  Hybrid 
locality 

5.  San 
Lorenzo 

6.  Choluteca 

7.  Chinandega 

Sample  size 

50 

83 

9 

15 

12 

78 

86 

Chromosomal 

morphs 

A 

p  =  98;  q  =  02 

p=  99;  q  =  01 

p  =  94 ;  q  =  06 

p  =  60;  q  =  40 

p  =  42;  q  — 58 

p  =  05;  q  =  95 

p  =  01;  q=  99 

B 

p  =  95;  q  — 05 

p=  95;  q  =  05 

p  =  78;  q  =  22 

p  =  53;  q  =  47 

p  =  29;  q  =  71 

p  =  01;  q  =  99 

p  =  00;  q  =  100 

C 

p=  100;  q  =  00 

p  =  100;  q  =  00 

p  =  94;  q  =  06 

p=  57;  q  =  43 

p=  33;  q  =  67 

p=  04;  q  =  96 

p  =  01;  q=  99 

the  B  chromosomes  are  used,  the  implications  are  different.  Chromosomal  data 
fit  the  pattern  of  introgression  in  which  some  chromosomes  are  allowed  to  enter 
the  “chromosome  pool”  of  another  type  by  hybridization  and  backcrossing, 
but  other  chromosomes  are  selected  against. 

The  pattern  of  chromosomal  morphs  across  this  contact  zone  closely  fits  the 
tension  zone  (White,  1973;  Key,  1974)  characteristic  of  stasipatric  speciation. 
In  evaluating  my  data  in  light  of  White’s  model,  several  points  need  to  be  made. 
First,  at  this  time  it  is  impossible  to  determine  if  this  zone  is  the  result  of  primary 
or  secondary  contact.  White’s  model  requires  that  the  zone  be  the  product  of 
primary  contact.  Second,  stasipatric  tension  zones  have  been  described  for  several 
species  (Bush,  1975),  and  a  suite  of  the  biological  characteristics  of  these  species 
do  not  fit  those  of  Uroderma.  In  species  with  low  vagility,  the  tension  zone  is 
usually  not  more  than  a  few  hundred  meters  wide;  in  Uroderma ,  species  with  high 
vagility,  the  zone  is  more  than  200  kilometers  in  breadth.  Third,  Uroderma  is 
K-selected,  whereas  other  species  with  tension  zones  are  R-selected. 

My  data  point  out  the  fact  that  tension  zones  need  not  be  composed  of  species 
characterized  by  low  vagility  and  R-selection.  Although  the  zone  of  inter¬ 
action  between  the  two  Uroderma  cytotypes  might  or  might  not  be  in  equilibrium, 
it  will  eventually  proceed  to  one  of  several  endpoints.  One  possibility  is  that  the  two 
cytotypes  could  develop  additional  isolating  mechanisms,  such  as  behavioral  or 
postmating,  and  evolve  into  two  species.  Another  possibility  is  the  replacement  of 
one  parental  type  by  the  other  via  the  mechanisms  of  competition  or  genetic 
swamping.  A  less  likely  outcome  could  be  the  survival  of  some  intermediate 
cytotype  with,  say,  42  chromosomes  (for  instance,  AABBcc).  At  any 
rate,  this  type  of  chromosomal  variation  undoubtedly  offers  a  unique  set  of 
possibilities  on  which  evolution  can  act.  The  unique  nature  of  these  biological 
circumstances  certainly  offers  a  rare  chance  to  observe  evolution  in  action. 

Electrophoretic  data  would  be  extremely  valuable  in  shedding  some  light  on 
the  history  of  Uroderma  populations  that  have  produced  this  tension  zone. 
Electrophoretic  data  indicate  that  when  two  species  have  been  derived  by  the 
classical  allopatric  model,  the  level  of  similarity  of  allozymes  is  usually  about  85 
per  cent  or  less  (Avise,  1974).  If  these  chromosomal  differences  accumulated 
during  a  long  allopatric  period,  it  could  be  predicted  that  these  two  chro¬ 
mosomal  races  should  have  accumulated  a  significant  number  of  fixed  allelic 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


131 


I  II  **  I  i  ii 


I!  il 


•  • 


I  •  *  •  * 


*  * 


*  * 


#  41 


»  § 


B 

Fig.  5. — Sample  patterns  of  C-bands  of  phyllostomatid  bats:  A,  Phyllostomus  elongatus ; 
B,  Enchisthenes  hcirtii. 


differences;  however,  if  the  process  that  gave  rise  to  the  current  condition  has  been 
like  that  proposed  by  White  (1968,  1973),  very  few  electrophoretic  differences 
should  be  detectable.  This  situation  is  currently  under  study  by  Ira  F.  Greenbaum. 

Miscellaneous  Cytogenetic  Studies 

In  addition  to  the  more  systematically  oriented  papers  discussed  above,  there 
have  been  a  few  more  detailed  studies  on  biochemical  aspects  involving  karyo¬ 
typic  data  for  phyllostomatid  bats. 

C-bands. — C-band  material  for  phyllostomatid  bats  is  described  in  enough 
species  that  general  trends  can  be  predicted  (Stock,  1975;  Patton,  1976;  Baker 
et  al.,  1978).  In  general,  phyllostomatid  bats  have  C-band  material  restricted 
to  the  centromeric  region.  The  amount  found  is  small,  similar  to  that  shown  in 
Fig.  5  for  Phyllostomus  and  Enchisthenes ,  respectively.  However,  in  some  species 
( Carollia  perspicillata  and  Choeroniscus  intermedius)  there  are  additional  portions 
of  the  karyotype  carrying  C-band  positive  material  (Stock,  1975).  Also,  see  the  dis¬ 
cussion  on  C-band  material  in  Uroderma  under  the  section  on  chromosomal  races. 

Nucleolar  organizer  regions. — Two  papers,  both  dealing  with  Carollia 
perspicillata  and  C.  castenea,  have  reported  studies  of  nucleolar  organizer 
regions  (NOR)  in  phyllostomatids  (Hsu  et  al.,  1975;  Goodpasture  and  Bloom, 
1975).  Hsu  et  al.  (1975)  used  DNA/rRNA  (ribosomal  RNA)  hybridization  to 
reveal  NOR’s.  In  the  karyotype  of  C.  perspicillata,  the  only  NOR  was  located  on 
s  the  X  chromosome;  their  studies  of  C.  castanea  were  made  on  a  transformed 
culture.  Hsu  et  al.  concluded  that  the  origin  of  the  NOR  on  the  Carollia  X/auto- 


132 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


somal  chromosome  was  from  the  X  and  not  the  translocated  autosomal  portion. 
Because  DNA-RNA  hybridization  is  difficult  and  expensive,  Goodpasture  and 
Bloom  (1975)  tested  the  feasibility  of  using  ammoniacal  silver  to  reveal  NOR’s. 
Their  methods  localized  NOR’s  at  the  same  points  as  did  the  methods  of  Hsu 
et  al.  The  same  individuals  were  studied  from  in  vitro  cultures.  Goodpasture  and 
Bloom  (1975)  present  theories  on  the  cytological  basis  for  silver  NOR  staining. 

Cesium  chloride  buoyant  densities. — Arrighi  et  al.  (1968,  1972)  reported  on 
cesium  chloride  buoyancy  in  phyllostomatid  bats.  Findings  are  summarized  in 
the  latter  paper.  Ten  species  of  phyllostomatid  bats  ( Anoura  geoffroyi,  Artibeus 
fallax=  A.  lituratus  in  Jones  and  Carter,  1976,  Artibeus  lituratus,  Carollia 
perspicillata,  Chiroderma  villosum,  Choeroniscus  intermedius,  Sturnira 
erythromos,  Sturnira  lilium ,  Sturnira  magna ,  and  Uroderma  bilobatum )  were 
studied  and  values  ranged  from  1.6982  for  Carollia  perspicillata  to  1.7005  for 
Anoura  geoffroyi  and  Sturnira  erythromos.  These  values  fall  within  those  given 
for  other  Microchiroptera  (1.696  to  1.702)  from  the  families  Rhinolophidae, 
Molossidae,  and  Vespertilionidae,  but  only  slightly  overlap  the  values  reported  for 
Megachiroptera  (1.694  to  1.697).  Of  the  families  of  Microchiroptera,  Phyl- 
lostomatidae  had  values  nearest  those  for  the  Megachiroptera.  Although  the 
magnitude  of  difference  between  the  suborders  is  small,  it  is  the  greatest  found 
between  suborders  of  mammals  and  is  interpreted  as  supporting  relatively 
ancient  lineages  for  the  two  suborders  (Arrighi  et  al.,  1972). 

X  chromosomes. — G-banded  X  chromosomes  for  a  variety  of  mammals 
(including  eight  species  of  phyllostomatids)  were  studied  by  Pathak  and  Stock 
(1974).  They  found  that  X  chromosomes  always  have  two  dark  staining,  trypsin 
resistant  bands  regardless  of  the  centromere  placement.  They  interpreted  these 
data  as  supporting  Ohno’s  (1967)  hypothesis  that  the  mammalian  X  chromosome 
is  extremely  conservative  in  genetic  constitution. 

Acknowledgments 

Laura  Kyle  and  Anette  Johnson  assisted  in  the  preparation  of  plates  and  Table 
1 .  I  am  especially  grateful  to  my  graduate  students  who  have  been  of  considerable 
assistance  in  accumulating  these  data.  I  also  thank  my  many  colleagues  who  aided 
in  the  collection  of  specimens,  read  drafts  of  the  manuscript,  and  offered 
criticisms.  This  work  was  made  possible  by  several  grants  from  the  National 
Science  Foundation,  of  which  the  most  recent  was  No.  DEB-76-20580. 

Literature  Cited 

Arrighi,  F.  E.,  J.  Bergendahl.,  and  M.  Mandel.  1968.  Isolation  and  characterization 
of  DNA  from  fixed  cells  and  tissues.  Exptl.  Cell  Res.,  50:47. 

Arrighi,  F.  E.,  W.  Z.  Lidicker,  Jr.,  M.  Mandel,  and  J.  Bergendahl.  1972.  Hetero¬ 
geneity  in  CsCl  buoyant  densities  of  Chiropteran  DNA.  Biochem.  Genet.,  6:27-30. 
Avise,  J.  C.  1974.  Systematic  value  of  electrophoretic  data.  Syst.  Zool.,  23:465-481. 
Baker,  R.  J.  1967.  Karyotypes  of  bats  of  the  family  Phyllostomidae  and  their  taxonomic 
implications.  Southwestern  Nat.,  12:407-428. 

- .  1970*7.  Karyotypic  trends  in  bats.  Pp.  65-96,  in  Biology  of  Bats  (W.  A.  Wimsatt, 

ed.),  Academic  Press,  New  York,  l:xxii+406  pp. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


133 


- .  1970/?.  The  role  of  karyotypes  in  phylogenetic  studies  of  bats.  Pp.  303-312,  in 

About  bats  (B.  H.  Slaughter  and  D.  W.  Walton,  eds.),  Southern  Methodist  Univ. 
Press,  Dallas,  vii  +  339  pp. 

- .  1973.  Comparative  cytogenetics  of  the  New  World  leaf-nosed  bats  (Phyl- 

lostomatidae).  Periodicum  Biologicum,  75:37-45. 

Baker,  R.  J.,  and  R.  A.  Bass.  1979.  Evolutionary  relationship  of  the  Phyllonycterinae  to 
the  glossophagine  genera,  Glossophaga  and  Monophyllus.  J.  Mamm.,  60: in 
press. 

Baker,  R.  J.,  and  W.  J.  Bleier.  1971.  Karyotypes  of  bats  of  the  subfamily  Carolliinae 
(Mammalia,  Phyllostomatidae)  and  their  evolutionary  implications.  Experientia, 
27:220. 

Baker,  R.  J.,  and  H.  H.  Genoways.  1972.  The  phyllostomatid  bat,  Vcunpyressa  brocki, 
in  Colombia.  Bull.  South.  California  Acad.  Sci.,  71:54. 

- .  1976.  A  new  species  of  Chiroderma  from  Guadeloupe,  West  Indies  (Chiroptera: 

Phyllostomatidae).  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  39:1-9. 

Baker,  R.  J.,  and  T.  C.  Hsu.  1970.  Further  studies  on  the  sex-chromosome  systems  of 
the  American  leaf-nosed  bats  (Chiroptera,  Phyllostomatidae).  Cytogenet., 
9:131-138. 

Baker,  R.  J.,  and  G.  Lopez.  1970«.  Chromosomal  variation  in  bats  of  the  genus  Uroderma 
(Phyllostomatidae).  J.  Mamm.,  51:786-789. 

- .  19706.  Karyotypic  studies  of  the  insular  populations  of  bats  on  Puerto  Rico. 

Caryologia,  23:465-472. 

Baker,  R.  J.,  and  V.  R.  McDaniel.  1972.  A  new  subspecies  of  Uroderma  bilobatum 
(Chiroptera:  Phyllostomatidae)  from  Middle  America.  Occas.  Papers  Mus., 
Texas  Tech  Univ.,  7:1-4. 

Baker,  R.  J.,  W.  R.  Atchley,  and  V.  R.  McDaniel.  1972a.  Karyology  and  mor¬ 
phometries  of  Peters’  tent-making  bat,  Uroderma  bilobatum  Peters  (Chiroptera, 
Phyllostomatidae).  Syst.  Zool.,  21:414-429. 

Baker,  R.  J.,  A.  L.  Gardner,  J.  L.  Patton.  19726.  Chromosomal  polymorphism  in  the 
phyllostomatid  bat,  Mimon  crenulatum  (Geoffroy).  Experientia,  28:969. 

Baker,  R.  J.,  H.  H.  Genoways,  W.  J.  Bleier,  and  J.  W.  Warner.  1973.  Cytotypes  and 
morphometries  of  two  phyllostomatid  bats,  Micronycteris  hirsuta  and  Vampyressa 
pusilla.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  17:1-10. 

Baker,  R.  J.,  W.  J.  Bleier,  and  W.  R.  Atchley.  1975.  A  contact  zone  between  karyotypi- 
cally  characterized  taxa  of  Uroderma  bilobatum  (Mammalia:  Chiroptera).  Syst. 
Zool.,  24:133-142. 

Baker,  R.  J.,  R.  A.  Bass,  and  M.  A.  Johnson,  n.d.  Evolutionary  implications  of  chro¬ 
mosomal  homology  in  four  genera  of  stenodermine  bats  (Phyllostomatidae: 
Chiroptera).  Evolution,  submitted. 

Becak,  M.  L.,  R.  F.  Batistic,  L.  D.  Vizotto,  and  W.  Becak.  1969.  Sex  determining 
mechanism  XYiY2  in  Artibeus  lituratus (Chiroptera,  Phyllostomidae).  Experi¬ 
entia,  25:81-83. 

Bickham,  J.  W.,  R.  J.  Baker.  1976.  Chromosome  homology  and  evolution  of  emydid 
turtles.  Chromosoma,  54:201-219. 

- .  1977.  Implications  of  chromosomal  variation  in  Rhogeessa  (Chiroptera: 

Vespertilionidae).  J.  Mamm.,  58:448-453. 

Bush,  G.  L.  1975.  Modes  of  animal  speciation.  Ann.  Rev.  Ecol.  and  Syst.,  1975:339-364. 

Cadena,  A.,  and  R.  J.  Baker.  1976.  Cariotipos  de  los  murcielagos  vampiros  (Chiroptera: 
Desmodinae).  Caldasia  II:  159-163. 

Capanna,  E.,  M.  V.  Civitelli.  1970.  Chromosomal  mechanisms  in  the  evolution  of 
chiropteran  karyotype.  Chromosomal  tables  of  Chiroptera.  Caryologia, 
23:79-111. 

Davis,  B.  L.,  R.  J.  Baker.  1974.  Morphometries,  evolution  and  cytotaxonomy  of  main¬ 
land  bats  of  the  genus  Macrotus  (Chiroptera:  Phyllostomatidae).  Syst.  Zool., 
23:26-39. 


134 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Davis,  W.  B.  1968.  Review  of  the  genus  Uroderma  (Chiroptera).  J.  Mamm.,  49:676-698. 

Duffey,  P.  A.  1972.  Chromosome  variation  in  Peromyscusr.  a  new  mechanism.  Science, 
176:1333-1334. 

Forman,  G.  L.,  R.  J.  Baker,  and  J.  D.  Gerber.  1968.  Comments  on  the  systematic 
status  of  vampire  bats  (family  Desmodontidae).  Syst.  Zool.,  17:417-425. 

Gardner,  A.  L.  1977.  Chromosomal  variation  in  Vampyressa  and  a  review  of  chro¬ 
mosomal  evolution  in  the  Phyllostomidae  (Chiroptera).  Syst.  Zool.,  26:300-318. 

Gardner,  A.  L.,  and  J.  P.  O’Neill.  1969.  The  taxonomic  status  of  Sturnira  bidens 
(Chiroptera:  Phyllostomidae)  with  notes  on  its  karyotype  and  life  history.  Occas. 
Papers  Mus.  Zool.,  Louisiana  State  Univ.,  38:1-8. 

Genoways,  H.  H.,  and  R.  J.  Baker.  1972.  Stenoderma  rufum.  Mammalian  Species, 
18:1-4. 

Goodpasture,  C.,  and  S.  E.  Bloom.  1975.  Visualization  of  nucleolar  organizer  regions  in 
mammalian  chromosomes  using  silver  staining.  Chromosoma,  53:37-50. 

Greenbaum,  I.  F.,  and  R.  J.  Baker.  1976.  Evolutionary  relationships  in  Macrotus 
(Mammalia:  Chiroptera):  biochemical  variation  and  karyology.  Syst.  Zool., 
25:15-25. 

Greenbaum,  I.  F.,  R.  J.  Baker,  and  D.  E.  Wilson.  1975.  Evolutionary  implications 
of  the  karyotypes  of  the  stenodermine  genera  Ardops,  Ariteus,  Phyllops  and 
Ectophylla.  Bull.  South.  California  Acad.  Sci.,  74:156-159. 

Hennig,  W.  1966.  Phylogenetic  systematics.  Univ.  Illinois  Press,  Urbana. 

Hsu,  T.  C.  and  K.  Benirschke.  1971.  An  atlas  of  mammalian  chromosomes.  Volume  5. 
Springer-Verlag,  New  York. 

- .  1973.  An  atlas  of  mammalian  chromosomes.  Volume  7.  Springer-Verlag, 

New  York. 

- .  1974.  An  atlas  of  Mammalian  chromosomes.  Volume  8.  Springer-Verlag, 

New  York. 

Hsu,  T.  C.,  R.  J.  Baker,  and  T.  Utakoji.  1968.  The  multiple  sex  chromosome  system  of 
American  leaf-nosed  bats  (Chiroptera,  Phyllostomidae).  Cytogenet.,  7:27-38. 

Hsu,  T.  C.,  S.  E.  Spirito,  and  M.  L.  Pardue.  1975.  Distribution  of  18  +  28S  ribosomal 
genes  in  mammalian  genomes.  Chromosoma,  53:25-36. 

Jones,  J.  K.,  Jr.,  and  D.  C.  Carter.  1976.  Annotated  checklist,  with  keys  to  subfamilies 

and  genera.  Pp.  7-38,  in  Biology  of  bats  of  the  New  World  family  Phyl- 

lostomatidae.  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.),  Spec.  Publ. 
Mus.,  Texas  Tech  Univ.,  10:1-218. 

Key,  K.  H.  L.  1974.  Speciation  in  the  Australian  Morabine  grasshoppers — taxonomy  and 
ecology.  Pp.  43-56,  in  Genetic  mechanisms  of  speciation  in  insects  (M.  J.  D. 
White,  ed.),  Sydney,  Australia,  and  New  Zealand  Book  Co.,  170  pp. 

Kiblisky,  P.  1969.  Chromosome  patterns  of  7  species  of  leaf-nosed  bats  of  Venezuela 
(Chiroptera — Phyllostomidae).  Experientia,  25:1203. 

Kniazeff,  A.  J.,  D.  Constantine,  W.  A.  Nelson-Rees,  D.  Schmidt,  and  R.  Owens.  1967. 

Studies  on  chiropteran  cell  lines.  41st  Tec.  Prog.  Rep.,  Naval  Biol.  Lab.  Suppl. 
Rept.,  CC-8:97-105. 

Lidicker,  W.  Z.,  Jr.  1962.  The  nature  of  subspecies  boundaries  in  a  desert  rodent  and  its 
implications  for  subspecies  taxonomy.  Syst.  Zool.,  11:160-171. 

Mascarello,  J.  T.,  A.  D.  Stock,  and  S.  Pathak.  1974.  Conservatism  in  the  arrangement 
of  genetic  material  in  rodents.  J.  Mamm.,  55:695-704. 

Miller,  G.  S.,  Jr.  1907.  The  families  and  genera  of  bats.  Bull.  U.S.  Nat.  Mus., 
57:xvii  +  282  pp. 

Nagorsen,  D.  W.,  and  R.  L.  Peterson.  1975.  Karyotypes  of  six  species  of  bats  (Chiroptera) 
from  the  Dominican  Republic.  Life  Sci.  Contrib.,  Royal  Ontario  Mus.,  28:1-8. 

Nelson-Rees,  W.  A.,  A.  J.  Kniazeff,  R.  J.  Baker,  and  J.  L.  Patton.  1968.  Intra¬ 
specific  chromosome  variation  in  the  bat,  Macrotus  waterhousii  Gray.  J.  Mamm., 
49:706-712. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


135 


Ohno,  S.  1967.  Sex  chromosomes  and  sex-linked  genes.  Springer-Verlag,  New  York, 
x  +  1 92  pp. 

Pathak,  S.,  and  A.  D.  Stock.  1974.  The  X  chromosomes  of  mammals:  karyological 
homology  as  revealed  by  banding  techniques.  Genet.,  78:703-714. 

Pathak,  S.,  T.  C.  Hsu,  and  F.  E.  Arrighi.  1973.  Chromosomes  of  Peromy sens  (Rodentia: 

Cricetidae).  IV.  The  role  of  heterochromatin  in  karyotypic  evolution.  Cyto- 
genet.  Cell  Genet.,  12:315-326. 

Patton,  J.  C.  1976.  Evolutionary  implication  of  the  G-banded  and  C-banded  karyotypes 
of  Phyllostomatoid  bats.  Unpublished  M.S.  thesis,  Texas  Tech  Univ.,  vi  +  349  pp. 

Patton,  J.  L.,  and  A.  L.  Gardner.  1971.  Parallel  evolution  of  multiple  sex-chromosome 
systems  in  the  phyllostomatid  bats,  Carollia  and  Choeroniscus.  Experientia, 
27:105. 

Pine,  R.  H.  1972.  The  bats  of  the  genus  Carollia.  Tech.  Monogr.,  Texas  Agric.  Exp. 
Sta.,  Texas  A&M  Univ.,  8: 1-125. 

Silva  Taboada,  G.,  and  R.  H.  Pine.  1969.  Morphological  and  behavioral  evidence  for  the 
relationship  between  the  bat  genus  Brachyphylla  and  the  Phyllonycterinae. 
Biotropica,  1:10-19. 

Smith,  J.  D.  1972.  Systematics  of  the  chiropteran  family  Mormoopidae.  Univ.  Kansas 
Publ.,  Mus.  Nat.  Hist.,  56:1-132. 

- .  1976.  Chiropteran  evolution.  Pp.  49-70,  in  Biology  of  bats  of  the  New  Work! 

family  Phyllostomatidae.  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter, 
eds.).  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  10:1-218. 

Stock,  A.  D.  1972.  Karyological  relationships  in  turtles  (Reptilia:  Chelonia).  Canadian 
J.  Genet.  Cytol.,  14:859-868. 

- .  1975.  Chromosome  banding  pattern  homology  and  its  phylogenetic  implications 

in  the  bat  genera  Carollia  and  Choeroniscus.  Cytogenet.  Cell  Genet.,  14:34-41. 

Stock,  A.  D.,  and  T.  C.  Hsu.  1973.  Evolutionary  conservatism  in  arrangement  of  genetic 
material.  Chromosoma,  43:21 1-224. 

Stock,  A.  D.,  F.  E.  Arrighi  and  K.  Stefos.  1974.  Chromosome  homology  in  birds: 

banding  patterns  of  the  chromosomes  of  the  domestic  chicken,  ring-necked  dove, 
and  domestic  pigeon.  Cytogenet.  Cell  Genet.,  13:410-418. 

Straney,  D.  O.,  M.  H.  Smith,  I.  F.  Greenbaum,  and  R.  J.  Baker.  1978.  Biochemical 
genetics.  Pp.  157-176,  in  Biology  of  Bats  of  the  New  World  Family  Phyllosto¬ 
matidae.  Part  III  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.),  Spec.  Publ. 
Mus.,  Texas  Tech  Univ.,  1 6: 1-442. 

White,  M.  J.  D.  1968.  Models  of  speciation.  Science,  159:1065-1070. 

- .  1973.  Animal  cytology  and  evolution.  3rd  Edition.  Cambridge  Univ.  Press., 

England,  961  pp. 

Wilson,  A.  C.,  G.  L.  Bush,  S.  M.  Case,  and  C.  M.  King.  1975.  Social  structuring  of 
mammalian  populations  and  rate  of  chromosomal  evolution.  Proc.  Nat.  Acad. 
Sci.,  72:5061-6065. 

Yonenaga,  Y.  1968.  Estudos  cromossomicos  em  especies  de  Chiroptera.  Ciencia  e 
Cultura,  20:172. 

Yonenaga,  Y.,  O.  Froto-Pessoa,  and  K.  R.  Lewis.  1969.  Karyotypes  of  seven  species  of 
Brazilian  bats.  Caryologia,  22:63-79. 


136 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


XX  XI!  XX  XX  XX  u  k» 

XK  XX  XX  XX  ft*  «*  *» 

««  M  • 

Plate  1 . — Karyotype  of  a  male  Lonchorhina  aurita  from  Trinidad. 


8  n  n  *i  «  a  « 

4*  ** 

Ah  A*  i*  no  AO  At  At 

fit  M  »•  ••  »»*  •• 

Plate  2. — Karyotype  of  a  female  Macrotus  waterhousii  from  Haiti. 

nn  U  on  XX  IS  XX 

Xl{  JfX  XX  »»  **  **  ** 

X3* 


Plate  3. — Karyotype  of  a  female  Micronycteris  brachyotis  from  Trinidad. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


137 


x*  OA 

/Ml  AO  no  (Ia  04  Ax  on 

^  1^  AA  •»*  ”  •*  ft 

Plate  4. — Karyotype  of  a  male  Micronycteris  hirsuta  from  Nicaragua. 


Aft  HU  IX  *x  Aft  A»  |« 

•»  mm 

(ift  XR  ha  ax 


Plate  5. — Karyotype  of  a  male  Micronycteris  megalot  is  from  Trinidad. 


K6  xx  f\j»  xx  x»  x a 

KM  AM  »* 

lift  flft  A/»  —  A 


Plate  6. — Karyotype  of  a  male  Micronycteris  minuta  from  Trinidad. 


138 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


U  88  88  88  **  XX 

XX  XX  X» 

f)»  to  to  8. 

Plate  7. — Karyotype  of  a  male  Micronycteris  nicefori  from  Trinidad. 

88  xx  x*  **  *«  *» 

K  K  •  «  MM 

ilArt/)  Ml  IfK  M 

AM  A  i|  «A  X* 

Plate  8. — Karyotype  of  a  male  Micronycteris  schmidtorum  from  Costa  Rica. 


M88K8  KUKxa  xk 

XX  XX  XX  *X  XX  xa  XX 

l 


Plate  9. — Karyotype  of  a  male  Mimon  cremilatum  from  Colombia. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


139 


KX  KK  XX  XX  xx  »* 

*>\  UK  MX  XX  xk  ««  ** 

"*  K  . 

Plate  10. — Karyotype  of  a  male  Phylloderma  stenops  from  Colombia. 

«!i  M  X»  KJI  MS  K#  l« 

XX  KX  *X  XX  X*  *« 
**  X. 

Plate  11. — Karyotype  of  a  male  Phyllostomus  discolor  from  Trinidad. 

MM  (tamii 

XX  XX  XX  XA  XX  XX  »x 

n 


Plate  12. — Karyotype  of  a  female  Phyllostomus  elongatus  from  Colombia. 


140 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


/WUX  KUtlxnu* 

X#  **  >«*  «*  xn  ** 

n. 

Plate  13. — Karyotype  of  a  male  Phyllostomus  hastat us  from  Trinidad. 


at «« is* 

Oft  00  AO  Aft  ft* 

Plate  14. — Karyotype  of  a  female  Tonal ia  bidens  from  Trinidad. 


Hi  ft!  XX  IX  Xt  n»  ** 

xx  XX  xx  xx  as  »• 

A  A 

Plate  15. — Karyotype  of  a  male  Tonatia  minuta  from  Trinidad. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


141 


K!  U  It  h  n  m  n 

XK  X«  xk  x*  Xfl  ** 

M  (14 

Plate  16. — Karyotype  of  a  female  Trachops  cirrhosus  from  Trinidad. 


%  XX  n  xh  xk  n  xx 

fix  XX  XX  XX  XX  XX  XX 

Kn 

Plate  17. — Karyotype  of  a  male  Vampyrum  spectrum  from  Trinidad. 

22  it  a  ii  M 
X*  »* 

16  16  it  M  XX  X. 

Plate  18. — Karyotype  of  a  male  Anoura  caudifer  from  Colombia. 


142 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Oft  X#  M  *tc  **  XX 


XX  mx 

it  il  iA  n*  H* 

Plate  19.— Karyotype  of  a  male  Anoura  cultrata  from  Costa  Rica. 


n 

/t  :«  Mi 

II  II  Aft  A*  Aft  i<  *u 

Plate  20. — Karyotype  of  a  female  Choeroniscus  godmani  from  Honduras. 


AA  ** 


ftfi  (SK  M  AA  AA  AA 

AX 


Plate  21. — Karyotype  of  a  female  Choeroniscus  intermedins  from  Trinidad. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


143 


Plate  22. — Karyotype  of  a  female  Choeronycteris  mexicana  from  Tamaulipas. 


n  n  n  n  n  t*  a 

XX  XX  X|  (A  H  *x  » 

”  X. 

Plate  23. — Karyotype  of  a  male  Glossophaga  soricina  from  Colombia. 


KK  U  „ 


nn  An 


x  x 


Plate  24. — Karyotype  of  a  female  Hylonycteris  underwood i  from  Costa  Rica. 


144 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


XX  XX  Al 

U  HD  ill  Afl  » 

Plate  25. — Karyotype  of  a  female  Lichonycteris  obscura  from  Nicaragua. 


il  ii  xx  H  ii » «i 

>*  i*  »a 

6A  fifi  ..  t. 

Plate  26. — Karyotype  of  a  male  Lionycteris  spitrrelli  from  Colombia. 


HH  till  KM 

nr-  m 


Plate  27. — Karyotype  of  a  male  Lonchophylla  robusto  from  Nicaragua. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


145 


XX 

AA  AA  Aft 

00  Oft  ft#  AA  Aft  Art  Art 

AO  OA  Art  »• 

Plate  28. — Karyotype  of  a  female  Lonchophylla  thomasi from  Colombia. 

18  M  XU  Un  xx  ix 

XX  XX  XX  XX  xr  aa 

«• 

Plate  29. — Karyotype  of  a  male  Monophyllus  redmani  from  Puerto  Rico. 


Ktt  AX  «s 


SIX 


* 

ttft. 


Plate  30. — Karyotype  of  a  male  Carollia  perspicillata  from  Colombia. 


146 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Hi!  88  88  xx  «x  xx  xx 

X8  XA  xx 

ns  «« 

<io  no  ao  »• 

Plate  31. — Karyotype  of  a  male  Rhinophylla  fischerae  from  Colombia. 

KK  AH  XX  XX  *Jt  «x  »4 

Aft  O  A  Xl 

Plate  32. — Karyotype  of  a  male  Rhinophylla  pumilio  from  Colombia. 

<*8  M  XX  xx  *x  xx 

XK  XX  K*  «*  »* 

U  XX  AX  li »* 

Plate  33. — Karyotype  of  a  male  Ametrida  centurio  from  Trinidad. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


147 


K  B  &  II  19  n  u 

SI  xa 

II  n  u  u  U 

Plate  34. — Karyotype  of  a  male  Ardops  nichollsi  from  Guadeloupe. 

XX  XX  XX  xx  xx 

XX  «x  ** 

Aft  Aft  xx  u  IU- 

Plate  35. — Karyotype  of  a  male  Ariteus  flavescens  from  Jamaica. 

XK  U  8R  kx  n  xx 

XX  a*  ah 

ftft  AA  Aft  KX  A.. 


Plate  36. — Karyotype  of  a  male  Artibeus  lituratus  from  Colombia. 


148 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


)'K  U  NK  Kfc  VJf  >tx  Jt* 

x  a  **  ** 

Aft  Aft  Aft  ft*  O" 

Plate  37. — Karyotype  of  a  male  Artibeus  pluieotis  from  Colombia. 

Cl  U  W  UK  M  XX 

XX  *n 

ill!  y  ~  I. 

Plate  38. — Karyotype  of  a  male  Chiroderma  improvisum  from  Guadeloupe. 

lc  t!  I*  XI  »*  **  «» 

IX  «* 

it  it  i<  1* 

Plate  39. — Karyotype  of  a  male  Chiroderma  salvini  from  Honduras. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


149 


H  ! 

n 

» 

U  si  si  st 

SS 

MM 

ii  M 

Plate  40- 

ii  U  5* 

-Karyotype  of  a  male  Ectophylla  alba  from  Costa  Rica. 

n  jx 

SI 

xy 

II 

XX 

XX 

x* 

Its  ns 

Plate  41.- 

fill 

—Karyotype  of  a 

11 

male  Enchisthenes  hartiii 

Ai  ft,. 

from  Colombia. 

nn  on 

no 

on 

on 

OA 

r*  n 

Plate  42. — Karyotype  of  a  male  Mesophylla  macconnelti  from  Trinidad. 


150 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


HI  Hn  8X  XX  U  9» 

XU  XX  *» 

M  u  n  .1,1  8.» 

Plate  43. — Karyotype  of  a  male  Phyllops  haitiensis  from  Haiti. 


U  M  Kt  K.h  u  n 

II  U  (A  >1  »  XI 

if. 

Plate  44. — Karyotype  of  a  male  Sphaeronycteris  toxophyllum  from  Colombia. 


KX  XX  XX  XX  XX  XX 

XX  X*  *K 

(MS  ftrt  no  xx  A. 

Plate  45. — Karyotype  of  a  male  Sturnira  erythromos  from  Colombia. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


151 


n  22  n  xs «  «  « 

S  U  X  *  X  21 

U  6»  io  M 

Plate  46. — Karyotype  of  a  female  Sturnira  mordax  from  Costa  Rica. 

XX  xx  xx  *> 

Ad  Aft  MAX  AX  Aft  AX 

AA  A*  XX 

6A  A*  X  A  MX  ft. 

Plate  47. — Karyotype  of  a  male  Uroderma  magnirostrum  from  Colombia. 

XX  nX  n  XX  XX  XX  XX 

**  aa 

Kft  Aft  hf> 


Plate  48. — Karyotype  of  a  female  Vampyressa  brocki  from  Colombia. 


152 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


M  XX  xx  xx  xx  ** « 

«ft  WoiiAM*  A. 

Plate  49. — Karyotype  of  a  male  Vampyressa  nymphaea  from  Honduras. 


U  xx 

Aft  flA  Oft  oo  aa  ** 

A. 

Plate  50. — Karyotype  of  a  male  Vampyressa  pusilla  from  Honduras. 


ftR  HO  00  lift  Aft  no  ,<A 

M  tifi  ** 

Plate  51. — Karyotype  of  a  male  Vampyressa  pusilla  from  Colombia. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


153 


U  KX  XX  XX  xx  xx  xx 

XX  Xtt  MM 

ftl UftAMA  Aa 

Plate  52. — Karyotype  of  a  male  Vampyrops  vittatus  from  Colombia. 

XI  II  Id  IX  88  ti  xx 

tx  XX  XX  a*  **  **  ** 

XX 

Plate  53. — Karyotype  of  a  female  Brachyphylla  caver narum  from  Puerto  Rico. 


81  88  XX  88  tf  XX  xx 

IX  *X  xx  *»  *«  **  " 


Plate  54. — Karyotype  of  a  male  Brachyphylla  nana  from  Haiti. 


154 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


ft  U  IK  H  n  n 

21  XX  xx  14  *»  *»  tx 


Plate  55. — Karyotype  of  a  male  Erophylla  sezekorni from  Puerto  Rico. 


XX  XX  XX  XX  x*  XX 

XA  XX  xx  X*  a*  x*  •« 

A*  K. 

Plate  56. — Karyotype  of  a  male  Phyllonycteris  aphylla  from  Jamaica. 


;(  n  »  si  is  xi  ii 

<1  XX  «  *»  "  “  !* 


Plate  57. — Karyotype  of  a  male  Phyllonycteris  poeyi  from  Haiti. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


155 


XI  lit  Kt  ss  n  g*  xx 

X.  XX  XX  XX  XX  ft*  X 

Plate  58. — Karyotype  of  a  male  Desmodus  rotundus  from  Veracruz. 

M  M  U  Xfi  8K  xx  xx 

KK  KX  U  XX  A&  Xft 

i?  «• 

Plate  59. — Karyotype  of  a  male  Diaemus  youngii from  Nicaragua. 

U  IS  n  88  n  n  it 

88  kx  x*  «  x*  **  ** 

**  s. 

Plate  60. — Karyotype  of  a  male  Diphylla  ecaudata  from  Veracruz. 


BIOCHEMICAL  GENETICS 


Donald  O.  Straney,  Michael  H.  Smith,  Ira  F.  Greenbaum, 

and  Robert  J.  Baker 


The  current  view  of  evolution  is  very  much  a  genetic  one.  Theoretical  develop¬ 
ments  since  the  rediscovery  of  Mendel’s  work  have  produced  an  intricate 
mathematical  theory,  integrating  genetic  and  ecologic  characteristics,  that 
provides  the  basis  for  our  understanding  of  the  evolutionary  process.  Within  this 
theoretical  framework  are  two  genetic  factors  of  critical  importance:  determination 
of  the  genetic  basis  of  fitness  and  the  genetic  structure  of  populations  in  space  and 
time.  Unfortunately,  information  about  these  two  factors  is  lacking  for  most 
groups  of  organisms.  The  first  is  nearly  impossible  to  establish  (Lewontin,  1974), 
and  the  second  requires  intensive  breeding  studies.  Until  recently,  the  spatial  and 
temporal  genetic  structure  of  natural  populations  had  been  described  only  for 
Drosophila  and  a  small  number  of  other  groups  (Dobzhansky,  1970).  In  order  to 
apply  theoretical  evolutionary  concepts  to  organisms  such  as  bats,  which  are 
difficult  to  breed  in  captivity,  it  has  been  necessary  to  assume  that  these  organisms 
behave  genetically  in  a  manner  similar  to  that  of  Drosophila. 

Most  species  of  phyllostomatid  bats  are  difficult  or  impossible  to  maintain  in 
captivity  in  the  numbers  required  for  genetic  breeding  studies  (see  Greenhall, 
1976).  In  addition,  lengthy  gestation  periods  and  low  productivity  make 
chiropterans,  in  general,  an  inefficient  group  with  which  to  work.  Bats  also  exhibit 
few  clear-cut  phenotypic  variants  within  populations  that  could  be  exploited  in 
genetic  studies,  as  has  been  done  with  Drosophila.  Thus,  the  genetic  properties 
of  chiropterans,  in  the  classical  sense,  are  unknown.  It  is  not  surprising  that, 
among  mammals,  easily  tractable,  prolific  and  variable  groups,  such  as  the  rodent 
genera  Mus  and  Peromyscus,  have  been  used  to  establish  genetic  baselines 
(Rasmussen,  1968). 

The  development  of  biochemical  techniques,  such  as  electrophoresis,  has 
enabled  genetic  studies  to  be  carried  out  at  the  protein  level,  thereby  circumventing 
many  of  the  traditional  problems  mentioned  above  concerning  maintaining  and 
breeding  animals.  Large  numbers  of  individuals  now  can  be  assayed  quickly,  even 
in  species  that  cannot  be  bred  in  the  laboratory,  to  give  baseline  data  documenting 
the  spatial  and  temporal  structure  of  natural  populations.  Breeding  studies  are 
needed  only  to  establish  the  inheritance  of  protein  banding  patterns,  and  for 
most  of  the  species  studied  so  far,  the  inheritance  of  these  banding  patterns 
appears  to  be  the  same  (Selander  et  al.,  1971,  Straney  et  al.,  1976a,  1976b).  Al¬ 
though  electrophoresis  and  other  biochemical  techniques  do  not  provide  a  com¬ 
plete  picture  of  evolutionary  genetics,  they  can  furnish  information  useful  in 
developing  models  of  evolution  and  do  have  the  potential  for  providing  data  that 
can  be  used  in  testing  phylogenetic  hypotheses.  Few  families  of  eutherian  mam¬ 
mals  are  as  ecologically  diverse  as  are  phyllostomatid  bats,  but  the  genetic  as- 


157 


158 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


pects  related  to  this  group’s  adaptive  radiation  are  poorly  understood.  It  is  from 
studies  of  organisms  such  as  the  Phyllostomatidae  that  information  on  the  rela¬ 
tionships  of  genetic,  ecological,  and  morphological  strategies  can  be  obtained. 
Study  of  phyllostomatid  genetics  though,  has  only  begun  and  is  limited  to  karyo¬ 
typic  (Baker,  this  volume)  and  biochemical  characters;  available  information 
points  more  toward  potential  questions  than  to  a  unified  picture  of  chiropteran 
genetics.  In  this  chapter,  we  review  the  published  works  on  biochemical  genetics 
of  phyllostomatid  bats  and  present  new  data  on  several  species  from  Trinidad. 

Literature 

Several  methods  have  been  used  to  study  the  biochemical  genetics  of  phyl- 
lostomatids.  Most  involve  electrophoresis  in  some  form  of  supporting  medium, 
such  as  cellulose  acetate,  polyacrilimide,  or  starch  gel.  Despite  differences  in 
medium,  the  process  is  the  same.  Proteins  in  tissue  extracts  are  placed  in  the 
medium  and  an  electric  current  applied  through  an  electrode  bridge.  The  pro¬ 
teins  are  ionized  by  the  buffer  used  in  the  electrode  bridge,  migrate  in  the 
electrical  field  in  characteristic  manners,  and  are  identified  by  means  of  appro¬ 
priate  histochemical  stains.  Differences  in  mobility  between  proteins  are  indicative 
of  variation  in  net  electric  charge  on  the  molecules.  Charge  variation  results 
from  changes  in  the  amino  acid  composition  of  the  proteins,  which  ultimately 
reflect  codon  differences  in  the  genes  involved.  Hence,  differences  in  mobility 
of  proteins  assayed  under  the  same  conditions  are  translatable  into  genetic  dif¬ 
ferences. 

The  earliest  examinations  of  chiropteran  biochemical  genetics  focused  on 
vespertilionids  and  were  conducted  by  Mitchell  (1966),  working  with  hemoglobin, 
and  Manwell  and  Kerst  (1966),  with  hemoglobin,  lactate  dehydrogenases, 
esterases,  and  general  tissue  proteins.  Both  papers  established  the  multiple 
component  structure  of  chiropteran  hemoglobin,  and  Manwell  and  Kerst  (1966) 
found  genetic  polymorphisms  in  several  species  that  involved  at  least  two  alleles 
at  the  lactate  dehydrogenase- 1  locus  and  several  alleles  at  the  esterase  and  tissue 
protein  loci.  Differences  in  protein  mobility  of  several  species  and  genera  were 
interpreted  as  genetic  variation  at  loci  encoding  these  proteins. 

Variation  in  bat  hemoglobins  has  been  studied  in  some  detail  by  Mitchell  (1970), 
Valdivieso  et  al.  (1969),  and  Tamsitt  and  Valdivieso  (1969).  Differences  in 
hemoglobin  molecules  were  found  primarily  at  the  familial  level,  although  within 
the  vespertilionids  examined  there  was  a  high  degree  of  variation  and  poly¬ 
morphism;  of  the  phyllostomatids  studied,  the  same  hemoglobin  moiety  was 
present.  Peptide  mapping  (Mitchell,  1970)  confirmed  the  identity  of  the  phyl¬ 
lostomatid  hemoglobins.  Desmodus  hemoglobin  (Tamsitt  and  Valdivieso,  1969) 
was  found  to  be  the  same  as  that  for  nine  other  species  of  phyllostomatids,  whereas 
hemoglobin  from  Pteronotus  was  unique,  results  consistent  with  current  taxonomic 
views  (Smith,  1972;  Jones  and  Carter,  1976).  Our  examination  of  samples  of 
phyllostomatids  from  Trinidad  (see  below)  suggests  that  variation  exists  in 
hemoglobins  of  some  species  of  this  family.  The  inheritance  of  this  variation  is 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


159 


not  clear  although  banding  patterns  suggest  allelic  variation  in  a  monomeric 
protein,  possibly  in  only  one  of  the  hemoglobin  chains. 

Valdivieso  and  Tamsitt  (1974)  examined  serum  proteins  of  18  species  from 
four  families  of  Neotropical  bats  and  were  able  to  isolate  four  to  eight  protein 
fractions.  Of  the  14  species  of  phyllostomatids  they  examined,  six  exhibited 
polymorphism  in  a  -globulins;  only  Artibeus  was  polymorphic  at  both  a  -  and  />- 
globulin  loci.  All  species  were  monomorphic  for  a -globulin.  Valdivieso  and 
Tamsitt  found  no  polymorphism  in  phyllostomatid  albumins;  however,  in  our 
samples  from  Trinidad,  albumin  is  the  single  most  variable  protein  locus  (see 
below).  Although  these  authors  noted  differences  in  albumins  between  species, 
genera,  and  families,  their  differences  are  not  concordant  with  our  data  (Table  1). 
Their  finding  that  the  albumin  of  Phyllostomus  hastatus  and  P.  discolor  differ 
from  all  other  phyllostomatids  appears  to  be  a  result  of  sampling  error.  Albumin 
allozymes  identical  to  those  of  Phyllostomus  were  present  in  other  phyllostomatids 
in  our  samples  (Table  1).  The  fact  that  in  their  sample  Molossus  albumins  were 
indistinguishable  from  those  of  some  phyllostomatids  is  probably  due  to  the  use 
of  cellulose  acetate  as  an  assay  medium.  Although  cellulose  acetate  makes  a 
quick  and  effective  medium  for  assaying  serum  protein  profiles,  the  accompany¬ 
ing  lack  of  resolution  makes  it  a  poor  system  for  surveys  of  genetic  variation. 
Their  (Valdivieso  and  Tamsitt,  1974)  conclusion  that  serum  protein  electrophore¬ 
sis  will  be  of  little  use  in  systematic  work  is  a  result  of  the  assay  medium  em¬ 
ployed,  the  number  of  species  examined,  and  sample  size. 

Straney  et  al.  (1976 a)  and  Greenbaum  and  Baker  (1976)  used  starch  gel  assay 
systems  to  examine  genetic  variation  at  17  and  21  loci,  respectively,  in  popu¬ 
lations  of  Macrotus.  In  45  individuals  sampled  from  a  population  of  M.  californicus 
in  Pima  County,  Arizona,  Straney  et  al.  described  six  polymorphic  loci,  but  the 
level  of  polymorphism  was  low,  with  no  locus  segregating  for  more  than  two 
alleles.  Indeed,  the  proportion  of  loci  in  the  heterozygous  state  in  the  average 
individual  (H)  in  this  population  was  0.03,  a  value  low  for  mammals  and  much 
less  than  that  found  in  Myotis  velifer{\\  =0.14;  Straney  et  al.,  \916a).  The  authors 
suggested  that  the  low  level  of  variation  in  Macrotus  was  consistent  with  the 
niche  width-variation  hypothesis,  as  modified  by  Selander  and  Kaufman  (1973). 

Greenbaum  and  Baker  (1976)  examined  genetic  variation  and  intra  and 
interspecific  similarity  in  Macrotus  californicus  and  M.  waterhousii  from  Arizona, 
Mexico,  and  Jamaica.  In  addition  to  the  polymorphisms  mentioned  above,  they 
described  others  at  two  gene  loci  in  populations  outside  of  Arizona.  Average 
population  heterozygosity  ranged  from  0.030  to  0.041  in  M.  californicus  and  from 
0.00  (for  specimens  from  an  interspecific  contact  locality)  to  0.043  in  M.  water¬ 
housii. 

Nei’s  genetic  distance  (D;  Nei  and  Roychoudhury,  1974)  reflects  the  number 
of  net  codon  differences  per  locus  between  a  pair  of  populations.  Genetic  distance 
between  populations  of  the  same  species  of  Macrotus  are  less  than  0.07.  Esti¬ 
mates  of  D  among  populations  of  Macrotus  are  within  the  range  reported  for 
other  mammals  (Greenbaum  and  Baker,  1976).  Jamaican  M.  waterhousii  are 


160 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1. —  Variation  in  albumin  in  Neotropical  phyllostomatid  bats.  Listed  are  those  species 
examined  both  in  this  study  and  by  Valdivieso  and  Tamsitt  (1974).  Entries  are  relative 
mobility  of  albumin  allozymes,  Artibeus  jamaicensis  taken  as  100.  Where  more  than  one 
allele  is  present  in  a  population,  mobilities  are  listed  in  decreasing  order  of  frequency. 


Species 

This  study 

Valdivieso  and  Tamsitt1 

Artibeus  jamaicensis 

100,  101,  103 

100,  106 

Artibeus  lituratus 

103,  100 

106 

Carol lia  perspicillata 

105,  104.5,  100 

106 

Phyllostomus  discolor 

104.5 

87.5 

Glossophaga  soricina 

111,  127 

94 

Desmodus  rotundas 

127 

100 

Sturnira  l ilium 

127 

100 

Walues  from  measurements  of  mobility  as  indicated  in  fig.  3  of  Valdivieso  and  Tamsitt  (1974).  Alb100 
is  taken  as  the  most  common  allozyme  in  A.  jamaicensis. 


12  times  as  distant  from  mainland  populations  of  this  species  as  the  latter  are 
among  themselves  (D  =  0.065  and  0.005,  respectively).  Although  this  difference 
involves  very  small  D-values  and  is  not  statistically  significant,  it  is  consistent 
with  the  view  that  Jamaican  populations  have  been  isolated  from  those  on  the 
mainland  for  some  time.  This  isolation  might  have  resulted  in  genetic  differenti¬ 
ation  of  Jamaican  populations  sufficient  to  warrant  recognizing  them  as  belong¬ 
ing  to  a  separate  subspecies,  a  conclusion  reached  by  Anderson  and  Nelson 
(1965)  based  on  morphological  analysis  of  M.  waterhousii  from  Jamaica  and 
Mexico. 

The  genetic  distance  between  species  of  Macrotus  is  substantial  (D  =  0.4);  at 
least  40  per  cent  of  the  loci  in  the  two  species  having  accumulated  codon  changes 
since  separation  from  a  common  ancestor.  This  value  is  high  for  congeneric 
species  of  mammals  and  is  near  the  value  reported  for  intergeneric  comparisons 
of  the  vespertilionids  Myotis  and  Pipistrellus  (Straney  et  al.,  1976/?).  Indeed,  this 
value  is  nearly  equal  to  that  found  separating  Glossophaga  and  Desmodus  (D  — 
0.35;  see  below),  members  of  different  phyllostomatid  subfamilies.  It  was  con¬ 
cluded  that  the  large  genetic  difference  between  M.  californicus  and  M.  water¬ 
housii  was  a  product  of  independent  evolution  during  a  long  period  of  separa¬ 
tion — current  parapatry  represents  secondary  contact.  Temporal  calibration  of 
Nei’s  D  in  phyllostomatids,  discussed  below,  suggests  that  these  species  have 
been  separated  for  approximately  10  million  years.  Yet,  during  this  time,  al¬ 
though  protein  loci  have  diverged,  morphological  change  has  been  slight  (Ander¬ 
son  and  Nelson,  1965;  Davis  and  Baker,  1974). 

The  electrophoretic  analysis  of  Macrotus  (Greenbaum  and  Baker,  1976)  clearly 
indicated  that  mainland  Macrotus  represent  two  species  and  that  Antillean 
populations  are  conspecific  with  Mexican  M.  waterhousii.  Their  study  suggests 
great  potential  for  electrophoretic  application  to  systematic  problems  on  an 
intrageneric  level.  Published  information  on  biochemical  genetics  of  phyllosto¬ 
matid  bats  establishes  the  presence  of  polymorphic  and  polytypic  genetic  variation 
in  members  of  the  family.  The  results  of  Greenbaum  and  Baker  (1976)  and 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


161 


Table  2. — Gene  loci  and  assay  systems  examined  in  Trinidad  phyllostomatids. 


Protein  System 

Buffer  system' 

pH 

Voltage 

Time  (hr.) 

a-Glycerophosphate  dehydrogenase  (a-GPD) 

Tris  citrate 

8.0 

130 

3.5 

Albumin  (ALB) 

Lithium  hydroxide 

8.1 

350 

5 

Alcohol  dehydrogenase  (ADH) 

Phosphate 

6.7 

130 

5 

Glutamic  oxaloacetic  transaminase- 1  (GOT-1)  Lithium  hydroxide 

8.1 

350 

5 

Glutamic  oxaloacetic  transaminase-2  (GOT-2)  Tris  citrate 

8.0 

130 

3.5 

Isocitrate  dehydrogenase  (IDH-1,  2) 

Tris  citrate 

6.7 

150 

5 

Indophenol  oxidase  (IPO) 

Lithium  hydroxide 

8.1 

350 

5 

Lactate  dehydrogenase- 1  (LDH-1) 

Lithium  hydroxide 

8.1 

350 

5 

Lactate  dehydrogenase-2  (LDH-2) 

Lithium  hydroxide 

8.1 

350 

5 

Malate  dehydrogenase- 1,  -2  (MDH-1,  -2) 

Tris  citrate 

6.7 

150 

5 

Phosphoglucomutase-1,  -2  (PGM-1,  -2) 

Tris  citrate 

6.7 

150 

5 

Phosphoglucose  isomerase-1,  -2  (PGI-1,  -2) 

Poulik 

8.5 

250 

3.5 

6-Phosphogluconate  dehydrogenase  (6-PGD) 

Tris  maleate 

7.4 

100 

5 

'Details  of  preparation  in  Selander  et  al.,  1971. 


Straney  et  al.  (1976a)  indicate  a  low  level  of  genetic  variation  in  the  average 
population  of  Macrotus.  The  level  of  divergence  observed  by  Greenbaum  and 
Baker  suggests  that  phyllostomatid  taxa  may  be  genetically  quite  distinct. 
New  data  collected  on  the  genetics  of  phyllostomatids  from  Trinidad,  summarized 
below,  allow  these  points  to  be  examined  in  more  detail. 

Implications  of  Genic  Variation  in  Phyllostomatids  from  Trinidad 

In  August,  1974,  we  collected  samples  of  14  species  of  phyllostomatid  bats  at 
six  localities  in  Trinidad.  Assay  systems  were  similar  to  those  described  by 
Straney  et  al.  (1976 a)  and  Greenbaum  and  Baker  (1976).  Table  2  lists  gene  loci 
examined  and  conditions  of  assays.  Several  proteins  were  examined  but  could 
not  be  interpreted  due  to  progressive  denaturation  (malic  enzyme- 1,  -2;  hemo¬ 
globin).  Esterases  presented  such  a  complex  pattern  that  it  was  not  possible  to 
establish  locus  homologies  and  these  proteins  have  been  disregarded. 

Table  3  presents  a  summary  of  gene  frequencies  in  the  populations  examined. 
In  many  cases  sample  sizes  are  quite  small  and  doubtless  some  polymorphic 
loci  were  missed.  Albumin  was,  as  mentioned  above,  the  most  polymorphic 
locus,  segregating  for  two  or  three  alleles  in  the  three  species  of  Artibeus  sampled, 
as  well  as  in  Chiroderma,  Carollia,  and  Glossophaga.  Other  loci  that  show 
relatively  high  levels  of  heterozygosity  are  1DH-1  (A.  jamaicensis  and  Anoura), 
a-GPD  (Carollia),  and  PGM-1  (Carollia).  All  other  variable  loci  either  are 
present  in  samples  too  small  to  give  fair  estimates  or  show  a  proportion  of 
heterozygotes  less  than  0.10. 


Genetics  and  Ecology 

Table  4  summarizes  heterozygosity  values  for  all  species  of  bats  thus  far 
examined.  Values  from  this  study  are  restricted  to  populations  with  sufficient 


162 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  3. —  Alleles  and  frequencies  (in  parentheses)  at  16  gene  loci  in 


Species 

Locality1 

N 

“  GPD 

Alb 

GOT-1 

GOT-2 

IDH-1 

1DH-2 

IPO 

Phyllostomatidae 

Ametrida  centurio 

5 

1 

75(1.00) 

105.5(1.00) 

83(1.00 

-100(1.00) 

800.00) 

-67(1.00) 

1000.00) 

Artibeus  cinereus 

1 

6 

100(0.92) 

62(0.08) 

102(0.92) 

101(0.08) 

56(  1 .00) 

-94(1.00) 

87(1.00) 

-67(1.00) 

2000.00) 

4 

7 

100(0.86) 

62(0.14) 

102(0.72) 

101(0.21) 

103(0.07) 

56(1.00) 

-94(1.00) 

87(1.00) 

-67(1.00) 

200(0.79) 

100(0.21) 

Artibeus  jamaicensis 

I 

9 

100(1.00) 

100(0.56) 

103(0.27) 

101(0.17) 

100(1.00) 

-1000.00) 

100(0.50) 

87(0.50) 

-100(1.00) 

1000.00) 

2 

30 

100(0.98) 

123(0.02) 

100(0.48) 

103(0.32) 

101(0.20) 

100(0.98) 

56(0.02) 

-1000.00) 

87(0.57) 

100(0.43) 

-  100(1.00) 

100(0.93) 

50(0.07) 

3 

10 

100(1.00) 

100(0.50) 

103(0.45) 

101(0.05) 

100(0.90) 

56(0.10) 

-100(1.00) 

100(0.50) 

87(0.50) 

-100(1.00) 

100(0.95) 

50(0.05) 

4 

4 

100(1.00) 

100(0.67) 

103(0.16) 

101(0.16) 

100(1.00) 

-1000.00) 

100(0.62) 

87(0.38) 

-100(1.00) 

1000.00) 

5 

4 

100(1.00) 

100(0.50) 

101(0.38) 

103(0.12) 

100(1.00) 

-1000.00) 

100(0.62) 

87(0.38) 

-  100(1.00) 

1000.00) 

Artibeus  lituratus 

1,  2,  3,  5 

7 

100(0.86 

62(0.14) 

103(0.93) 

100(0.07) 

100(0.93) 

56(0.07) 

-94(1.00) 

87(1.00) 

-67(1.00) 

2000.00) 

Chiroderma  villosuni 

5 

3 

100(0.67) 

123(0.33) 

106(1.00) 

111(0.67) 

61(0.33) 

-94(1.00) 

87(1.00) 

-67(1.00) 

1000.00) 

Sturnira  (Species  “A”) 

1,  3 

2 

108(1.00) 

98(1.00) 

56(1.00) 

-1000.00) 

50(1.00) 

-100(1.00) 

1000.00) 

Uroderma  bilob  at  um 

3,  4,  5 

5 

123(1.00) 

106(1.00) 

56(1.00) 

-500.00) 

80(0.60) 

100(0.40) 

-67(1.00) 

1000.00) 

Vampryrops  helleri 

1,  3,  4 

8 

108(1.00) 

100.5(1.00) 

56(1.00) 

-94(1.00) 

87(1.00) 

-67(1.00) 

200(0.94) 

100(0.06) 

Carollia  perspicillata 

1 

30 

123(0.73) 

146(0.24) 

108(0.03) 

105(0.98) 

104.5(0.02) 

136(0.97) 

100(0.03) 

-100(0.90) 

-50(0.10) 

67(1.00) 

-67(1.00) 

-30(1.00) 

3 

5 

123(0.80) 

146(0.20) 

105(1.00) 

136(0.90) 

100(0.10) 

- 100(0.60) 

-  50(0.40) 

67(1.00) 

-67(1.00) 

-30(1.00) 

4 

10 

123(0.75) 

108(0.15) 

146(0.10) 

105(0.95) 

100(0.05) 

136(0.85) 

100(0.15) 

-1000.00) 

67(1.00) 

-67(1.00) 

-300.00) 

5 

12 

123(0.88) 

146(0.08) 

108(0.04) 

105(0.88) 

104.5(0.08) 

100(0.04) 

136(1.00) 

-100(0.96) 

-50(0.04) 

67(1.00) 

-67(1.00) 

-300.00) 

Phyllostomus  discolor 

3 

1 

123(1.00) 

104.5(1.00) 

100(1.00) 

-125(1.00) 

77(0.50) 

60(0.50) 

-135(1.00) 

900.00) 

Phyllostomus  hast  at  us 

1,  3 

2 

123(1.00) 

104(1.00) 

17(1.00) 

-125(1.00) 

70(1.00) 

-133(1.00) 

900.00) 

Glossophaga  soricina 

1,  4,  5 

5 

108(0.90) 

123(0.10) 

107(1.00) 

56(1.00) 

-50(1.00) 

87(1.00) 

-67(1.00) 

-25(1.00) 

3 

14 

108(0.65) 

123(0.35) 

107(1.00) 

56(1.00) 

-500.00) 

87(1.00) 

-67(1.00) 

-25(1.00) 

Anoura  geoffroyi 

6 

30 

169(1.00) 

101(1.00) 

66(1.00) 

-94(1.00) 

87(0.94) 

90(0.03) 

77(0.03) 

-67(1.00) 

150(0.97) 

250(0.03) 

Desmodus  rotundus 

2.  4 

4 

123(1.00) 

107(1.00) 

56(1.00) 

-31(1.00) 

60(1.00) 

-67(1.00) 

-25(1.00) 

Molossidae 

Molossus  molossus 

2 

30 

177(1.00) 

104.1(1.00) 

63(1.00) 

-50(1.00) 

73(1.00) 

-133(1.00) 

75(1.00) 

Natal  idae 

Natalus 

6 

30 

100(1.00) 

99(0.52) 

99.5(0.48) 

5(1.00) 

-97(1.00) 

73(1.00) 

-133(1.00) 

-200(1.00) 

1.  Locality  designations  are: 

1, 

Las  Cuevas,  St.  George  Co.; 

2,  Maracas 

Valley,  2 

mi.  N  (by 

road)  St. 

Joseph,  St.  George  Co.;  3,  Guayaguayare,  Mayaro  Co.;  4,  Maracas  Valley,  12  mi.  N  (by  road)  St.  Joseph, 
St.  George  Co.;  5,  2  mi.  E,  3  mi.  S  San  Raphael,  St.  George  Co.;  6,  Tamana  Cave,  St.  Andrew  Co. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


163 


bats  from  Trinidad.  N  is  sample  size;  locus  designations  are  as  in  Table  2. 


LDH-l 

LDH-2 

MDH-1 

MDH-2 

PGM-1 

PGM-2 

PGM 

PG1-2 

6PGD 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

175(1.00) 

6000.00) 

117(1.00) 

100(1.00) 

-100(1.00) 

100(0.92) 

60(0.08) 

-100(1.00) 

240(0.58) 

100(0.42) 

-100(1.00) 

1000.00) 

1000.00) 

100(0.92) 

40(0.08) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(0.79) 

240(0.21) 

-100(1.00) 

100(1.00) 

1000.00) 

100(0.97) 

40(0.03) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

1000.00) 

1000.00) 

100(0.83) 

166(0.17) 

100(1.00) 

-100(0.98) 

-50(0.02) 

100(1.00) 

-100(1.00) 

100(0.98) 

240(0.02) 

-  100(1.00) 

100(1.00) 

1000.00) 

100(0.98) 

166(0.02) 

100(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

1000.00) 

1000.00) 

1000.00) 

100(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

1000.00) 

1000.00) 

100(0.88) 

166(0.02) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

1000.00) 

1000.00) 

1000.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

1000.00) 

1000.00) 

97(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-1000.00) 

1000.00) 

1000.00) 

1000.00) 

84(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

380(0.75) 

640(0.25) 

-183(1.00) 

2000.00) 

8000.00) 

1000.00) 

100(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-1000.00) 

1000.00) 

100(1.00) 

1000.00) 

98(1.00) 

-100(1.00) 

100(1.00) 

-  100(1.00) 

100(1.00) 

-1000.00) 

100(1.00) 

1000.00) 

1000.00) 

91(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

240(0.97) 

380(0.03) 

-1000.00) 

62(1.00) 

0(1.00) 

1000.00) 

91(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

240(1.00) 

-100(1.00) 

62(1.00) 

0(1.00) 

1000.00) 

91(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

240(0.90) 

380(0.10) 

-1000.00) 

62(1.00) 

0(1.00) 

100(1.00) 

91(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

240(0.92) 

380(0.08) 

-1000.00) 

62(1.00) 

0(1.00) 

1000.00) 

100(1.00) 

-  100(1.00) 

100(1.00) 

-100(1.00) 

100(0.50) 

240(0.50) 

-  183(1.00) 

160(1.00) 

550(1.00) 

100(1.00) 

100(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

240(0.75) 

100(0.25) 

-183(1.00) 

1600.00) 

5500.00) 

1000.00) 

84(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

225(1.00) 

-1000.00) 

206(1.00) 

1000(1.00) 

1660.00) 

84(1.00) 

-100(1.00) 

100(1.00) 

-100(1.00) 

225(1.00) 

-1000.00) 

206(  1 .00) 

1000(1.00) 

166(1.00) 

96(1.00) 

-100(1.00) 

80(1.00) 

-100(1.00) 

380(1.00) 

-1000.00) 

175(1.00) 

7000.00) 

90(1.00) 

84(1.00) 

+  100(1.00) 

100(1.00) 

-100(1.00) 

380(1.00) 

-1000.00) 

206(1.00) 

10000.00) 

1660.00) 

68(1.00) 

0(1.00) 

40(0.98) 

62(0.02) 

-133(1.00) 

38(1.00) 

-183(1.00) 

600(1.00) 

170(0.98) 

190(0.02) 

73(1.00) 

-200(1.00) 

59(1.00) 

-133(1.00) 

400(1.00) 

-2000.00) 

238(1.00) 

5000.00) 

2000.00) 

164 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


sample  size  (N>  15)  to  permit  relatively  unbaised  calculation  of  gene  frequencies. 
H-values  listed  are  those  expected  under  Hardy-Weinberg  assumptions,  from 
which  none  of  the  phyllostomatid  samples  deviate  significantly.  Johnson  (1974) 
has  suggested  that  enzymes  in  key  regulatory  positions  in  metabolic  pathways  are 
more  variable  than  those  in  nonregulatory  positions,  and  that  enzymes  with 
variable  substrates  show  highest  heterozygostities.  There  is  no  consistent  agree¬ 
ment  of  the  chiropteran  data  with  this  hypothesis  (Table  4).  Only  six  of  the  11 
species  have  higher  heterozygosities  in  regulatory  enzymes  than  in  nonregulatory 
ones.  We  agree  with  Selander  (1976)  that  Johnson’s  hypothesis  will  not  in  itself 
account  for  heterozygosity  differences  seen  between  loci.  Unfortunately,  John¬ 
son’s  hypothesis  does  not  deal  with  general  protein  loci,  which  we  found  to  be  the 
most  variable  in  the  phyllostomatids.  General  proteins  usually  have  exhibited 
low  levels  of  polymorphism  in  other  mammals  (Selander,  1976). 

The  data  presented  in  Table  4  suggest  that  phyllostomatids  differ  from  species 
of  Myotis  in  having  lower  levels  of  genic  heterozygosity.  The  frequency  dis¬ 
tributions  of  per  locus  heterozygosity  (h)  differ  between  these  groups  (Fig.  1). 
The  average  locus  in  the  phyllostomatids  examined  has  a  heterozygosity  of  0.036, 
whereas  for  Myotis  this  value  is  0.1 17.  The  pattern  seen  among  phyllostomatids 
is  very  near  to  that  observed  in  a  variety  of  rodents  (Fig.  1 ;  data  for  esterases  are 
excluded  from  this  figure).  When  the  h-\ alues  for  phyllostomatids  and  Myotis 
are  compared  in  an  Analysis  of  Variance  (after  arcsine  transformation),  the 
difference  is  highly  significant  (P<  0.001).  Phyllostomatids  possess  more 
monomorphic  loci  than  do  species  of  Myotis  and  do  not  show  a  second  frequency 
peak  for  loci  with  high  heterozygosity.  Most  of  the  loci  contributing  to  this  second 
peak  in  Myotis  are  not  in  Hardy-Weinberg  equilibrium  (Straney  et  al.,  1976 a). 

The  patterns  in  Fig.  1  suggest  that  phyllostomatids  might  have  levels  of 
heterozygosity  equivalent  to  those  observed  in  rodents.  This  is  not  apparent  when 
H-values  presented  here  are  compared  (average  H  for  rodents,  0.059),  because 
esterases  account  for  43  per  cent  of  rodent  H-values  (Selander,  1976).  Some 
Myotis  populations,  however,  are  more  similar  in  heterozygosity  levels  to  in¬ 
vertebrates  (average  H  =  0.12;  Selander,  1976),  but  this  is  not  true  of  vespertilio- 
nids  as  a  group.  Pipistrellus  populations  exhibit  low  genetic  variability,  and  it  has 
been  suggested  that  this  results  from  demographic  factors  (Table  4;  Straney 
et  al.,  19766).  Preliminary  data  on  California  vespertilionids  indicate  that  other 
species  also  have  low  variability  (J.  L.  Patton,  personal  communication). 

Levels  of  genic  variability  in  phyllostomatids,  and  at  least  some  species  of 
Myotis,  differ  greatly,  and  it  is  likely  that  other  evolutionary  characteristics  do 
as  well.  A  number  of  factors  could  produce  the  differences  in  heterozygosity  ob¬ 
served  between  phyllostomatids  and  Myotis-.  stochastic  processes,  gene  flow, 
adaptation  to  microgeographic  conditions,  and  the  grain  of  experienced  environ¬ 
ments  (Levins,  1968;  Soule,  1976).  Differences,  on  a  much  lower  level,  also  are 
apparent  within  the  phyllostomatids  examined  (Table  4).  Artibeus,  Glossophaga, 
and  Carollia,  the  most  common  species  in  our  collections,  differ  greatly  in  levels 
of  polymorphism.  Population  bottlenecks,  inbreeding,  and  drift  are  not  suffi- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


165 


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166 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  1. — Per  cent  occurrence  of  loci  with  different  levels  of  heterozygosity  in  rodents 
(summarized  in  Smith  et  al,  1978),  phyllostomatid  bats  (this  study),  and  bats  of  the  genus 
Myotis  (Table  4).  N  is  number  of  individuals. 

cient  to  explain  the  low  levels  of  heterozygosity  in  Glossophaga,  Anoura,  and 
Carollia,  compared  to  the  relatively  high  levels  in  Artibeus,  because  all  four 
genera  are  widespread,  highly  vagile,  and  abundant.  Isolation  of  the  population 
characteristics  that  might  be  responsible  for  differences  in  heterozygosity  is  not 
possible  using  genetic  data  alone.  Only  genetic  studies  coupled  with  extensive 
ecological  investigations  will  provide  the  information  needed  to  address  this 
point,  and  then  only  if  temporal  trends  also  are  examined. 

Differences  in  heterozygosity  may  index  more  subtle  differences  in  population 
characteristics.  The  data  presented  above  suggest  that  different  species  of  bats 
have  been  exposed  to  different  evolutionary  forces,  which  are  dictated  by 
differences  in  population  structure.  Although  we  are  unable  at  this  point  to 
determine  why  variation  in  population  structure  exists  or  what  evolutionary 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


167 


forces  effect  these  differences,  it  is  clear  that  genetic  models  of  chiropteran  pop¬ 
ulations  must  account  for  several  distinct  patterns  of  variation. 

Future  studies  of  ecological  genetics  in  bats  should  pay  particular  attention  to 
spatiotemporal  structure  of  populations.  With  proper  experimental  design,  it  is 
possible  to  estimate  deme  size,  effective  population  size,  and  migration  rate  using, 
for  example,  Kirby’s  (1976)  analysis  of  Wright’s  F-statistics.  More  important 
than  estimates  of  these  values,  though,  is  an  estimate  of  their  variability  through 
time.  Bat  populations  are  conceivably  temporally  unstable  in  composition,  due 
in  part  to  their  vagility  and  roosting  habits.  Turner’s  (1975)  studies  of  Desmodus 
in  Costa  Rica  indicate  that  vampire  populations  can  be  either  ephemeral  or 
relatively  stable  depending  on  where  the  bats  roost.  It  is  important  to  know  on 
what  scale  this  temporal  variability  acts  as  well  as  which  ecological  factors,  such 
as  roost  site,  can  alter  its  periodicity.  Species  differences  in  these  parameters  are 
to  be  expected  in  a  group  as  diverse  as  the  phyllostomatids,  and  comparative 
studies  will  be  necessary  to  indicate  to  what  degree  morphological  and  ecological 
diversity  is  reflected  in  population  structure.  The  evolutionary  process  proceeds 
only  within  the  limits  set  by  the  spatiotemporal  structure  of  the  populations 
involved.  Hence,  a  useful  approach  to  understanding  patterns  of  population 
differentiation,  speciation,  and  phyletic  evolution  in  different  lineages  is  to 
determine  to  what  extent  structural  differences  in  populations  determine  different 
evolutionary  strategies.  Structural  parameters  of  populations  are  major  deter¬ 
minants  of  the  fate  of  new  mutants,  the  permanence  of  polymorphisms,  and  the 
speed  with  which  adaptive  change  can  be  effected. 

Genetic  Phyletics 

An  alternative  to  using  traditional  characteristics  for  reconstructing  the 
evolutionary  history  of  a  group  is  to  employ  measures  of  genetic  comparisons 
between  taxa.  Because  evolution  can  be  expressed  as  the  change  in  genomes 
through  time,  genetic  comparisons  can  be  used  to  estimate  the  degree  of  divergence 
between  taxa.  With  the  advent  of  biochemical  assay  systems  this  has  become 
possible.  As  genetic  comparisons  dependent  upon  breeding  studies  cannot  be 
used  to  compare  taxa  above  the  species  level  in  most  mammals,  the  early  interest 
in  electrophoresis  of  bat  proteins  was,  in  part,  systematic. 

Manwell  and  Kerst  (1966),  Valdivieso  et  al.  (1969),  Valdivieso  and  Tamsitt 
(1974),  Tamsitt  and  Valdivieso  (1969),  and  Mitchell  (1970),  all  working  with 
one  or  at  most  a  few  proteins,  concluded  that  electrophoretic  comparisons  would 
be  of  little  use  in  chiropteran  systematics  below  the  family  level.  These  studies 
did,  however,  find  confirming  evidence  for  placing  the  mormoopids 
( Pteronotus  and  Mormoops )  into  a  family  separate  from  phyllostomatids  and 
for  the  inclusion  of  the  vampires  as  a  subfamily  in  the  Phyllostomatidae.  However, 
phylogenetic  conclusions  based  on  a  few  biochemical  characters  cannot 
be  expected  to  be  any  more  accurate  than  those  based  on  a  few  morphological 
characters  (Avise  et  al.,  1974).  Biochemical  data  used  to  indicate  phylogenetic 
relationships  are  based  on  the  assumption  that  the  loci  sampled  are  representative 


168 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


+ 

1001- 
98  - 
97 


LDH-I 


0 


+ 


0 


:§ 

£ 


§ 


$ 


Fig.  2. — Diagramatic  representation  of  banding  patterns  of  four  protein  gene  loci  in  seven 
species  of  stenodermine  bats. 


of  the  genome  as  a  whole.  The  magnitude  of  sampling  error,  and  the  resolving 
power  of  genetic  divergence  estimates,  is  a  direct  function  of  the  number  of  loci 
examined  (Nei,  1976).  Thus,  electrophoretic  comparisons  utilizing  only  a  few 
loci  provide  data  that  must  be  approached  with  caution. 

It  is  possible  that,  with  a  small  group  of  closely  related  taxa,  biochemical  data 
for  a  few  loci  will  give  quite  useful  information.  The  utility  of  this  information, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


169 


Anoura 


Fig.  3. — Wagner  tree  calculated  from  Nei’s  D  (X  100).  Numbers  are  the  amount  of  diver¬ 
gence  between  branch  points  and  represent  the  minimum  number  of  net  nucleotide  changes 
per  100  loci  accumulated  along  the  connecting  branch. 


however,  will  depend  on  the  sample  of  loci  examined.  Fig.  2  illustrates  banding 
patterns  of  four  gene  products  for  the  seven  species  of  stenodermine  bats  we  have 
examined  from  Trinidad.  Although  these  four  loci  are  sufficient  to  identify  all 
seven  species  electrophoretically,  they  are  insufficient  for  calculation  of  genetic 
distance  values,  because  D-v alues  have  large  errors  when  based  on  only  a  few 
loci  (Nei  and  Roychoudhury,  1974). 

In  Fig.  3  we  present  a  phylogenetic  estimate  of  the  relationships  among  14 
phyllostomatid  species,  based  on  the  examination  of  17  gene  loci.  The  genetic 
distances  between  taxa,  upon  which  this  tree  is  based,  are  summarized  in  Table  5. 
Seventeen  loci  certainly  are  only  a  small  fraction  of  the  phyllostomatid  genome. 
The  sampling  error  associated  with  these  divergence  values  is  not  small  (Nei, 
1 976),  and  the  tree  in  Fig.  3  must  be  evaluated  in  this  light.  It  also  should  be  pointed 
out  that  this  technique  overestimates  similarity,  and  additional  refinement  and 
the  inclusion  of  loci  such  as  esterases  should  reveal  further  separation  of  taxa. 
We  present  these  preliminary  data  as  a  starting  point  for  additional  work. 

Farris’  (1972)  modified  Wagner  algorithm  for  Nei’s  distance  was  used  to 
construct  the  tree  in  Fig.  3.  This  method  does  not  assume  that  evolutionary  rates 
are  the  same  in  all  lines  of  descent,  as  does  the  use  of  an  unweighted  pair-group 
method  for  constructing  phenograms.  The  modified  Wagner  method  partitions 
the  genetic  distance  between  taxa  into  branch  lengths  of  the  paths  connecting 
them.  This  is  done  in  such  a  way  that  the  resulting  estimates  of  branch  lengths  are 


170 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


minimum  estimates  of  the  amount  of  change  between  cladistic  events.  Because  the 
tree  is  based  on  Nei’s  D,  the  branch  lengths  can  also  be  interpreted  as  the  minimum 
number  of  net  codon  changes  per  locus  since  a  particular  cladistic  event.  Thus, 
according  to  Fig.  3,  Artibeus  cinereus  and  Vampyrops  share  an  immediate 
common  ancestor.  Since  the  cladistic  splitting  of  the  two,  we  estimate  that 
Vampyrops  has  accumulated  22  net  codon  changes  per  100  loci,  whereas  A. 
cinereus  has  accumulated  a  minimum  of  three.  Because  these  taxa  share  a  common 
ancestor,  the  difference  in  divergence  is  also  a  difference  in  evolutionary  rate 
along  the  two  branches.  One  of  the  striking  characteristics  of  the  tree  in  Fig.  3  is 
that  the  branch  lengths  are  unequal,  implying  that  the  rates  of  evolution  have  not 
been  the  same  in  all  lines  of  descent.  This  is  consistent  with  the  argument  given 
above  that  differences  in  levels  of  genetic  variability  within  phyllostomatid  species 
mirror  underlying  differences  in  population  structure,  thereby  differentially 
affecting  evolutionary  potential. 

The  root  in  Fig.  3  has  been  placed  using  Farris’  (1972)  minimum  variance 
criterion.  This  is  an  iterative  procedure  whereby  the  root  is  placed  in  the  position 
that  minimizes  the  variance  in  divergences  of  terminal  taxa  from  the  hypothetical 
ancestor  of  the  group  as  a  whole.  There  are  three  major  lineages  apparent  when 
the  root  is  placed:  1)  stenodermines,  2)  Phyllostomus  and  Carollia,  and  3)  glos- 
sophagines,  Desmodus,  and  Sturnira.  The  average  divergence  of  these  three 
lineages  from  the  ancestor  is  similar  (mean,  86,  76,  and  85  codon  changes  per 
100  loci,  respectively).  An  analysis  of  variance  of  within  and  between  lineage 
effects  on  divergence  indicates  that  100  per  cent  of  the  variance  in  divergence 
present  in  Fig.  3  is  within  lineages.  As  we  can  demonstrate  no  differences  in 
evolutionary  rate  between  lineages,  we  can  use  the  average  divergence  of  the 
lineages  (82  codon  changes  per  100  loci)  to  estimate  the  age  of  the  family.  Nei’s 
D  is  a  linear  function  of  time  (Nei,  1976),  and  studies  by  Avise  and  Ayala 
(1975,  1976)  indicate  that  genetic  distance  is  by  and  large  independent  of 
cladistic  history.  Sarich  (1977)  has  calibrated  Nei’s  D  against  his  albumin  clock 
estimate  of  divergence  time  and  has  provided  us  with  the  conversion  equation 
1.0  D—  28  million  years  (for  branch  length,  1.00  =  56  million  years).  Using  this 
conversion,  we  estimate  that  the  diversification  of  the  family  occurred  40  mil¬ 
lion  years  ago  during  the  early  Oligocene.  Because  this  is  a  minimum  estimate  of 
age,  the  age  estimated  is  of  diversification  not  origin,  and  the  estimate  is  not 
without  sampling  error,  we  feel  that  these  data  are  comparable  with  Koopman’s 
(1976)  and  Smith’s  (1976)  conclusion  that  the  late  Oligocene  is  the  latest  that 
the  family  could  have  arisen. 

Within  the  error  of  our  estimates,  the  lineages  represented  in  Fig.  3  appear 
to  have  arisen  at  the  same  time.  These  lineages  are  not  well  defined,  except  for  the 
relatively  compact  stenodermine  lineage,  and  there  is  no  evidence  of  a  “  Macrotus- 
like”  and  “ Phy l lostom ws-like”  (Smith,  1976)  dichotomy  within  our  sample. 
Genera  hypothesized  as  belonging  to  one  lineage  or  the  other  are  intermixed  in 
Fig.  3  (compare  Smith,  1976,  fig.  2).  Even  though  our  inability  to  distinguish 
this  dichotomy  may  be  an  artifact  of  sampling,  we  think  it  best  to  assume  that  the 
major  adaptive  trends  within  the  family  are  of  coeval  origin. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


171 


Stenodermines 

The  discreteness  of  the  stenodermine  lineage  in  Fig.  3  probably  results  from 
more  extensive  sampling  of  members  of  this  subfamily.  The  radiation  of  this 
group  appears  to  be  an  early  one,  the  line  leading  to  Ametrida  diverged  perhaps 
20  million  years  ago  in  the  late  Miocene.  Artibeus  is  a  basal  taxon  for  the  rest 
of  the  subfamily  represented  here  and  two  separate  lineages  derive  from  it.  The 
three  species  of  Artibeus  have  undergone  little  divergence  from  their  respective 
common  ancestors  whereas  the  two  lineages  involving  Vampyrops  and  Uroderma- 
Chiroderma  have  evolved  at  a  much  faster  rate.  These  results  suggest  that 
Artibeus  is  a  paraphyletic  taxon. 

With  effort,  it  is  possible  to  identify  Smith’s  (1976)  “short-faced,  long-faced” 
dichotomy  in  our  phylogram.  The  “long-faced”  lineage  is  polyphyletic  in  our 
reconstruction  although  the  three  members  of  this  group  (  Vampyrops,  Uroderma, 
and  Chiroderma )  are  derived  from  a  single  genus,  Artibeus.  Furthermore,  our 
phylogenetic  hypothesis  suggests  that  short-faced  is  the  primitive  condition  for 
stenodermines.  We  have  examined  too  few  genera  to  be  certain  of  this  point, 
but  the  data  at  hand  indicate  that  long  faces  represent  parallel  derived  characters. 

Our  sample  of  stenodermine  taxa,  however,  is  sufficient  to  suggest  a  polarity 
for  Baker’s  (1973)  phylogeny  of  the  subfamily  based  on  gross  karyotypic 
characters.  His  fig.  5  is  quite  similar  to  our  Fig.  3  if  the  root  of  his  phylogram  is 
displaced  to  the  right  and  if  one  ignores  the  absence  of  Sturnira.  Karyotypically, 
Chiroderma  and  Uroderma  are  not  related  as  closely  to  each  other  as  elec¬ 
trophoretic  data  indicate;  further  study  could  identify  additional  areas  of  dis¬ 
agreement.  It  is,  however,  reassuring  to  find  the  same  basic  phylogenetic  frame¬ 
work  emerging  from  two  different  and  independent  data  sources. 

Phyllostomus  and  Carollia 

There  is  little  that  can  be  said  of  the  association  of  Carollia  and  Phyllostomus 
presented  in  Fig.  3.  These  two  genera  are  not  closely  related  but  probably  do 
represent  a  distinct  lineage  within  the  family.  Walton  and  Walton  (1968)  suggested 
a  similar  relationship  based  on  their  study  of  postcranial  osteology.  There  is  no 
indication  in  our  data  of  close  phylogenetic  ties  between  Carollia  and  Glossophaga 
(sensu  Smith,  1976). 

The  divergence  of  the  two  species  of  Phyllostomus  appears  to  have  occurred  8 
million  years  ago  during  the  mid-Pliocene  (D=0.29).  The  morphological  and 
ecological  differences  between  P.  hastatus  and  P.  discolor  are  much  greater  than 
those  between  the  two  species  of  Macrotus  studied  by  Greenbaum  and  Baker 
(1976),  even  though  the  latter  are  separated  by  a  greater  genetic  distance  (D  = 
0.41-0.50).  This  represents  another  of  the  growing  number  of  cases  where  genetic 
and  morphological  measures  of  divergence  are  found  to  be  discordant  (King 
and  Wilson,  1975;  Avise,  1976). 

Glossophagines,  Desmodus,  and  Sturnira 

This  group  forms  the  most  heterogeneous  branch  of  our  phylogenetic  tree,  and 
the  relationships  within  it  are  difficult  to  reconcile  with  morphological  evidence 


172 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


and  current  concepts  of  phyllostomatid  systematics.  Anoura  and  Glossophaga  are 
somewhat  closely  related,  based  on  electrophoretic  data,  although  this  association 
is  overshadowed  by  the  greater  amount  of  protein  evolution  along  the  Anoura 
branch.  Our  placement  of  Sturnira  is  at  variance  with  current  taxonomic  opinion. 
Walton  and  Walton  (1968)  postulated  a  relationship  between  Sturnira  and  the 
glossophagines,  following  a  comparison  of  postcranial  morphology.  Addition 
of  more  genera  to  this  data  set  would  not  result  in  a  closer  association  of  Sturnira 
and  the  stenodermines  because  additional  data  would  not  decrease  the  large 
genetic  distances  between  these  groups  (Table  5).  Based  on  our  electrophoretic 
sample,  we  are  left  with  the  conclusion  that  Sturnira  is  not  genetically  a  stenoder- 
mine  bat  and  is  not  closely  related  to  any  one  of  the  lineages  represented  in  this 
study. 

A  close  relationship  between  Desmodus  and  the  glossophagines,  based  on 
chromosomal,  immunological,  and  sperm  morphology  data,  was  proposed  by 
Forman  et  al.  (1968).  Our  data  also  suggest  such  a  relationship  between  Glos¬ 
sophaga  and  Desmodus  (Fig.  3;  Table  5).  Because  of  the  difference  in  evolution¬ 
ary  rates  along  the  two  branches,  it  is  difficult  to  estimate  the  age  of  this  diver¬ 
gence,  but  we  suggest  that  it  is  10  million  years.  This  is  consistent  with  the  fossil 
record  to  the  extent  that  fossil  desmodontines  are  not  known  prior  to  about  1.5 
million  years  bp  (Hutchison,  1967). 

An  overview  of  the  genic  and  morphological  data  from  this  family  suggests 
that  there  are  several  examples  where  there  is  discordance  in  the  rates  of  evolu¬ 
tion  of  genic  and  classical  morphological  characters.  One  hypothesis  that  at¬ 
tempts  to  reconcile  genetic  and  morphological  data  assumes  that  the  morphologi¬ 
cal  modifications  leading  to  a  specialized  taxon  have  been  due  to  changes  in 
regulatory  genes  affecting  developmental  pathways.  Such  changes,  which  one 
would  not  expect  to  be  reflected  in  the  structural  genes  assayed  in  electrophoresis, 
could  result  in  major  and  rapid  morphological  evolution.  This  form  of  quantum 
evolution  ( sensu  Simpson,  1953)  has  recently  been  invoked  by  King  and  Wilson 
(1975)  to  explain  the  small  genetic  distance  between  Homo  and  Pan.  If  this  hypo¬ 
thesis  reflects  the  true  path  of  evolution  followed  in  these  discordant  examples, 
we  would  predict,  following  King  and  Wilson  (1975),  that  DNA  hybridization 
between  such  taxa  would  show  similarity  in  the  unique  DNA  fraction  consistent 
with  that  found  electrophoretically  and  a  larger  difference  in  the  presumably 
regulatory  medium  repeated  DNA  fraction. 

Phylogenetic  reconstruction  is  as  much  a  science  as  it  has  been  portrayed  an 
art.  One  proceeds  by  constructing  hypotheses  of  relationships  from  different 
data  sources  and  searching  for  one  that  subsumes  the  others  and  provides  an 
explanation  of  their  differences.  This  consistent  hypothesis  is  accepted  as  “true” 
either  until  a  more  general  one  is  produced  or  conflicting  data  are  found.  The 
phylogenetic  hypotheses  of  Smith  (1976)  and  those  reflected  by  the  checklist  of 
Jones  and  Carter  (1976)  are  not  in  accordance  with  the  genetic  relationships 
indicated  by  our  electrophoretic  data.  We  do  not  view  these  electrophoretic 
results  as  a  procrustean  bed  of  truth  into  which  the  morphological  evidence 
must  be  forced  in  agreement.  Rather,  they  generate  a  phylogenetic  hypothesis 


Table  5. — Nei’s  genetic  distance  (D,  upper  half  matrix)  and  Rogers’  genetic  similarity  (S,  lower  half  matrix)  for  hat  populations  from  Trinidad. 
Where  more  than  one  population  of  a  species  is  listed,  numerical  designations  are  as  in  Table  3.  I  indicates  an  infinite  value  for  D  ( Nei's  genetic 

identity  1=0.00). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


173 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


sufficiently  different  from  others  that  have  been  proposed  to  indicate  that  “the 
great  deal  of  uncertainty  and  contradictory  evidence”  (Smith,  1976)  surrounding 
phyllostomatid  phylogency  will  continue  in  the  future.  We  still  lack  an  hypothesis 
of  the  phylogeny  of  the  Phyllostomatidae  that  is  consistent  with  available  data 
and  that  also  identifies  the  evolutionary  processes  producing  the  differences 
between  morphological  and  genetic  findings. 

[Note  added  in  galley. — Additional  work  by  us  suggests  that  the  distance  we 
report  between  Glossophaga  and  Desmodus  is  too  low.  Examination  of  new  ma¬ 
terial,  both  at  Lubbock  and  Berkeley,  shows  that  Desmodus  and  Glossophaga 
share  very  few  alleles.  ] 


Conclusions 

Biochemical  genetics  has  proven  valuable  in  evolutionary  biology  through  the 
characterization  of  population  structure  in  space  and  time  and  through  generation 
of  phylogenetic  hypotheses.  By  examining  the  genetic  structure  of  populations, 
important  evolutionary  parameters  can  be  identified  and  quantified  to  provide  a 
bridge  between  genetic  phylogenies  and  more  traditional  evolutionary  recon¬ 
structions.  The  study  of  chiropteran  genetics  is  only  10  years  old;  yet,  in  that  time 
it  has  provided  information  that  both  challenges  and  supports  the  traditional  view 
of  chiropteran  evolution.  The  dynamics  of  population  structure  of  vespertilionid 
and  phyllostomatid  bats  does  not  appear  to  be  the  same,  although  studies  of 
temporal  structure  will  be  necessary  to  confirm  this  conclusion.  The  mode  of 
evolution,  as  reflected  by  electrophoretic  parameters,  appears  to  be  different 
between  some  lineages  of  phyllostomatids,  particularly  the  desmodontines.  When 
more  genetic  data  are  available  on  phyllostomatid  bats,  an  integration  of  genetic, 
karyotypic,  and  morphological  data  should  produce  a  consistent  model  of 
evolution  in  this  group,  which  might  be  surprising  in  its  complexity. 

Literature  Cited 

Anderson,  S.,  and  C.  E.  Nelson.  1965.  A  systematic  revision  of  Mcicrotus  (Chiroptera). 
Amer.  Mus.  Novit.,  2212:1-39. 

Avise,  J.  C.  1975.  Systematic  value  of  electrophoretic  data.  Syst.  Zool.,  23:465-481. 

- .  1976.  Genetic  differentiation  during  speciation.  Pp.  106-122,  in  Molecular 

Evolution  (F.  J.  Ayala,  ed.),  Sinauer  Associates,  Sunderland,  Mass.,  x  +  277  pp. 
Avise,  J.  C.,  and  F.  J.  Ayala.  1975.  Genetic  change  and  rates  of  cladogenesis.  Genetics, 
81:757-773. 

- .  1976.  Genetic  differentiation  in  speciose  versus  depauperate  phylads:  evidence 

from  the  California  minnows.  Evolution,  30:46-58. 

Avise,  J.  C.,  M.  H.  Smith,  and  R.  K.  Selander.  1974.  Biochemical  polymorphism  and 
systematics  in  the  genus  Peromyscus.  VI.  The  boylii  species  group.  J.  Mamm., 
55:751-763. 

Baker,  R.  J.  1973.  Comparative  cytogenetics  of  new  world  leaf-nosed  bats  (Phyl¬ 
lostomatidae).  Period.  Biol.,  75:37-45. 

Crow,  J.  F.,  and  M.  Kimura.  1970.  An  introduction  to  population  genetics  theory. 
Harper  and  Row,  New  York,  xiv  +  591  pp. 

Davis,  B.  L.,  and  R.  J.  Baker.  1974.  Morphometries,  evolution  and  cytotaxonomy  of 
mainland  bats  of  the  genus  Mcicrotus  (Chiroptera:  Phyllostomatidae).  Syst.  Zool., 
23:26-39. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


175 


Dobzhansky,  T.  1970.  Genetics  of  the  evolutionary  process.  Columbia  Univ.  Press, 
New  York,  xi  +  505  pp. 

Ferris,  J.  S.  1972.  Estimating  phylogenetic  trees  from  distance  matrices.  Amer.  Nat., 
106:645-668. 

Forman,  G.  L.,  R.  J.  Baker,  and  J.  D.  Gerber.  1968.  Comments  on  the  systematic 
status  of  vampire  bats  (Family  Desmodontidae).  Syst.  Zool.,  17:417-425. 

Greenbaum,  I.  F.,  and  R.  J.  Baker.  1976.  Evolutionary  relationships  in  Macrotus 
(Mammalia:  Chiroptera):  biochemical  variation  and  karyology.  Syst.  Zool., 
25:15-25. 

Greenhall,  A.  M.  1976.  Care  in  captivity.  Pp.  89-131,  in  Biology  of  bats  of  the  New 
World  family  Phyllostomatidae.  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C. 
Carter,  eds.),  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  10:1-218. 

Hutchison,  J.  H.  1967.  A  pleistocene  vampire  bat  ( Desmodus  stock i)  from  Potter  Creek 
Cave,  Shasta  County,  California.  Paleobios,  3:1-6. 

Johnson,  G.  B.  1974.  Enzyme  polymorphism  and  metabolism.  Science,  184:28-37. 

Jones,  J.  K.,  Jr.,  and  D.  C.  Carter.  1976.  Annotated  checklist,  with  keys  to  subfamilies 
and  genera.  Pp.  7-38,  in  Biology  of  bats  of  the  New  World  family  Phyllostomatidae. 
Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.),  Spec.  Publ.  Mus., 
Texas  Tech  Univ.,  10:1-218. 

King,  M.  C.,  and  A.  C.  Wilson.  1975.  Evolution  at  two  levels  in  humans  and  chim¬ 
panzees.  Science,  188:107-1 16. 

Kirby,  G.  C.  1976.  Heterozygote  frequencies  in  small  populations.  Theoret.  Pop. 
Biol.,  8:31-48. 

Koopman,  K.  F.  1976.  Zoogeography.  Pp.  39-47,  in  Biology  of  bats  of  the  New  World 
family  Phyllostomatidae.  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter, 
eds.),  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  10:1-218. 

Levins,  R.  1968.  Evolution  in  changing  environments.  Princeton  Univ.  Press,  Prince¬ 
ton,  New  Jersey,  x  +  120  pp. 

Lewontin,  R.  C.  1974.  The  genetic  basis  of  evolutionary  change.  Columbia  Univ. 
Press,  New  York,  xiii  +  346  pp. 

Manwell,  C.,  and  K.  V.  Kerst.  1966.  Possibilities  of  biochemical  taxonomy  of  bats 
using  hemoglobin,  lactate  dehydrogenase,  esterases  and  other  proteins.  Comp. 
Biochem.  Physiol.,  17:741-754. 

Maxon,  L.  R.,  and  A.  C.  Wilson.  1974.  Convergent  morphological  evolution  detected  by 
studying  proteins  of  tree  frogs  in  the  Hyla  eximia  group.  Science,  185:66-68. 

Miller,  G.  S.  1907.  The  families  and  genera  of  bats.  Bull.  U.S.  Nat.  Mus.,  57:1-282. 

Mitchell,  G.  C.  1970.  An  electrophoretic  comparison  of  hemoglobins  in  bats.  Comp. 
Biochem.  Physiol.,  35:667-677. 

Mitchell,  H.  A.  1966.  Multiple  haemoglobins  in  bats.  Nature,  210: 1067-1068. 

Nei,  M.  1976.  Mathematical  models  of  speciation  and  genetic  distance.  Pp.  723-766, 
in  Population  genetics  and  ecology  (S.  Karlin  and  E.  Nevo,  eds.).  Academic  Press, 
New  York,  xiv  +  832  pp. 

Nei,  M.,  and  A.  K.  Roychoudhury.  1974.  Sampling  variances  of  heterozygosity  and 
genetic  distance.  Genetics,  76:379-390. 

Powell,  J.  R.  1975.  Protein  variation  in  natural  populations  of  animals.  Ann.  Rev. 
Ecol.  Syst.,  6:79-119. 

Rasmussen,  D.  I.  1968.  Genetics.  Pp.  340-372,  in  Biology  of  Peromyscus  (Rodentia) 
(J.  A.  King,  ed.),  Amer.  Soc.  Mammal.  Spec.  Publ.,  2:ii  +  593  pp. 

Rogers,  J.  S.  1972.  Measure  of  genetic  similarity  and  genetic  distance.  Univ.  Texas 
Publ.,  7213:145-153. 

Sarich,  V.  M.  1977.  Rates,  sample  sizes  and  the  neutrality  hypothesis  for  electrophoresis 
in  evolutionary  studies.  Nature,  265:24-28. 

Selander,  R.  K.  1976.  Genetic  variation  in  natural  populations.  Pp.  21-45,  in  Molecular 
evolution  (F.  J.  Ayala,  ed.),  Sinauer  Associates,  Sunderland,  Mass.,  x-f-277  pp. 


176 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Selander,  R.  K.,  and  D.  W.  Kaufman.  1973.  Genetic  variability  and  strategies  of 
adaptation  in  animals.  Proc.  Nat.  Acad.  Sci.,  U.S.A.,  70:1875-1877. 

Selander,  R.  K.,  M.  H.  Smith,  S.  Y.  Yang,  W.  E.  Johnson,  and  J.  B.  Gentry.  1971.  Bio¬ 
chemical  polymorphism  and  systematics  in  the  genus  Peroymscus.  I.  Variation  in 
the  old-field  mouse  ( Peromyscus  polinotus).  Studies  in  Genetics  VI,  Univ.  Texas 
Publ.,  7103:49-90. 

Simpson,  G.  G.  1953.  The  Major  Features  of  Evolution.  Colombia  Univ.  Press,  New  York, 
xx  +  434  pp. 

Smith,  J.  D.  1972.  Systematics  of  the  chiropteran  family  Mormoopidae.  Misc.  Publ. 
Mus.  Nat.  Hist.,  Univ.  Kansas,  56:1-132. 

- .  1976.  Chiropteran  evolution.  Pp.  49-69,  in  Biology  of  bats  of  the  New  World 

family  Phyllostomatidae.  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter, 
eds.).  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  10:1-218. 

Smith,  M.  H.,  M.  N.  Manlove,  and  J.  Joule.  1978.  Genetic  organization  in  space 
and  time.  Pp.  99-113,  in  Populations  of  small  mammals  under  natural  conditions 
(D.  P.  Snyder,  ed.).  Spec.  Publ.  Pymatuning  Lab.  Ecol.,  Univ.  Pittsburgh. 

Soule,  M.  1976.  Allozyme  variation:  its  determinants  in  space  and  time.  Pp.  60-77, 
in  Molecular  evolution  (F.  J.  Ayala,  ed.),  Sinauer  Associates,  Sunderland,  Mass., 
x  +  277  pp. 

Straney,  D.  O.,  M.  H.  Smith,  R.  J.  Baker,  and  I.  F.  Greenbaum.  1976a.  Biochemical 
variation  and  genic  similarity  in  Myotis  velifer  and  Macrotus  californicus.  Comp. 
Biochem.  Physiol.,  54B:243-248. 

Straney,  D.  O.,  M.  J.  O'Farrell,  and  M.  H.  Smith.  19766.  Biochemical  genetics  of 
Myotis  californicus  and  Pipistrellus  hesperus  from  southern  Nevada.  Mammalia, 
40:344-347. 

Tamsitt,  J.  R.,  and  D.  Valdivieso.  1969.  Hemoglobin  electrophoresis  in  the  systematics 
of  bats  (Microchiroptera).  Occas.  Papers  Life  Sci.,  Royal  Ontario  Mus.,  14:1-12. 

Turner,  D.  C.  1975.  The  vampire  bat.  Johns  Hopkins  Univ.  Press,  Baltimore,  x+  145  pp. 

Valdivieso,  D.,  and  J.  R.  Tamsitt.  1974.  Electrophoretic  patterns  of  serum  proteins  of 
neotropical  bats  (Chiroptera).  Contrib.  Life  Sci.,  Royal  Ontario  Mus.,  98:1-24. 

Valdivieso,  D„  J.  R.  Tamsitt,  and  E.  Conde-de  Pino.  1969.  Electrophoretic  properties 
of  neotropical  bat  hemoglobins.  Comp.  Biochem.  Physiol.,  30:117-122. 

Walton,  D.  W.,  and  G.  W.  Walton.  1968.  Comparative  osteology  of  the  pelvic  and 
pectoral  girdles  of  the  Phyllostomatidae  (Chiroptera:  Mammalia).  J.  Grad. 
Res.  Center,  Southern  Methodist  Univ.,  37:1-35. 


SPERM  MORPHOLOGY 


G.  Lawrence  Forman  and  Hugh  H.  Genoways 


Bishop  and  Austin  (1957)  in  their  study  of  variation  in  mammalian  spermatozoa 
suggested  that  the  sperm  of  each  mammalian  species  was  probably  unique. 
Although  complete  volumes  have  been  written  on  the  ultrastructure  of  spermatozoa 
(for  example  Baccetti,  1970),  particularly  of  humans  and  domestic  animals, 
there  is  still  relatively  little  information  available  on  the  comparative  gross 
morphology  of  spermatozoa.  McFarlane  (1963),  Forman  (1968),  and  Forman 
et  al.  (1968)  made  significant  contributions  to  our  understanding  of  the  use  of 
sperm  morphology  in  establishing  systematic  and  phylogenetic  relationships  of 
birds  and  mammals.  However,  there  have  been  very  few  similar  studies  published 
to  this  date. 

The  use  of  sperm  morphology  as  a  systematic  character  among  mammals,  is 
relatively  new,  beginning  with  the  study  of  British  murid  rodents  by  Friend  (1936) 
Other  studies  dealing  with  rodent  sperm  include  those  of  Braden  (1959),  Hirth 
(1960),  Wooley  and  Beaty  (1967),  Genoways  (1973),  Helm  and  Bowers  (1973), 
and  Linzey  and  Layne  (1974).  Hughes  (1964,  1965)  compared  the  morphology 
of  sperm  of  18  species  of  marsupials  representing  five  families,  and  Biggers  and 
Delamater  (1965)  and  Biggers  (1966)  reported  on  the  spermatozoa  of  several 
genera  of  American  marsupials.  Griffiths  (1968)  presented  data  on  the  sperm 
of  the  echidna  and  Bedford  (1967)  reported  observations  on  the  fine  structure 
of  the  spermatozoa  of  two  primates  in  addition  to  man.  An  especially  important 
contribution  is  that  of  Martin  et  al.  (1975).  They  used  scanning  electron 
microscopy  to  compare  spermatozoa  of  16  species  of  primates  representing 
four  families  and  concluded  that  sperm  morphology  might  be  valuable  in  gaining 
better  understanding  of  intrageneric  relationships  among  primates. 

Six  studies  have  described  the  sperm  of  Rhinolophus  ferrumequinum,  and 
Hirth  (1960),  Fawcett  and  Ito  (1965),  Wimsatt  et  al.  (1966),  and  Forman 
(1968)  reported  on  various  aspects  of  the  spermatozoa  of  species  of  vespertilionid 
bats.  Forman  (1968)  was  the  first  to  present  information  on  the  sperm  of  members 
of  the  family  Phyllostomatidae.  In  his  study,  he  presented  information  on  eight 
species  representing  four  of  the  six  subfamilies.  In  the  same  year,  Forman  et  al. 
(1968)  reported  on  two  additional  phyllostomatid  species,  Desmodus  rotundus 
and  Diphylla  ecaudata,  of  a  fifth  subfamily,  the  Desmodontinae. 

Over  the  past  seven  years,  we  have  accumulated  data  on  the  sperm  of 
phyllostomatid  bats  in  the  course  of  several  other  studies  of  this  family.  This  has 
resulted  in  material  for  35  species,  28  of  which  have  not  been  studied  previously. 
Through  new  staining  techniques,  we  also  have  been  able  to  acquire  new 
information  on  the  seven  species  for  which  some  data  were  presented  previously. 
The  results  of  our  studies  and  their  systematic  implications  are  discussed  below. 


177 


178 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Methods  and  Materials 

The  spermatozoa  of  35  species  belonging  to  six  subfamilies  of  phyllostomatids 
were  examined.  To  obtain  spermatozoa,  the  epididymides  of  freshly-killed  bats 
were  removed.  A  small  amount  of  fluid  containing  mature  sperm  was  taken  and 
suspended  in  an  isotonic  solution  of  sodium  citrate.  A  few  drops  of  the  sodium 
citrate  and  spermatozoon  solution  were  placed  on  a  microscope  slide  and  allowed 
to  air-dry.  Dilution  of  the  spermatozoa  with  sodium  citrate  was  necessary  so  that 
individual  spermatozoa  would  be  dispersed  for  study  and  photographing. 
Spermatozoa  on  slides  were  fixed  with  a  solution  of  one  part  acetic  acid  and 
four  parts  absolute  methyl  alcohol.  Slides  were  allowed  to  fix  for  10  to  15  seconds 
and  then  shaken  dry.  Fixing  for  a  longer  period  resulted  in  destruction  of  the 
acrosome. 

Slides  were  stained  with  Toluidine  Blue  O  and  counterstained  with  PAS. 
Counterstaining  resulted  in  delineation  of  the  acrosomal  material  so  that  the 
outline  of  the  headcap  could  be  observed.  The  procedure  outlined  below  was 
followed  in  staining  slides: 

1.  fix  in  solution  of  acetic  acid  and  methyl  alcohol; 

2.  rinse  three  times  in  distilled  water; 

3.  place  in  15%  Periodic  Acid  for  10  minutes; 

4.  rinse  in  tap  water  for  10  minutes; 

5.  rinse  briefly  in  distilled  water; 

6.  place  in  Schiffs'  Reagent  for  10  minutes; 

7.  rinse  in  metabisulfite  with  three  changes  at  three  minutes  each; 

8.  rinse  in  tap  water  for  5  minutes; 

9.  rinse  briefly  in  distilled  water; 

10.  place  in  .02%  Toluidine  Blue  O  for  30  minutes; 

1 1.  place  in  acetone  for  2  minutes; 

12.  place  in  solution  of  acetone  plus  xylene  (1:1)  for  2  minutes; 

13.  place  in  xylene  for  two  changes  at  3  minutes  each; 

14.  mount  using  cover  slip  and  Permount. 

The  following  characters  were  measured:  total  length  of  head,  length  of 
acrosome,  nuclear  length,  head  width,  midpiece  length.  The  mean,  range  (in 
parentheses),  and  one  standard  deviation  for  the  aforementioned  characters  are 
given  beyond  in  the  species  descriptions  whenever  possible.  Measurements  were 
taken  by  means  of  a  Unitron  Filar  widefield  dial  micrometer  attached  to  an  AO 
microstar  Series  10  research  microscope.  Measurements  are  given  in  microns. 

The  terms  dorsal  and  ventral  refer  to  the  flattened  surfaces  of  the  head  and 
midpiece,  whereas  lateral  refers  to  the  narrow  sides  of  the  sperm.  Length  of 
head  included  both  the  acrosome  and  nuclear  area.  Width  of  the  head  was  measured 
as  the  distance  between  extremities  when  observed  in  dorsal  or  lateral  view.  The 
tails  of  sperm  were  not  considered  in  this  study. 

Characters  considered  in  this  study  included:  shape  of  head;  shape  of  apices  of 
acrosome  and  nucleus;  shape  of  base  of  head;  symmetry  of  acrosome  and  head; 
length  of  acrosome  as  compared  with  nucleus;  location  of  posterior  edge  of 
acrosome;  placement  of  the  attachment  of  the  neck  and  midpiece  to  head;  relative 
amount  of  acrosome  anterior  to  nucleus;  thickness,  relative  length,  and  degree 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


179 


Table  1. — Calculated  ratios  comparing  the  dimensions  of  the  spermatozoa  of  35  species  of 

phyllostomatid  hats. 


Midpiece 

length/ 

head 

Species  length 

Head 

length/ 

head 

width 

Head 

length/ 

acrosome 

length 

Midpiece 

length/ 

acrosome 

length 

Nuclear 

length/ 

head 

width 

Midpiece 

length/ 

nuclear 

length 

Head 

length/ 

nuclear 

length 

Nuclear 

length/ 

acrosome 

length 

Micronycteris  megalotis 

1.91 

1.53 

1.78 

3.41 

1.19 

2.45 

1.28 

1.40 

Micronycteris  nicefori 

2.01 

1.18 

1.71 

3.44 

1.11 

2.13 

1.06 

1.62 

Macrotus  waterhousii 

2.00 

1.29 

1.50 

3.00 

0.82 

3.12 

1.56 

0.96 

Tonatia  bidens 

2.44 

1.56 

1.25 

3.05 

1.25 

Mimon  crenulatum 

1.66 

1.38 

1.62 

2.69 

1.03 

2.23 

1.35 

1.20 

Phyllostomus  discolor 

1.73 

1.46 

1.67 

2.89 

1.12 

2.25 

1.30 

1.28 

Glossophaga  soricina 

2.12 

1.19 

1.19 

2.53 

0.90 

2.83 

1.33 

0.90 

Anoura  geoffroyi 

1.44 

1.28 

1.82 

2.62 

0.98 

1.89 

1.31 

1.39 

Choeronycteris  mexicana 

Carollia  brevicauda 

1.51 

1.46 

1.48 

2.24 

0.97 

2.27 

1.50 

0.99 

Carollia  perspicillata 

1.63 

1.56 

1.59 

2.60 

1.07 

2.39 

1.46 

1.09 

Sturnira  lilium 

1.92 

1.65 

1.71 

3.27 

1.17 

2.71 

1.41 

1.21 

Sturnira  tildae 

1.81 

1.59 

1.73 

3.13 

1.27 

2.26 

1.25 

1.38 

Uroderma  bilobatum 

1.86 

1.48 

2.30 

4.29 

1.08 

2.56 

1.37 

1.68 

Vampyrops  helleri 

1.72 

1.62 

1.60 

2.76 

1.24 

2.26 

1.31 

1.22 

Vampyrodes  caraccioli 

1.69 

1.64 

1.76 

2.98 

1.25 

2.21 

1.31 

1.35 

Chiroderma  improvisum 

Chiroderma  trinitatum 

1.82 

1.39 

1.62 

2.95 

1.18 

2.23 

1.23 

1.32 

Mesophylla  macconnelli 

1.65 

1.36 

1.63 

2.69 

1.03 

2.19 

1.32 

1.23 

Artibeus  cinereus 

1.94 

1.29 

1.42 

2.75 

1.07 

2.35 

1.21 

1.17 

Artibeus  toltecus 

1.75 

1.57 

1.66 

2.91 

1.23 

2.23 

1.28 

1.30 

Artibues  jamaicensis' 

1.94 

1.36 

1.64 

3.17 

1.09 

2.42 

1.25 

1.31 

Artibeus  lituratus 

1.73 

1.48 

1.45 

2.51 

1.11 

2.30 

1.33 

1.09 

Ardops  nichollsi 

2.09 

1.35 

1.76 

3.67 

1.03 

2.76 

1.32 

1.33 

Phillops  haitiensis 

1.79 

1.45 

1.73 

3.11 

1.11 

2.34 

1.30 

1.33 

Ariteus  flavescens 

1.97 

1.41 

1.63 

3.21 

1.04 

2.73 

1.36 

1.04 

Stenoderma  rufum 

1.86 

1.43 

1.57 

2.91 

1.08 

2.46 

1.32 

1.08 

Centurio  senex 

1.72 

1.22 

1.66 

2.85 

1.01 

2.07 

1.21 

1.37 

Brachyphylla  cavernarum 

Erophylla  bombifrons 

1.44 

1.42 

1.49 

2.15 

1.09 

1.88 

1.30 

1.14 

Erophylla  sezekorni 

1.59 

1.59 

1.70 

2.80 

1.15 

2.19 

1.38 

1.23 

Phyllonycteris  poeyi 

1.34 

1.40 

1.55 

2.09 

1.03 

1.82 

1.35 

1.15 

Desmodus  rotundus 

2.47 

1.74 

1.58 

3.91 

1.42 

3.03 

1.23 

1.29 

Diaemus  youngii 

2.23 

1.80 

1.75 

3.91 

1.45 

2.78 

1.23 

1.41 

Diphylla  ecaudata 

2.10 

1.32 

1.58 

3.32 

2.09 

1.00 

1.58 

of  tapering  of  midpiece.  Table  1  gives  statistical  ratios  based  on  measurements 
taken.  Figs.  1-5  compare  the  total  head  length,  nuclear  length,  and  midpiece 
length  of  the  species  studied.  Voucher  specimens  are  deposited  in  The  Museum 
of  Texas  Tech  University  (TTU)  and  Carnegie  Museum  of  Natural  History  (CM). 
Most  specimens  were  collected  under  a  grant  from  the  National  Science  Foundation 
(GB-41 105)  to  Robert  J.  Baker  and  Hugh  H.  Genoways. 

Accounts  of  Species 
Subfamily  Phyllostomatinae 
Micronycteris  megalotis  (Gray,  1 842) 

Description  (Fig.  1A). — Head  oval,  rear  portion  tapered  slightly  but  con¬ 
siderably  more  than  that  of  Macrotus\  bilaterally  symmetrical;  apex  narrowly 
rounded;  acrosome  no  wider  than  nucleus;  base  slightly  convex;  nuclear  portion 


180 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


PH  YLLOSTOMATINAE 


Fig.  1. — Sperm  of  six  phyllostomatine  bats.  A)  Micronycteris  megalot  is",  B)  Micronycteris 
nicefori;  C)  Macrotus  waterhousii',  D)  Tonatia  b ideas',  E)  Mimon  crenulatum',  F)  Phyllostomus 
discolor.  Scale  equals  5  microns. 

has  blunt  apex,  more  rounded  than  that  of  acrosome;  acrosome  longer  than  nucleus 
and  constituting  a  substantial  portion  of  the  head  length;  head  length  4.46(4.19- 
4.65)  ±0.138,  4.87(4.56-5.12)  ±0.237,  acrosome  length  3.00(2.79-3.07) 

±0.102,  2.73(2.42-2.98)  ±0.188,  nuclear  length  3.65(3.44-3.91)  ±0.160, 
3.81(3.17-3.19)  ±0.072,  head  width  2.92(2.79-3.07)  ±0.088,  3.19(3.07-3.35) 
±0.091.  Neck  short,  joins  head  midway  at  base  of  head.  Midpiece  extremely 
thin,  relatively  long,  length  9.45(9. 1 1  -9.95)  ± 0.286,  9.32(8.84-9.58)  ± 0.25 1 . 

Remarks. — Morphology  of  the  sperm  head  of  Micronycteris  megalotis  is 
substantially  different  from  that  of  Macrotus  waterhousii,  with  the  sperm  head 
of  M.  megalotis  considerably  narrower  than  that  of  M.  waterhousii. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


181 


Specimens  examined. — Trinidad:  Blanchisseuse,  St.  George,  1  (TTU  23754);  Maracas, 
St.  George,  1  (TTU  23759). 

Micronycteris  nicefori  Sanborn,  1949 

Description  (Fig.  IB). — Head  wider  than  that  of  M.  megalotis,  more  rounded; 
bilaterally  symmetrical;  base  flattened,  not  convex;  acrosome  substantially 
shorter  than  nucleus,  in  sharp  contrast  to  condition  found  in  M.  megalotis;  nuclear 
portion  extremely  rounded;  apex  of  acrosome  and  nucleus  similar  in  shape;  head 
length,  4.00(3.72-4.37)  ±0.299,  acrosome  length  2.34(2.23-2.60)  ±0.145, 
nuclear  length  3.78(3.62-3.91)  ±0.092,  head  width  3.40(2.98-3.72)  ±0.177. 
Neck  short,  not  joining  head  midway  along  base.  Midpiece  extremely  narrow, 
difficult  to  distinguish  from  tail;  length  8.04(7.91-8.18)  ±0.1 15. 

Remarks. — Morphology  of  the  spermatozoa  of  M.  nicefori  is  similar  to  that 
of  M.  megalotis  but  does  differ  in  several  ways.  Most  noticeably,  the  acrosome 
is  shorter  than  the  nucleus  in  M.  nicefori  but  longer  than  the  nucleus  in  M. 
megalotis.  M.  nicefori  also  has  a  wider  sperm  head  than  megalotis  and  a  flattened 
rather  than  convex  base  of  head. 

Specimen  examined. — Trinidad:  2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  23768). 

Macrotus  waterhousii  Gray,  1843 

Description  (Fig.  1C). — Head  not  rounded,  triangular;  bilaterally  symmetrical; 
base  strongly  convex;  apex  of  acrosome  broadly  rounded,  bullet-shaped;  posterior 
border  of  acrosome  sharply  defined;  acrosome  no  wider  than  nucleus  and  similar 
in  length;  nuclear  portion  small,  with  extremely  blunt  apex,  and  more  rounded  than 
apex  of  acrosome;  head  length  3.73(3.53-4.00)  ±0.150,  3.67(3.44-3.81)  ±0.100, 
nuclear  length  2.39(2.32-2.70)  ±0.132,  2.49(2.32-2.79)  ±0.156,  acrosome 
length  2.49(2.32-2.70)  ±0.178,  2.23(2. 14-2.32)  ±0.068,  head  width  2.90(2.70- 
3.16)  ±0.156,  2.95(2.79-3.07)  ±0.1 12.  Neck  short,  joining  head  midway  at 
base  of  head.  Midpiece  extremely  short;  demarcation  with  tail  distinctive;  length 
7.46(7.34-7.63)  ±0.1 12,  7.66(7.16-7.91)  ±0.183. 

Remarks. — The  form  of  the  sperm  head  in  this  species  is  unique  with  no 
comparable  conformation  found  in  any  other  genus.  Also  of  interest  is  the  ex¬ 
tremely  short  midpiece. 

Specimens  examined. — Jamaica:  Green  Grotto,  2  mi.  E  Discovery  Bay,  St.  Ann  Parish, 
3  (TTU  21501-02,  21504). 

Tonatia  bidens  (Spix,  1 823) 

Description  (Fig.  ID). — Head  rounded  to  broadly  oval;  acrosome  can  con¬ 
tribute  markedly  to  total  length  of  head;  acrosome  bilaterally  symmetrical,  rear 
terminus  only  slightly  beyond  apex  of  nucleus;  apex  of  acrosome  broadly  rounded 
but  less  so  than  nucleus;  acrosome  considerably  shorter  than  nucleus  and  never 
wider  than  nucleus;  nucleus  rounded,  with  extremely  blunt  apex;  base  of  head 
concave;  head  length  approximately  4.64(4.46-4.84),  nuclear  length  3.72(3.58- 
4.00),  head  width  2.98(2.88-3.07).  Neck  relatively  long  and  slightly  off  center  of 


182 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


point  of  attachment  to  head.  Midpiece  relatively  long,  anterior  portion  broad, 
tapering  sharply  posteriorly;  length  1 1.36(9.49-1  1.25). 

Remarks. — Among  the  phyllostomatines,  the  head  of  the  spermatozoon 
of  T.  bidens  is  most  similar  in  general  shape  (acrosome  and  nucleus)  to 
Micronycteris  nicefori  and  Phyllostomus  discolor. 

Specimen  examined. — Trinidad:  2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  23794). 

Minion  crenulatum  (E.  Geoffroy  St.-Hilaire,  1810) 

Description  (Fig.  IE). — Head  bluntly  rounded;  acrosome  keel-shaped,  ex¬ 
tremely  asymmetrical;  acrosome  slightly  broader,  at  widest  point,  than  nucleus; 
acrosome  terminates  posteriorly  about  midway  along  length  of  nucleus,  adding 
about  25  per  cent  to  length  of  head;  nucleus  slightly  longer  than  acrosome;  nuclear 
portion  extremely  rounded,  apex  narrowly  rounded  terminating  in  broad  point; 
base  of  nucleus  rounded  but  slightly  concave;  head  length  5.42(5. 1 2-5.86)  ±  0. 1 94, 
acrosome  length  3.34(3. 16-3.53)  ±0.1 34;  nuclear  length  4.02(3.91-4.09)  ±0.068, 
head  width  3.92(3.72-4.09)  ±0.1 19.  Neck  short  with  attachment  to  head  slightly 
off  center.  Midpiece  of  moderate  breadth  anteriorly;  moderate  length;  length 
8.98(8.56-9.39)  ±0.213. 

Remarks. — The  sperm  head  of  Mimon  differs  in  general  morphology  from  both 
Macrotus  and  Micronycteris  and  is  exceptionally  large.  The  asymmetry  of  the 
acrosome  is  in  striking  contrast  to  the  generally  symmetrical  acrosome  of  other 
phyllostomatines. 

Specimen  examined. — Trinidad:  2  mi.  E  San  Rafael,  St.  George,  1  (TTU  23770). 
Phyllostomus  discolor  (Wagner,  1 843) 

Description  (Fig.  IF). — Head  narrowly  rounded;  acrosome  only  slightly 
asymmetrical,  shorter  than  nucleus,  and  terminating  posteriorly  about  half-way 
along  length  of  nucleus;  acrosome  slightly  wider,  at  widest  point,  than  is  nucleus; 
nucleus  triangular  in  shape  with  broad  base,  apex  narrowly  rounded,  pointed;  base 
of  nucleus  slightly  concave;  head  length  5.19(4.93-5.58)  ±0.239,  acrosome  length 
3.1 1(2.79-3.44)  ±0.240,  nuclear  length  3.99(3.53-4.37)  ±0.230,  head  width 
3.55(3.26-3.72)  ±0.159.  Neck  extremely  short,  junction  with  head  considerably 
off  center;  joins  head  on  same  side  as  most  distinct  portions  of  the  apex  of  the 
acrosome.  Midpiece  of  moderate  length,  thin,  tapering  gradually  to  distinctive 
junction  with  tail;  length  8.98(8.56-9.58)  ±0.316. 

Remarks. — The  head  of  the  spermatozoon  of  Phyllostomus  discolor  has 
morphological  similarities  with  both  Mimon  and  Micronycteris  but  is  identical 
to  neither;  the  head  is  most  similar  to  that  of  M.  nicefori  except  that  the  acrosome 
is  slightly  asymmetrical.  The  nucleus  is  narrower  than  in  Mimon  with  broad, 
triangular  base  as  in  M.  nicefori. 

Previous  study. — Two  specimens  from  Nicaragua  (Forman,  1968:905). 

Specimen  examined. — Trinidad:  Las  Cuevas,  St.  George,  1  (TTU  23777). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


183 


Subfamily  Glossophaginae 
Glossophaga  soricina  (Pallas,  1766) 

Description  (Fig.  2A). — Head  extremely  small,  short,  and  quite  rounded; 
base  of  head  broad  giving  a  shovellike  shape;  base  has  well-developed  concavity; 
apex  of  acrosome  nearly  symmetrical,  being  somewhat  more  narrowly  rounded 
than  the  broadly  rounded  apex  of  nucleus;  acrosome  nearly  as  long  as  nucleus; 
posterior  limit  of  acrosome  considerably  behind  midpoint  of  nucleus;  only  a  small 
portion  of  acrosome  occurs  anterior  to  nucleus;  acrosome  never  wider  than  nucleus; 
head  length  3.80(3.53-4.00)  ±0.162,  acrosome  length  3. 19(3.09-3.26)  ±0.202, 
nuclear  length  2.86(2.70-3.26)  ±0.268,  head  width  3.19(3.07-3.26)  ±0.091. 
Neck  moderate  in  length,  junction  with  head  only  slightly  off  center.  Midpiece 
extremely  broad,  tapering  gradually  posteriorly;  junction  with  tail  quite  distinctive; 
length  8.08(7.63-8.46)  ±0.316. 

Remarks. — Sperm  morphology  in  this  species  is  notably  similar  to  that  of 
Anoura;  heads  are  extremely  small  compared  to  those  of  most  other  species. 

Previous  study. — Four  specimens  from  Chiapas  (Forman,  1968). 

Specimens  examined. — Veracruz:  4  km.  W,  5  km.  S  Sontecomapa,  1  (TTU  28900); 
Yucatan:  Merida,  1. 

Anoura  cultrata  Handley,  1960 

Description  (after  Forman,  1968). — Head  rounded,  its  breadth  approximately 
seven-eighths  of  length,  broadest  in  basal  region,  bluntly  rounded  at  apex;  base 
slightly  concave  (the  acrosome  was  not  examined  in  the  previous  study).  Neck  not 
observed.  Midpiece  short  when  compared  to  length  of  tail;  width  uniform  through¬ 
out. 

Remarks. — The  spermatozoa  of  Anoura  cultrata  are  distinct  from  those  of 
Glossophaga  soricina.  The  head  is  broader  in  A.  cultrata  than  in  G.  soricina,  the 
ratio  of  length  to  breadth  being  1.15  as  opposed  to  1.28  in  G.  soricina  (Forman, 
1968). 

Previous  study. — Two  specimens  from  Panama  (Forman,  1968). 

Anoura  geoffroyi  Gray,  1838 

Description  (Fig.  2B,  2C). — Head  quite  rounded;  base  slightly  convex; 
acrosome  slightly  asymmetrical,  with  apex  occasionally  somewhat  pointed; 
acrosome  shorter  than  nucleus  and  contributing  markedly  to  total  head  length; 
acrosome  only  slightly  broader  than  nucleus  at  widest  point;  apices  of  acrosome 
and  nucleus  usually  broadly  rounded,  that  of  the  nucleus  particularly  so;  head 
length  3.92(3.53-4.09)  ±0.184,  4.05(3.91-4.37)  ±0.151,  acrosome  length  2.23 
(2.05-2.32)  ±0.09,  2.23(2.05-2.42)  ±0.1 16,  nuclear  length  3.08(2.79-3.44)  ± 
0.216,  3.09(2.88-3.35)  ±0.165,  head  width  3.14(2.88-3.26)  ±0.128,  3.16(2.98- 
3.35)  ±0.104.  Neck  of  moderate  length,  junction  with  head  slightly  off  center; 
attachment  to  head  on  same  side  as  longest  portion  of  acrosome.  Midpiece  ex- 


184 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


GLOSSOPHAGINAE 


CAROLLI  IN  AE 


1 


E 

Fig.  2. — Sperm  of  some  glossophagine 
B-C)  Anoura  geoffroyr,  D)  Choeronvcteris 
perspicilhita.  Scale  equals  5  microns. 


F 


and  carol liine  bats.  A)  Glossophaga  soricina, ; 
mexicana;  E)  Carollia  brevicauda ;  F)  Carollici 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


185 


tremely  wide  at  anterior  end,  tapering  abruptly  towards  posterior  end;  junction 
with  tail  distinctive;  length  5.57(4.93-6.14)  ±0.358,  5.84(5.58-6.05)  ±0.149. 

Remarks. — The  spermatazoon  of  this  species  is  quite  similar  to  that  of 
Glossophaga  soricina ,  the  only  species  of  the  genus  examined. 

Specimens  examined. — Hidalgo:  13  km.  WSW  Tehuetlan,  2  (TTU  15477-78).  Trinidad: 
2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  23802);  Las  Cuevas,  St.  George,  1  (TTU 
23798). 

Choeronycteris  mexicana  Tschudi,  1844 

Description  (Fig.  2D). — Head  oval,  somewhat  triangular  or  shovel  shaped; 
extremely  large  (in  length  and  breadth);  acrosome  symmetrical,  relatively  long, 
posterior  terminus  well  posterior  to  midpoint  of  head,  and  apex  broadly  rounded; 
acrosome  difficult  to  distinguish  from  nucleus,  blending  in  at  the  sides  of  the  head; 
acrosome  adds  only  slightly  to  total  length  of  head;  nucleus  extremely  rounded, 
apex  rounded;  base  concave,  corners  rounded;  head  length  5.09(4.74-5.58)  ± 
0.259,  acrosome  length  3.37(3.26-3.44)  ±0.089,  nuclear  length  4.26(4.00-4.46) 
±0.158,  head  width  3.99(3.62-4. 19)  ±0. 145.  Neck  short,  attached  to  base  of 
head  nearly  at  its  midpoint.  Midpiece  narrow,  moderate  length,  tapering  only 
slightly  posteriorly;  length  8.59(8.37-9.02)  ±0.182. 

Remarks. — Spermatozoa  from  Choeronycteris  mexicana  are  easily  dis¬ 
tinguishable  by  their  larger  size  from  those  of  other  glossophagines.  Glossoph- 
agines  examined  to  date  appear  relatively  consistent  and  uniform  in  sperm  mor¬ 
phology. 

Specimen  examined. — Tlaxcala:  5  km.  E,  3  km.  N  Tlaxcala,  1  (TTU  25347). 

Subfamily  Carolliinae 
Carollia  castanea  H.  Allen,  1890 

Description  (after  Forman,  1968:909). — Head  rounded,  somewhat  heart- 
shaped;  apex  broadly  rounded;  base  concave  and  symmetrical,  narrowing 
laterally  at  point  of  junction  with  neck  (acrosome  not  observed  in  this  study). 
Neck  short  but  distinct;  junction  with  head  near  center  of  base.  Midpiece  short, 
anterior  end  at  distinct  angle  to  base  of  head,  tapering  only  slightly  posteriorly. 

Remarks. — A  spiraled  midpiece  was  observed  in  this  species,  confirming  the 
existence  of  such  a  structure  in  at  least  one  member  of  the  Phyllostomatidae 
(Forman,  1968). 

Previous  study. — Three  specimens  from  Panama  (Forman,  1968). 

Carollia  brevicauda  (Schinz,  1821) 

Description  (Fig.  2E). — Head  rounded;  acrosome  long,  posterior  border  located 
from  midway  to  two-thirds  back  along  the  length  of  the  nucleus;  acrosome  slightly 
asymmetrical  and  terminating  in  broadly  rounded  apex;  acrosome  extremely  large 
and  longer  than  nucleus,  possibly  somewhat  wider  than  nucleus  at  its  widest 
point;  nucleus  rounded  with  broadly  rounded  apex;  base  of  head  slightly  concave; 


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head  length  5.22(4.84-5.49)  ±0.180,  acrosome  length  3.53(3.26-3.81)  ±0.167, 
nuclear  length  3.48(3.26-3.72)  ±0. 1 1 8,  head  width  3.58(3.44-3.81)  ±0.135. 
Neck  of  moderate  length;  attachment  to  head  off  center,  with  attachment  on  same 
side  as  longest  portion  of  acrosome.  Midpiece  narrow,  moderate  length,  tapering 
gradually  to  posterior;  junction  with  tail  distinctive;  length  7.90(7.53-8.28)  ± 
0.208. 

Remarks. — Overall  shape  of  the  sperm  head  in  C.  brevicauda  is  more  rounded, 
wider,  and  generally  greater  in  size  than  that  of  C.  perspicillata.  C.  brevicauda 
shares  several  characteristics  with  C.  perspicillata ,  including  an  acrosome  that  is 
often  longer  than  the  nucleus  and  a  nucleus  that  is  rounded  with  a  broadly  rounded 
apex. 

Specimen  examined. — Veracruz:  4  km.  W,  5  km.  S  Sontecomapa,  1  (TTU  28901). 
Carollia  perspicillata  (Linnaeus,  1758) 

Description  (Fig.  2F). — Head  relatively  narrow  (because  significant  amount 
of  acrosome  is  anterior  to  apex  of  nucleus;  portion  of  acrosome  anterior  to  apex 
of  nucleus  may  exceed  30  per  cent  of  total  length  of  head);  acrosome  slightly 
asymmetrical,  as  long  as  or  slightly  longer  than  the  nucleus  in  many  cases; 
acrosome  terminates  posteriorly  about  40  to  50  per  cent  of  way  back  along  the 
length  of  the  nucleus;  acrosome  only  slightly  wider  than  nucleus  at  its  widest  point; 
nucleus  rounded,  base  concave,  and  apex  broadly  rounded;  head  length  5.23 
(5.02-5.39)  ±0.103,  acrosome  length,  3.29(3.07-3.53)  ±0. 148,  nuclear  length 
3.58(3.26-3.81)  ±0.201,  head  width  3.35(3.16-3.53)  ±0.131.  Neck  short, 
attached  to  base  of  head  slightly  off  center.  Midpiece  of  moderate  length,  gradually 
tapering;  junction  with  tail  distinctive;  length  8.55(8.1 8-9. 1 1 )  ± 0.28 1 . 

Remarks. — Morphology  of  the  spermatozoon  of  Carollia  perspicillata  resembles 
that  of  Micronycteris  megalotis,  but  the  head  differs  in  several  respects  from  that 
of  C.  brevicauda.  Large  sperm  heads  might  be  characteristic  of  the  genus 
Carollia. 

Specimens  examined. — Quintana  Roo:  14  km.  NE  Playa  del  Carmen,  1  (TTU  18421); 
Trinidad:  Blanchisseuse,  St.  George,  1  (TTU  23859). 

Subfamily  Stenoderminae 
Sturnira  lilium  (E.  Geoffroy  St. -Hilaire,  1810) 

Description  (Fig.  3A). — Head  large,  relatively  narrow  oval;  acrosome 
symmetrical,  shorter  than  nucleus;  acrosome  large,  terminating  anteriorly  in 
moderately  rounded  apex  and  posteriorly  about  halfway  along  length  of  nucleus; 
distinctive  portion  of  acrosome  lies  anterior  to  nucleus;  acrosome  may  be  narrower 
at  base  than  nucleus  at  its  widest  point  or  they  may  be  of  equal  breadth;  nucleus 
oval,  apex  more  broadly  rounded  than  that  of  the  acrosome;  base  extremely  narrow 
(relative  to  greatest  breadth  of  nucleus)  and  concave;  head  length  5.15(4.93- 
5.49)  ±0.179,  acrosome  length  3.02(2.70-3.16)  ±0.150,  nuclear  length  3.64 
(3.44-4.00)  ±0.158;  head  width  3. 12(2.98-3.26)  ±0.085.  Neck  moderate  in 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


187 


length,  attached  to  head  slightly  off  center.  Midpiece  long,  stains  dark;  broad  at 
anterior  end,  sharply  tapering  posteriorly;  junction  with  tail  distinctive;  length 
9.87(9.39-10.14)  ±0.224. 

Remarks. — The  overall  similarity  of  sperm  from  Sturnira  lilium  to  that  found 
in  other  stenodermines  supports  the  inclusion  of  this  genus  within  the  subfamily. 

Previous  study. — Two  specimens  from  Chiapas  (Forman,  1968). 

Specimens  examined. — Trinidad:  2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  23901); 
Blanchisseuse,  St.  George,  1  (TTU  23899). 

Sturnira  tildae  de  la  Torre,  1959 

Description  (Fig.  3B). — Head  similar  in  structure  to  S.  lilium  but  differs  from 
it  in  several  ways;  base  of  head  less  concave  than  that  of  S.  lilium  and  sometimes 
lacking  concavity;  apex  of  acrosome  symmetrical,  as  much  as  half  of  the  acrosome 
occurring  anterior  to  nucleus;  acrosome  covers  only  a  very  small  portion  of  the 
nucleus;  nucleus  ovoid;  head  length  4.81(4.56-5.02)  ±0.121,  4.82(4.65-4.93) 
±0.151,  acrosome  length  2.78(2.51-3.07)  ±0.149,  2.43(2.23-2.70)  ±0.177, 
nuclear  length  3.85(3.62-4.37)  ±0. 1 86,  3.78(3.44-4.09)  ±0.237,  head  width 
3.02(2.88-3.26)  ±0.136,  3.00(2.79-3.35)  ±0.162.  Neck  relatively  long,  attached 
to  middle  of  base.  Midpiece  slightly  shorter  than  that  of  S.  lilium,  extremely 
narrow,  and  tapering  slightly  posteriorly;  length  8.71(8.28-9.1 1)  ±0.250, 
8.81(8.37-9.02)  ±0.293. 

Remarks. — Spermatazoa  of  Sturnira  tildae  differ  from  those  of  species  in  this 
genus  mainly  in  that  base  of  head  is  less  concave  and  midpiece  shorter.  The  small 
acrosome  may  be  unique  to  S.  tildae,  but  that  possibility  awaits  examination  of 
the  acrosome  of  Sturnira  ludovici.  The  nucleus  is  similar  in  configuration  to  that 
of  Artibeus  cine  reus. 

Specimens  examined. — Trinidad:  2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  23907); 
Blanchisseuse,  St.  George,  1  (TTU  23904). 

Sturnira  ludovici  Anthony,  1 924 

Description  (after  Forman,  1968). — Head  much  as  in  S.  lilium,  differing  only 
in  proportions;  apex  blunt;  no  concavity  in  base  (acrosome  not  examined).  Neck 
not  discernible.  Midpiece  broad,  nonhelical,  and  long. 

Remarks. — The  gross  morphology  of  spermatozoa  of  Sturnira  ludovici  is 
similar  to  that  of  S.  lilium.  However,  according  to  measurements  given  by  Forman 
(1968),  length  of  sperm  head  and  length  of  midpiece  are  greater  in  S.  ludovici. 

Previous  study. — Eight  specimens  from  Panama  (Forman,  1968). 

Uroderma  bilobatum  Peters,  1866 

Description  (Fig.  3C). — Head  similar  in  overall  morphology  to  that  of 
Artibeus  jamaicensis;  relatively  narrow;  acrosome  symmetrical  or  slightly 
asymmetrical,  narrowly  rounded  at  apex;  acrosome  notable  in  being  extremely 
short  terminating  posteriorly  one-third  or  less  the  way  along  the  length  of  the 


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STENODERMIN  AE 


Fig.  3. — Sperm  of  some  stenodermine  bats.  A)  Sturnira  l ilium;  B)  Sturnira  tildae ;  C) 
Vroderma  bilobatum;  D)  Vcimpyrops  helleri;  E)  Vampyrodes  caraccioli ;  F)  Chiroderma 
improvisum ;  G)  Chiroderma  trinitatum ;  H)  Mesophylla  macconnelli.  Scale  equals  5  microns. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


189 


nucleus;  approximately  half  of  acrosome  visible  anterior  to  nuclear  apex;  acrosome 
also  appearing  to  be  narrower  in  width  than  the  nucleus;  nucleus  ovoid;  base  of 
nucleus  flattened  or  slightly  concave  with  pointed  corners;  head  length  4.56 
(4.09-4.84)  ±0.23,  acrosome  length  1.98(1.67-2.32)  ±0.27,  nuclear  length 
3.32(3.16-3.53)  ±0. 12,  head  length  3.08(2.98-3.16)  ±0.08.  Neck  extremely 
short,  junction  with  head  well  off  center.  Midpiece  of  moderate  length;  thin  but 
tapering  slightly  posteriorly;  length  8.49(7.91-8.84)  ±0.29. 

Remarks. — Morphology  of  the  sperm  head  of  Uroderma  resembles  most 
closely  that  of  Artibeus,  particularly  A.  jamaicensis.  The  acrosome  of  this  species 
is  unusually  short  and  covers  an  extremely  small  portion  of  the  nucleus.  The 
flattened  base  of  the  head  is  an  unusual  feature. 

Specimen  examined. — Trinidad:  Guayaguayare,  Mayaro,  1  (TTU  24046). 

Vampyrops  helleri  Peters,  1867 

Description  (Fig.  3D). — Head  long  and  narrow,  nucleus  relatively  long  com¬ 
pared  with  other  species;  acrosome  narrow  and  asymmetrical  (terminus  of  apex 
on  same  side  of  head  as  the  attachment  of  the  midpiece  to  the  head),  appears  ter  be 
slightly  narrower  than  nucleus;  apex  of  acrosome  narrowly  rounded  and  may  be 
somewhat  pointed;  posterior  limit  of  acrosome  terminates  midway  along  the  length 
of  the  nucleus;  a  substantial  portion  of  acrosome  occurs  anterior  to  the  apex  of 
the  nucleus;  nucleus  strongly  ovoid  with  rounded  base  that  is  strongly  concave; 
apex  of  nucleus  rounded;  head  length  5.54(5.39-5.77)  ±0.14,  acrosome  length 
3.46(3.16-3.72)  ±0.26,  nuclear  length  4.22(4.09-4.37)  ±0.1 1,  head  width 
3.41(3.26-3.53)  ±0.14.  Neck  short,  junction  with  head  only  slightly  off  center. 
Midpiece  long,  extremely  thin;  junction  with  tail  distinctive;  length  9.54(8.74- 
10.14)  ±0.41. 

Remarks. — Structure  and  size  of  the  sperm  head  within  this  species  is  unique 
among  those  studied  because  it  is  unusually  long;  it  closely  resembles  that  of 
Artibeus  jamaicensis. 

Specimen  examined. — Trinidad:  Guayaguayare,  Mayaro,  1  (TTU  24063). 

Vampyrodes  caraccioli  (Thomas,  1 889) 

Description  (Fig.  3E). — Head  most  complete  oval  of  any  phyllostomatid 
studied  with  base  of  head  extremely  narrow;  head  egg-shaped,  long,  relatively 
narrow,  similar  in  size  but  slightly  smaller  than  that  of  Vampyrops-,  nucleus  and 
acrosome  usually  with  a  symmetrical  apex  at  anterior  end,  apices  narrowly 
rounded  or  pointed,  acrosomal  apex  especially  pointed;  acrosome  usually 
symmetrical  and  equal  in  width  to  nucleus,  in  some  cases  nucleus  appears  to  be 
only  slightly  longer  than  accompanying  acrosome;  posterior  limit  of  acrosome 
sometimes  behind  midpoint  of  nucleus;  substantial  portion  of  acrosome  occurs 
anterior  to  apex  of  nucleus;  base  of  head  extremely  narrow  and  flattened  to 
concave,  with  pointed  corners;  head  length  5.25(4.84-5.49)  ±0.202,  acrosome 
length  2.98(2.79-3.16)  ±0.13,  nuclear  length  4.02(3.72-4.28)  ±0.16,  head  width 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


3.21(3.07-3.44)  ±0. 1 3.  Neck  extremely  short,  attachment  to  base  of  head  only 
slightly  off  center  or  is  centered.  Midpiece  of  moderate  length  and  breadth; 
length  8.89(8.28-9.21)  ±0.39. 

Remarks. — Head  morphology  is  unique  in  being  long  and  having  an  unusually 
narrow  apex  and  base.  Sperm  resembles  somewhat  that  of  Vampyrops,  but  unlike 
Vampyrops,  Vampyrodes  has  a  symmetrical  acrosome  and  an  extremely  narrow 
and  flattened  head  base. 

Specimen  examined. — Trinidad:  Blanchisseuse,  St.  George,  1  (TTU  24060). 


Chiroderma  improvisum  Baker  and  Genoways,  1976 

Description  (Fig.  3F). — Head  similar  to  that  of  C.  trinitatum,  but  slightly  less 
rounded;  acrosome  sometimes  appears  to  be  asymmetrical,  short,  and  with  a  small 
portion  extending  anterior  to  nucleus;  posterior  limit  of  acrosome  lies  in  front  of 
midpoint  of  nucleus  and  appears  less  arched  than  in  C.  trinitatum ;  nucleus  ovoid, 
apex  considerably  more  rounded  than  the  more  pointed  apex  of  the  acrosome; 
base  of  head  asymmetrical,  but  less  so  than  in  C.  trinitatum ;  base  slightly  concave; 
head  length  4.74(4.37-5.30)  ±0.28,  acrosome  length  2.65(2.60-2.79)  ±0.08, 
nuclear  length  3.96(3.81-4.19)  ±0.14,  head  width  3.17(2.88-3.26)  ±0.18. 
Neck  relatively  long,  junction  with  head  well  off  center  as  in  C.  trinitatum.  Mid¬ 
piece  of  moderate  breadth,  tapering  posteriorly;  length  8.64(7.53-9.95)  ±0.71. 

Remarks. — Although  similar  to  that  of  Chiroderma  trinitatum,  the  sperm  head 
in  C.  improvisum  is  slightly  less  rounded,  its  base  less  asymmetrical,  and  it  pos¬ 
sesses  a  shorter  acrosome.  The  spermatozoa  of  species  of  Chiroderma  can  be 
distinguised  easily  from  other  stenodermines. 

Specimen  examined. — Guadeloupe:  2  km.  S,  2  km.  E  Baie-Mahault,  Basse-Terre, 
1  (TTU  19900). 

Chiroderma  trinitatum  Goodwin,  1958 

Description  (Fig.  3G) — Head  morphology  generally  variable;  shape  ovoid 
to  rounded;  nucleus  ovoid  with  pointed  apex;  acrosome  nearly  symmetrical, 
short,  with  apex  only  slightly  more  rounded  than  that  of  nucleus;  terminal  border 
of  acrosome  appears  to  be  slightly  arched  with  apex  directed  anteriorly;  acrosome 
terminates  posteriorly  at  midpoint  of  nucleus  and  extends  anteriorly  only  very 
slightly  beyond  apex  of  nucleus;  base  of  head  flattened  or  very  slightly  concave 
and  is  unusual  in  being  asymmetrical  with  the  greatest  posterior  extension 
occurring  on  the  side  of  the  head  that  is  in  contact  with  the  neck;  base  of  head 
narrower  than  girth  of  head,  with  corners  pointed;  head  length  4.87(4.56-5.39) 
±0.26,  acrosome  length  3.00(2.70-3.35)  ±0.23,  nuclear  length  3.97(3.62- 
4.28)  ±0.25,  head  width  3.37(3.07-3.62)  ±0.175.  Neck  relatively  long,  junction 
with  head  well  off  center  and  nearly  to  the  edge  of  base  of  head.  Midpiece  thin, 
tapering  gradually  posteriorly  and  short  relative  to  length  of  head;  length  8.84 
(8.56-9.1 1)  ±0.21. 

Remarks. — The  morphology  of  the  spermatozoa  head  in  this  species,  although 
variable,  is  distinctly  different  from  that  of  other  stenodermines.  Only  a  very  small 


.BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


191 


portion  of  the  acrosome  extends  anterior  to  nucleus,  the  base  of  the  head  is 
asymmetrical,  and  the  point  of  midpiece  attachment  is  substantially  off  center. 

Specimen  examined. — Trinidad:  2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  24026). 

Mesophylla  macconnelli  Thomas,  1901 

Description  (Fig.  3H). — Head  relatively  long  and  narrow,  not  large;  acrosome 
with  pointed  asymmetrical  apex,  tip  of  apex  on  same  side  of  head  as  attachment 
of  midpiece;  acrosome  short  and  an  extremely  small  portion  of  it  occurs  anterior 
to  the  apex  of  the  nucleus;  posterior  limit  of  acrosome  slightly  anterior  to  mid¬ 
point  of  nucleus;  acrosome  considerably  shorter  than  the  nucleus  (often  only 
slightly  more  than  half  its  length)  and  the  same  breadth  as  the  nucleus  at  its 
posterior  limit;  nucleus  ovoid,  apex  symmetrical;  base  of  head  flattened  with 
slight  concavity;  base  of  head  narrower  than  its  girth,  asymmetrical  with  corner 
nearest  the  midpiece  being  more  pointed  than  the  other;  head  length  4.71(4.56- 
5.02)  ±0.14,  4.68(4.28-4.93)  ±0.19,  acrosome  length  2.73(2.51-2.88)  ±0.12, 
2.64(2.51-2.88)  ±0.13,  nuclear  length  4.01(3.62-4.19)  ±0.15,  3.99(3.81- 

4.37)  ±0.22,  head  width  3.13(2.98-3.34)  ±0.12,  3.25(3.07-3.44)  ±0.10.  Neck 
relatively  long,  junction  with  head  well  off  center  and  near  the  pointed  corner  of 
the  head  base.  Midpiece  short,  broad  anteriorly,  tapering  abruptly  posteriorly; 
junction  with  tail  indistinct;  length  7.61(7.25-7.92)  ±0.23,  7.66(7.25-8.18) 
±0.27. 

Remarks. — Most  notable  among  the  characteristics  of  sperm  from  Mesophylla 
is  the  minute  amount  of  acrosome  anterior  to  the  nuclear  apex  and  the  unusual 
asymmetry  of  the  base  of  the  head.  The  head  is  somewhat  similar  to  that  of 
Phyllostomus  discolor,  but  the  base  and  apex  of  the  nucleus  are  dissimilar. 
An  extremely  short  midpiece  distinguishes  M.  macconnelli  from  other  stenoder- 
mines,  with  the  exception  of  Centurio. 

Specimen  examined. — Trinidad:  Guayaguayare,  Mayaro,  2  (TTU  24039,  24044). 

Artibeus  cinereus  (Gervais,  1855) 

Description  (Fig.  4A). — Head  broad  in  midsection,  tapering  distinctively  both 
anteriorly  and  posteriorly;  acrosome  extremely  pointed,  nearly  cone-shaped, 
slightly  shorter  than  nucleus,  and  terminating  posteriorly  about  midway  along 
nucleus;  nucleus  rounded;  base  of  head  slightly  convex  or  often  lacking  concavity, 
base  of  head  notably  rounded  at  the  corners;  head  length  4.59(4.28-4.84)  ±0.495, 
acrosome  length  2.93(2.51-3.26)  ±0.339,  nuclear  length  3.62(3.35-3.91)  ±0.104, 
head  width  3.15(2.98-3.26)  ±0.084.  Neck  short,  junction  with  head  very  slightly 
off  center.  Midpiece  broad  anteriorly,  tapering  gradually  posteriorly;  length  8.74 
(8.37-9.02)  ±0.342. 

Remarks. — Sperm  morphology  in  this  species  is  very  similar  to  that  of  Artibeus 
jamaicensis,  Ardops  nichollsi,  and  Ariteus  flavescens.  The  most  unusual  feature 
is  the  extremely  pointed,  exceptionally  tapered  apex  to  the  symmetrical  acrosome. 

Specimens  examined. — Trinidad:  Guayaguayare,  Mayaro,  1  (TTU  23924);  2  mi.  E  San 
Rafael,  St.  George,  1  (TTU  23936). 


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STENODER  M I N  AE 


B 


E  F  G  H 

Fig.  4. — Sperm  of  some  stenodermine  bats.  A)  Artibeus  cinereus ;  B)  Artibeus  toltecus, 
C)  Artibeus  jamaicensir,  D)  Artibeus  lituratus ;  E)  Ardops  nichollsi ;  F)  Phyllops  haitiensis , 
G)  Ariteus  flavescens ,  H)  Stenoderma  rufum;  I)  Centurio  senex.  Scale  equals  5  microns. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


193 


Artibeus  toltecus  (Saussure,  1 860) 

Description  (Fig.  4B). — Head  quite  long,  appearing  relatively  narrow,  similar 
to  other  species  of  Artibeus ;  nucleus  ovoid  with  relatively  narrow  apex  and  base; 
acrosome  symmetrical  and  pointed  at  apex;  posterior  limit  of  acrosome  extending 
to  midway  along  length  of  nucleus;  head  length  4.96(4.84-5.12)  ±0.10,  acrosome 
length  2.99(2.79-3.07)  ±0.09,  nuclear  length  3.88(3.62-4.19)  ±0.20,  head 
width  3.16(2.98-3.35)  ±0.12.  Neck  short,  junction  with  head  well  off  center. 
Midpiece  short,  compared  to  length  of  head,  and  narrow;  tapering  posteriorly; 
length  8.69(8.37-9.02)  ±0.21. 

Remarks. — General  shape  of  head  similar  to  Ardops,  Ariteus,  and  other 
species  of  Artibeus,  particularly  A.  lituratus\  however,  the  head  in  general  is  less 
rounded  than  in  other  species.  Heads  of  spermatozoa  from  A.  toltecus  are  longer 
than  in  other  stenodermines. 

Specimen  examined. — Veracruz:  4  km.  W,  5  km.  S  Sontecomapa,  1  (TTU  28902). 

Artibeus  jamaicensis  Leach,  1821 

Description  (Fig.  4C). — Head  similar  in  morphology  to  that  of  Ardops  and 
Ariteus-,  acrosome  usually  symmetrical,  but,  if  asymmetrical,  only  slightly  so; 
apex  of  acrosome  narrowly  rounded  to  nearly  pointed;  portion  of  acrosome 
anterior  to  nucleus  always  less  than  in  Ariteus  and  Ardops-,  nucleus  narrowly 
rounded  at  apex;  base  of  nucleus  broad  and  slightly  concave;  head  length  4.48 
(4.28-4.65)  ±0.1 19,  acrosome  length  2.74(2.51-2.98)  ±0.148,  nuclear  length 
3.59(3.35-4.00)  ±0.159,  head  width  3.30(3. 16-3.44)  ±0.089.  Neck  short, 
junction  with  head  off  center.  Midpiece  nearly  twice  head  length,  thick  anteriorly, 
and  tapering  posteriorly;  length  8.69(8.09-9.21)  ±0.316. 

Remarks. — Morphology  of  the  heads  of  spermatozoa  from  A.  jamaicensis  is 
quite  similar  to  that  of  both  Ariteus  and  Ardops,  but  the  portion  of  the  acrosome 
anterior  to  the  nucleus  was  always  less  in  A.  jamaicensis.  The  acrosome  has  less 
symmetry  than  other  species  of  Artibeus  that  have  been  examined. 

Previous  study. — One  specimen  from  Dominica  and  one  specimen  from 
Nayarit  (Forman,  1968). 

Specimen  examined. — Haiti:  1  km.  E  Lebrun,  Dept,  du  Sud,  1  (TTU  22649). 


Artibeus  lituratus  (Olfers,  1818) 

Description  (Fig.  4D). — Head  similar  to  other  Artibeus-,  acrosome  relatively 
larger  (as  compared  with  nucleus)  than  that  of  other  species  within  the  genus; 
acrosome  only  slightly  shorter  than  the  nucleus,  with  somewhat  narrowly  rounded, 
symmetrical  apex;  acrosome  distinctly  triangular,  its  posterior  limit  consistently 
well  behind  the  midpoint  of  the  nucleus;  acrosome  sometimes  slightly  narrower 
than  nucleus,  otherwise  equivalent  in  width  at  its  posterior  limit;  distinctive  por¬ 
tion  of  acrosome  found  anterior  to  nuclear  apex;  apex  of  nucleus  rounded  but 
rarely  as  narrowly  as  acrosome;  base  of  head  asymmetrical  with  corner  nearest 
neck  slightly  more  posterior  than  the  rounded  corner  on  the  other  side  of  the  base; 


194 


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base  slightly  concave;  head  length  4.77(4.46-5.21)  ±0.229,  acrosome  length 
3.30(3.16-3.53)  ±0.132,  nuclear  length  3.59(3.35-3.72)  ±0.103,  head  width 
3.23(3.07-3.35)  ±0.140.  Neck  relatively  long,  junction  with  head  off  center. 
Midpiece  length  similar  to  other  species  of  Artibeus ;  tapering  gradually  pos¬ 
teriorly;  junction  with  tail  quite  distinctive;  length  8.27(7.91-8.46)  ±0.158. 

Remarks. — Head  morphology  of  sperm  of  A.  lituratus  is  similar  to  that  of 
other  species  of  Artibeus  but  is  most  like  A.  toltecus,  A.  jamaicensis,  and 
Vampyrops  helleri. 

Previous  study. — Two  specimens  from  Chiapas  (Forman,  1968). 

Specimen  examined. — Trinidad:  Guayaguayare,  Mayaro,  1  (TTU  24010). 

Ardops  nichollsi  (Thomas,  1891) 

Description  (Fig.  4E). — Head  bullet  shaped  with  pointed  apex;  acrosome 
asymmetrical  (but  sometimes  nearly  symmetrical);  apex  pointed  or  very  narrowly 
rounded;  a  moderate  portion  of  acrosome  extends  forward  beyond  nucleus;  some 
acrosomes  narrower  than  nucleus;  acrosome  shorter  than  nucleus,  terminating 
posteriorly  at  a  point  slightly  anterior  to  midpoint  of  nucleus;  nucleus  extremely 
rounded  at  apex;  base  broad  and  deeply  concave;  head  length  4.25(4.00-4.65) 
±0.150,  4.31(3.81-4.56)  ±0.262,  acrosome  length  2.42(2.32-2.60)  ±0.132, 
2.58(2.42-2.70)  ±0.1 17,  nuclear  length  3.22(3.07-3.44)  ±0.125,  3.37(3.16- 
3.53)  ±0.1 15,  head  width  3.14(2.88-3.44)  ±0.260,  3.03(2.88-3.16)  ±0.096. 
Neck  short,  junction  with  head  off  center.  Midpiece  of  moderate  length,  thin, 
gradually  tapering  posteriorly;  junction  with  tail  not  distinctive;  length  8.88(8.74- 
9.30)  ±0.192,  8.54(8.09-9.02)  ±0.277. 

Remarks. — The  symmetry  of  the  acrosome  appears  to  be  variable  in  this 
species.  In  some  spermatozoa,  acrosomes  are  asymmetrical,  but  in  others, 
nearly  symmetrical.  Spermatozoa  are  similar  to  those  of  Ariteus  and  Artibeus. 

Specimens  examined. — Guadeloupe:  1  km.  S  Basse-Terre,  Basse-Terre,  1  (TTU  20816); 
1  km.  N,  1  km.  W  St.  Franfois,  Grande-Terre,  1  (TTU  20847). 

Phyllops  haitiensis  (J.  A.  Allen,  1908) 

Description  (Fig.  4F). — Head  usually  somewhat  triangular  in  shape;  acrosome 
only  slightly  asymmetrical;  posterior  terminus  of  acrosome  at  midpoint  of 
nucleus;  substantial  portion  of  acrosome  occurring  anterior  to  the  apex  of  the 
nucleus;  acrosome  shorter  than  nucleus  with  similar  morphology  and  placement 
(orientation)  on  the  nucleus  as  Artibeus,  Ardops,  and  Ariteus-,  nucleus  rounded 
with  broadly  rounded  apex;  base  of  nucleus  with  rounded  corners  and  a  slight 
concavity  or  no  concavity  in  center  of  basal  border;  head  length  4.90(4.28- 
5. 12)  ±0.23,  4.82(4.65-5.12)  ±0.13,  acrosome  length  2.80(2.51-3.07)  ±0.16, 
2.78(2.60-2.98)  ±0.14,  nuclear  length  3.76(3.62-3.91)  ±0.20,  3.70(3.53- 

3.91)  ±0.11,  head  width  3.57(3.35-3.72)  ±0.13,  3.32(3.26-3.44)  ±0.06.  Neck 
extremely  short,  junction  with  head  only  slightly  off  center.  Midpiece  of  moderate 
length  and  breadth,  tapering  only  slightly  posteriorly;  junction  with  tail  distinctive; 
midpiece  length  8.74(8.37-9.30)  ±0.31,  8.64(8.37-9.1 1)  ±0.23. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


195 


Remarks. — The  morphology  of  the  sperm  head  of  Phyllops  is  similar  to  that 
of  Artibeus,  Ariteus,  and  Ardops.  Nuclear  morphology  is  most  like  that  of 
Artibeus  cinereus,  but  the  base  of  the  nucleus  is  less  concave  than  in  most  species 
of  Artibeus. 

Specimens  examined. — Haiti:  2  km.  N,  2  km.  E  Lebrun,  Dept,  du  Sud,  1  (TTU  22672); 
1  km.  S,  1  km.  E  Legrun,  Dept,  du  Sud,  1  (TTU  22697);  4  km.  S  Lebrun,  Dept,  du  Sud, 
1  (TTU  22733). 

Ariteus  flavescens  (Gray,  1831) 

Description  (Fig.  4G). — Head  nearly  identical  in  morphology  to  that  of 
Ardops  nichollsi-,  triangular;  acrosome  extremely  pointed  at  apex  and  acrosome 
can  be  asymmetrical  or  symmetrical;  acrosome  shorter  than  nucleus;  base  of  head 
broad  and  concave;  head  length  4.60(4.37-4.84)  ±0.156,  acrosome  length  2.83 
(2.60-3.16)  ±0.233,  nuclear  length  3.27(2.88-3.53)  ±0.208,  head  width  3.49 
(3.26-3.62)  ±0.136.  Neck  short,  junction  with  head  off  center.  Midpiece  of 
moderate  breadth  anteriorly,  tapering  posteriorly;  junction  with  tail  distinctive; 
length  9.08(8.56-9.30)  ±0.304. 

Remarks. — The  head  of  spermatozoa  from  this  species  bears  a  striking  re¬ 
semblance  to  that  of  Ardops  nichollsi.  The  two  also  are  extremely  similar  in 
dimensions  of  the  nucleus,  acrosome,  and  length  of  midpiece. 

Specimens  examined. — Jamaica:  Queenhythe,  St.  Ann  Parish,  1  (TTU  21774);  Duanvale, 
Trelawny  Parish,  1  (TTU  21781). 

Stenoderma  rufum  Desmarest,  1 820 

Description  (Fig.  4H). — Head  most  similar  in  shape  to  those  of  Ariteus, 
Ardops,  and  Artibeus-,  more  or  less  triangular,  both  nucleus  and  acrosome 
generally  symmetrical;  acrosome  short  and  usually  quite  pointed  at  apex;  acrosome 
usually  narrower  at  base  than  is  nucleus  at  its  widest  point;  acrosome  can  be 
slightly  asymmetrical  at  apex  in  that  sometimes  it  is  offset  to  side  of  head  with 
attachment  to  neck;  one-third  to  half  of  acrosome  occurring  anterior  to  the  apex 
of  nucleus;  posterior  border  of  acrosome  lies  anterior  to  midpoint  of  nucleus; 
nucleus  nearly  triangular  with  broadly  rounded  apex  and  quite  rounded  corners 
at  the  base;  base  slightly  concave  to  nearly  flattened;  head  length  4.58(4.19- 
4.84)  ±0.18,  4.48(4.37-4.65)  ±0.13,  acrosome  length  2.92(2.60-3.26)  ±0.16, 
2.81(2.42-2.98)  ±0.17,  nuclear  length  3.46(3.26-3.81)  ±0.13,  3.56(3.44- 

3.81)  ±0.11,  head  width  3.20(2.88-3.35)  ±0.15,  3.21(3.07-3.35)  ±0.10.  Neck 
relatively  long,  junction  with  base  of  head  moderately  off  center.  Midpiece 
relatively  broad,  tapering  gradually  posteriorly;  junction  of  midpiece  and  tail 
distinctive;  length  8.50(8.18-8.84)  ±0.20,  8.33(8.09-8.65)  ±0.16. 

Remarks. — Head  of  sperm  in  this  species  is  most  similar  to  that  of  Ariteus 
flavescenes,  Ardops  nichollsi,  and  members  of  the  genus  Artibeus  but  is  dis¬ 
tinguishable  from  all  of  them.  The  most  unusual  feature  of  the  spermatozoa  of  this 
species  is  the  narrowness  of  the  acrosome  relative  to  the  breadth  of  the  nucleus. 
Also,  the  nucleus  and  acrosome  are  extremely  similar  in  outline,  a  situation  rarely 
observed. 


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Specimens  examined. — Puerto  Rico:  El  Verde,  2  (TTU  22361,  22362). 


Centurio  senex  Gray,  1 842 

Description  (Fig.  41). — Head  short,  nuclear  portion  extremely  rounded; 
acrosome  symmetrical  with  extremely  pointed  apex,  forming  an  isosceles  triangle, 
as  wide  as  nucleus;  posterior  limit  of  acrosome  lies  in  front  of  center  of  nucleus; 
acrosome  shorter  than  nucleus;  moderate  portion  of  acrosome  occurs  anterior 
to  the  nuclear  apex,  which  is  narrowly  rounded;  nucleus  usually  as  wide  as  it  is 
long  with  its  anterior  border  often  appearing  flattened  on  either  or  both  sides;  base 
of  head  flattened  or  even  slightly  convex,  giving  base  a  rounded  appearance;  head 
length  4.44(4. 19-4.74)  ±0.20,  acrosome  length  2.68(2.42-3.07)  ±0.28,  nuclear 
length  3.68(3.44-4.00)  ±0. 1 8,  head  width  3.65(3.35-3.91)  ±0.17.  Neck  long, 
junction  with  head  well  off  center.  Midpiece  extremely  thin,  short;  length  7.36 
(7.34-7.91)  ±0.20. 

Remarks. — The  morphology  of  the  sperm  head  in  Centurio  senex  is  distinctive 
and  unique.  The  acrosome  is  extremely  pointed,  the  nucleus  nearly  circular. 
Perhaps  the  greatest  contrast  in  degree  of  pointedness  of  nuclear  and  acrosomal 
apices  is  observed  in  this  species. 

Specimen  examined. — Trinidad:  Blanchisseuse,  St.  George,  1  (TTU  24019). 

Subfamily  Phyllonycterinae 
Brachyphylla  cavernarum  Gray,  1 834 

Description  (Fig.  5A). — Head  of  moderate  length,  narrow;  acrosome 
symmetrical,  considerably  shorter  than  nucleus,  and  with  its  posterior  limit  well 
anterior  to  midpoint  of  nucleus;  nucleus  more  ovoid  than  that  of  Ardops,  Ariteus, 
and  Artibeus\  base  slightly  concave;  head  length  4.60,  5.12,  acrosome  length 
2.79,  2.79,  nuclear  length  3.26,  3.53,  head  width  2.79,  1.98.  Neck  short,  junction 
with  head  near  center.  Midpiece  of  moderate  width,  long,  tapering  posteriorly; 
junction  with  tail  distinctive. 

Remarks. — The  sperm  of  Brachyphylla  is  different  from  other  phyllonycterines 
and  does  not  possess  features  generally  found  among  other  members  of  the  sub¬ 
family  (for  example,  Brachyphylla  differs  in  shape  and  size  of  the  acrosome, 
relative  length  of  the  midpiece,  symmetry  of  the  head). 

Specimens  examined. — Guadeloupe:  1  km.  S  Basse-Terre,  Basse-Terre,  1  (TTU  20966); 
1  km.  N,  1  km.  W  St.  Francois,  Grande-Terre,  1  (TTU  20976). 

Erophylla  bombifrons  (Miller,  1899) 

Description  (Fig.  5B). — Head  extremely  long,  ovoid  and  generally  robust; 
acrosome  large  and  encompassing  a  distinctive  portion  of  the  head;  acrosome 
with  slight  asymmetry,  anteriormost  limit  of  apex  on  the  same  side  of  head  as 
attachment  of  tail,  and  with  an  apex  quite  similar  in  shape  to  that  of  the  nucleus; 
acrosome  only  slightly  wider  than  the  nucleus,  terminating  posteriorly  just  beyond 
midpoint  of  nucleus;  acrosome  only  slightly  shorter  than  nucleus;  nucleus  broad 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


197 


PHYLLONYCTERINAE 


DESMODONTINAE 


E  F  G 


Fig.  5. — Sperm  of  some  phyllonycterine  and  desmodontine  bats.  A)  Brachyphylla 
cavernarum;  B)  Erophylla  bombifrons\  C)  Erophylla  sezekornr,  D)  Pbyllonycteris  poeyr, 
E)  Desmodus  rotundas ,  F)  Diaemus  youngii;  G)  Diphylla  ecaudata.  Scale  equals  5  microns. 

and  usually  rounded,  apex  symmetrical;  base  of  nucleus  strongly  asymmetrical 
and  concave,  with  corner  nearest  attachment  of  midpiece  often  less  rounded  than 
other  corner;  head  length  5.14(4.84-5.30)  ±0.148,  acrosome  length  3.45(3.26- 
3.62)  ±0.142,  nuclear  length  3.95(3.62-4.09)  ±0.146,  head  width  3.62(3.53- 


198 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


3.81)  ±0.09.  Neck  appears  extremely  long;  junction  with  base  of  head  off  center. 
Midpiece  broad  anteriorly,  tapering  abruptly;  length  7.42(7.07-8.37)  ±0.379. 

Remarks. — The  head  of  the  sperm  of  Erophylla  bombifrons  is  similar  to  that 
of  Phyllonycteris  poeyi',  however,  the  acrosome  of  E.  bombifrons  is  smaller  and 
not  so  asymmetrical.  The  midpiece  of  this  species  is  exceptionally  thick  at  its 
anterior  end. 

Specimens  examined. — Puerto  Rico:  1  mi.  W  Corozal,  2  (TTU  22426,  22429). 


Erophylla  sezekorni  (Gundlach,  1861) 

Description  (Fig.  5C). — Head  narrow  and  long,  oval  in  general  shape;  acrosome 
exceedingly  asymmetrical  with  apex  on  same  side  of  head  as  midpiece  attachment; 
acrosome  slightly  wider  than  nucleus  at  its  widest  point;  posterior  terminus  of 
acrosome  at  or  slightly  posterior  to  midpoint  of  head;  acrosome  shorter  than 
nucleus  by  small  amount  and  with  apex  more  narrowly  rounded  than  that  of 
nucleus;  nucleus  an  egg-shaped,  rounded  oval  with  broadly  rounded  apex;  base 
of  head  rounded  or  slightly  concave;  amount  of  acrosome  anterior  to  nucleus 
variable  but  generally  amount  is  moderate;  head  length  4.74,  4.84,  acrosome 
length  2.79,  2.98,  nuclear  length  3.44,  3.53,  head  width  2.98,  3.07.  Neck  moderate 
in  length,  junction  with  head  slightly  off  center.  Midpiece  short,  broad  anteriorly 
(but  considerably  less  so  than  in  E.  bombifrons),  and  tapering  gradually  pos¬ 
teriorly;  junction  with  tail  indistinct;  length,  7.53. 

Remarks. — The  head  of  the  sperm  of  Erophylla  sezekorni  is  like  that  of  E. 
bombifrons  but  is  more  similar  to  that  of  Phyllonycteris  poeyi  in  general  char¬ 
acteristics.  The  acrosome  in  Erophylla  is  much  smaller  than  in  Phyllonycteris 
and  with  considerably  less  exposed  acrosome  than  in  sperm  of  Phyllonycteris.  The 
thickened  area  of  the  tail  just  distal  to  the  midpiece  in  P.  poeyi  was  not  observed 
in  either  species  of  Erophylla. 

Specimen  examined. — Jamaica:  Orange  Valley,  St.  Ann  Parish,  1  (TTU  21894). 

Phyllonycteris  poeyi 

Description  (Fig.  5D). — Head  extremely  long  and  broad  because  of  enormous 
asymmetrical  acrosome;  acrosome  slightly  wider  than  long  with  apex  extremely 
broad  and  on  same  side  of  head  as  midpiece  attachment;  apex  of  acrosome  even 
more  removed  from  the  midline  of  nucleus  than  midpiece,  with  result  that  the 
apex  is  often  so  far  off  center  as  to  be  outside  the  axis  of  the  nucleus;  acrosome 
broadest  of  any  phyllostomatid  studied  and  broader  than  nucleus;  acrosome 
terminates  posteriorly  slightly  beyond  the  midpoint  of  nucleus;  nucleus  bilaterally 
symmetrical  except  for  base;  nucleus  a  broad  oval,  being  slightly  longer  than 
acrosome;  base  of  nucleus  concave,  and  of  moderate  breadth,  apex  rounded;  head 
length  6.42(6. 14-6.98)  ±0.214,  6.67(6.32-6.88)  ±0.204,  acrosome  length 

4. 13(3.8 1-4.50)  ±0.2 15,  4.56(4.28-5.02)  ±0.234,  nuclear  length  4.74(4.56- 
5.02)  ±0.156,  4.73(4.46-5.02)  ±0.201,  head  width  4.60(4.19-4.74)  ±0.169, 
4.57(4.19-4.74)  ±0.157.  Neck  short,  junction  with  head  off  center.  Midpiece  of 
moderate  length;  broad  anteriorly  and  tapering  posteriorly;  unusual  tapered 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


199 


thickening  of  tail  just  distal  to  junction  of  tail  and  midpiece;  length  8.63 
(8.18-8.84)  ±0.204,  8.63(8.28-8.93)  ±0.237. 

Remarks. — The  sperm  of  P.  poeyi  exhibits  several  unique  characteristics.  The 
acrosome  has  an  unusual  morphology  including  having  the  apex  far  offset  and 
being  the  broadest  of  any  species  studied.  This  is  the  only  species  examined  in 
which  over  half  of  the  area  of  the  acrosome  occurs  anterior  to  the  apex  of  the 
nucleus.  There  is  an  unusual  thickening  in  the  tail  of  all  specimens  that  occurs  just 
distal  to  the  junction  of  the  tail  and  midpiece;  the  thickened  area  tapers  posteriorly 
into  a  narrow  tail. 

Specimens  examined. — Haiti:  1  km.  E  Lebrun,  Dept,  du  Sud,  1  (TTU  22773);  1  km. 
S  Lebrun,  Dept,  du  Sud,  1  (TTU  22782);  4  km.  S  Lebrun,  Dept,  du  Sud,  1  (TTU  22798). 

Subfamily  Desmodontinae 
Desmodus  rotundus  (E.  Geoffroy  St. -Hilaire,  1810) 

Description  (Fig.  5E). — Head  long,  narrow,  and  extremely  ovoid  with  narrowly 
rounded  apex  and  narrow  base;  acrosome  long,  terminating  posteriorly  well 
behind  midpoint  of  nucleus,  apex  symmetrical;  most  of  acrosome  in  contact 
with  nucleus,  only  an  extremely  minute  portion  anterior  to  nuclear  apex;  viewed 
dorsally,  nucleus  comprises  most  of  head;  acrosome  no  wider  than  nucleus,  apex 
of  acrosome  slightly  more  rounded  than  that  of  nucleus;  base  of  head  quite  narrow, 
with  distinctive  concavity  at  junction  with  neck;  head  length  4.71(4.46-4.93)  ± 
0.183,  acrosome  length  2.98(2.88-3.07)  ±0.067,  nuclear  length  3.84(3.62-4.09) 
±0.162,  head  width  2.71(2.51-2.88)  ±0.103.  Neck  extremely  short;  attaches  at 
center  of  head.  Midpiece  extremely  long,  thickened  or  even  flared  at  neck;  tapers 
gradually  posteriorly;  junction  with  tail  moderately  distinctive;  length  11.64 
(1 1.16-12.18)  ±0.277. 

Remarks. — The  heads  of  the  spermatozoa  of  Desmodus  rotundus  show  much 
greater  symmetry  than  other  phyllostomatid  subfamilies.  The  other  unique 
features  of  the  sperm  of  this  species  include  the  relatively  long  and  narrow  head, 
long  midpiece  that  is  flared  at  the  anterior  end,  and  an  acrosome  closely  attached 
to  the  nucleus. 

Previous  study. — Two  specimens  from  Nicaragua  (Forman  et  al.,  1968). 

Specimens  examined. — Trinidad:  2  mi.  N,  2  mi.  E  Valencia,  St.  Andrew,  1  (TTU  24086); 
Blanchisseuse,  St.  George,  1  (TTU  24080). 

Diaemus  youngii  (Jentink,  1 893) 

Description  (Fig.  5F). — Head  very  similar  in  structure  to  that  of  Desmodus 
rotundus,  however,  acrosome  protrudes  well  anterior  of  apex  of  nucleus;  acrosome 
symmetrical,  relatively  narrow  compared  to  D.  rotundus ,  and  with  posterior 
limit  often  well  in  front  of  the  midpoint  of  the  nucleus;  apex  of  acrosome  somewhat 
more  rounded  than  that  of  the  nucleus;  nucleus  longer  than  acrosome;  nucleus 
nearly  identical  to  that  of  Desmodus  except  base  is  concave  or  flattened;  head 
length  5.61(5.21-5.95)  ±0.249,  acrosome  length  3.20(2.98-3.53)  ±0.170, 
nuclear  length  4.50(4.28-4.74)  ±0.135,  head  width  3.1  1(2.98-3.35)  ±0.104. 


200 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Neck  extremely  short,  junction  with  head  at  center  or  very  slightly  off  center. 
Midpiece  extremely  long  and  extremely  broad  anteriorly;  tapering  abruptly  then 
gradually  posteriorly;  length  12.51(1 1.81-12.83)  ±0.255. 

Remarks. — Sperm  of  Diaemus  youngii  is  very  similar  to  that  of  Desmodus 
rotundus  but  quite  different  from  the  sperm  of  the  third  member  in  the  subfamily, 
Diphylla  ecaudata.  The  difference  in  head  length  between  Desmodus  and  Diaemus 
is  due,  in  part,  to  the  position  of  the  acrosome  on  the  nucleus.  The  midpiece  of 
Diaemus  is  longer  than  any  other  species  of  phyllostomatid  studied  and  appears 
to  lack  the  flared  anterior  end  found  in  the  sperm  of  Desmodus. 

Specimen  examined. — Trinidad:  La  Brea,  St.  Patrick,  1  (CM  45371). 


Diphylla  ecaudata  Spix,  1823 

Description  (Fig.  5G). — Head  clearly  a  shovel-shaped,  extremely  broad, 
rounded  triangle;  acrosome  closely  applied  to  front  of  nucleus  as  in  Desmodus ; 
acrosome  barely  anterior  to  the  nuclear  apex  (in  some  cases  it  cannot  be  seen); 
acrosome  large,  generally  assumes  shape  of  the  nucleus  at  its  apex  but  can  be 
more  pointed;  acrosome  terminates  posteriorly  well  beyond  the  midpoint  of  the 
nucleus  as  in  Desmodus ;  acrosome  the  same  width  as  the  nucleus  throughout  most 
of  its  length;  nucleus  considerably  longer  than  acrosome,  its  base  asymmetrical, 
broad,  with  corners  somewhat  pointed;  a  distinctive  depression  in  base  of  head  at 
junction  with  neck;  head  length  4.57(4.37-4.84)  ±0.160,  acrosome  length  2.89 
(2.70-3.16)  ±0.154,  nuclear  length  4.22(4.02-4.63)  ±0.154,  head  width  3.46 
(3.26-3.62)  ±0.126.  Neck  slightly  longer  and  somewhat  broader  than  other 
vampires;  attachment  to  base  of  head  at  one  comer  of  base.  Midpiece  long,  broad 
anteriorly  and  tapering  gradually  posteriorly;  junction  with  tail  not  distinctive; 
length  9.60(9.21-10.14)  ±0.294. 

Remarks. — Morphology  of  the  sperm  head  of  Diphylla  ecaudata  is  quite 
different  from  the  other  two  species  of  vampires — most  distinctive  is  the  great 
breadth  of  the  nucleus  and  the  attachment  of  the  head  farther  off  center  than  noted 
for  any  other  species  examined. 

Previous  study. — Two  specimens  from  Nicaragua  (Forman  et  al.,  1968). 

Specimen  examined. — Yucatan:  3  km.  S,  1  km.  W  Calcehtoc,  1  (TTU  18447). 

Discussion 

The  spermatozoa  of  35  species  representing  all  six  of  the  subfamilies  of  the 
Phyllostomatidae  were  examined  in  this  study.  Descriptions  of  three  additional 
phyllostomatid  species  are  available  in  the  literature  (Forman,  1968).  The 
morphology  of  all  species  studied  is  basically  similar,  and  this  serves  to  distinguish 
members  of  the  Phyllostomatidae  from  those  of  other  families  of  bats.  The 
acrosome  proved  to  be  the  most  variable  structure,  more  variable  than  even  the 
nuclear  region. 

Below  we  will  discuss  the  relationships  by  subfamily  that  were  observed  in  this 
work. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


201 


Phyllostomatinae. — Acrosomes  within  this  subfamily  were  almost  universally 
asymmetrical  and  always  extended  well  anterior  to  the  nuclear  apex.  Sperm 
from  Mimon  crenulatum  and  Macrotus  waterhousii  were  most  dissimilar  from 
other  members  of  the  subfamily  and  from  each  other.  Mimon  possesses  a 
strikingly  enlarged  and  asymmetrical  acrosome,  whereas  Macrotus  is  characterized 
by  the  unusual  configuration  of  the  nucleus,  particularly  by  its  unique  broad  base. 
Sperm  of  Phyllostomus,  Micronycteris,  and  Tonatia  were  quite  similar,  and 
Phyllostomus  and  Micronycteris  were  characterized  further  among  the  phyllosto- 
matines  by  a  relatively  long  midpiece. 

Glossophaginae. — Heads  of  the  spermatozoa  from  this  subfamily  were  rather 
rounded.  Sperm  from  Choeronycteris  showed  a  larger  head  and  a  substantially 
longer  midpiece  than  either  Anoura  or  Glossophaga.  Anoura  was  distinguished 
from  other  glossophagines  by  a  more  strongly  concave  base  to  the  head  and  from 
other  phyllostomatids  by  an  unusually  short  midpiece. 

Spermatozoa  were  found  to  be  no  more  variable  within  this  subfamily  than 
they  were  among  the  phyllostomatines  or  desmodontines.  Therefore,  sperm 
morphology  does  not  support  the  contentions  based  on  karyology  (Baker,  1967), 
dental  anatomy  (Phillips,  1971),  and  immunologic  comparisons  (Gerber  and 
Leone,  1971)  that  the  glossophagines  are  a  polyphyletic  grouping. 

Carolliinae. — The  sperm  of  three  species  of  the  genus  Carollia  that  have  been 
studied  were  similar,  with  the  nuclei  being  quite  rounded.  However,  the  species 
can  be  distinguished  from  each  other  based  on  overall  head  morphology. 

Stenoderminae. — Morphology  of  the  sperm  heads  of  stenodermines  was  highly 
variable.  Acrosomes  varied  from  pointed  and  nearly  symmetrical  ( Centurio )  to 
broadly  rounded  at  the  apex  and  strongly  asymmetrical  ( Chiroderma ).  There 
was  considerable  variability  in  the  point  of  attachment  of  the  neck  and  midpiece 
to  the  base  of  the  head  and  ranged  from  nearly  central  attachment  to  attachment 
near  the  edge  of  the  base  of  the  head.  However,  the  length  and  breadth  of  the  mid¬ 
piece  of  stenodermines  was  similar,  except  for  Mesophylla,  in  which  the  mid¬ 
piece  was  shorter  than  in  other  species. 

Sperm  from  Ardops,  Ariteus,  Stenoderma,  Phyllops,  and  Artibeus  were 
alike  in  size  and  morphology  of  the  nucleus  and  acrosome.  Members  of  the  first 
four  genera  are  Antillean  endemics  characterized  by  shortened  rostra  and  white 
spots  on  their  shoulders.  These  genera  are  believed  to  have  resulted  from  a  single 
invasion  of  the  Antilles  (Baker  and  Genoways,  1978)  with  subsequent  radiation. 
Morphology  of  the  sperm  supports  this  hypothesis  and  also  suggests  that  members 
of  this  group  may  share  a  close  ancestor  with  members  of  the  genus  Artibeus. 
Uroderma  bilobatum  is  similar  in  morphology  to  members  of  this  group,  except 
that  in  Uroderma  the  base  of  the  head  is  flattened  and  has  pointed  corners. 

Sperm  heads  of  Centurio  senex  were  unusually  triangular  in  form  with  the  base 
of  the  head  unusually  broad.  In  members  of  the  genus  Vampyrops,  the  nucleus 
was  extremely  long,  but  in  Vampyrodes,  the  distinguishing  feature  was  the  narrow 
base  of  the  head.  In  addition  to  the  shortened  midpiece,  Mesophylla  is  char¬ 
acterized  by  the  strongly  asymmetrical  base  of  the  head. 


202 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


The  sperm  of  Chiroderma  improvisum  and  C.  trinitatum  were  the  most 
unique  in  head  morphology  among  the  stenodermines  examined.  In  both  species, 
only  a  very  small  portion  of  the  acrosome  extends  beyond  the  apex  of  the  nucleus. 
Furthermore,  the  base  of  the  nucleus  is  asymmetrical  with  the  greatest  posterior 
extension  occurring  on  the  side  of  the  head  that  is  in  contact  with  the  neck.  The 
sperm  of  these  two  species  are  similar  but  C.  improvisum  can  be  distinguished 
from  C.  trinitatum  by  the  head  of  the  former  being  slightly  less  rounded,  acro¬ 
some  shorter,  and  base  of  head  less  asymmetrical. 

Until  recently,  members  of  the  genus  Sturnira  were  placed  in  a  separate  sub¬ 
family,  Sturnirinae.  However,  recent  authors  (Baker,  1967;  Slaughter,  1970; 
Jones  and  Carter,  1976)  have  placed  them  in  the  subfamily  Stenoderminae.  The 
morphology  of  the  sperm  of  the  three  species  described  herein  were  similar,  all 
being  characterized  by  nearly  symmetrical  acrosomes.  Sperm  head  morphology 
of  species  of  Sturnira  was  most  similar  to  that  of  stenodermines,  among  the  sub¬ 
families  we  examined,  and  we  believe  our  data  support  placement  of  members 
of  the  genus  Sturnira  in  the  subfamily  Stenoderminae.  Although  the  sperm  of  the 
three  species  of  Sturnira  were  similar,  they  could  be  distinguished  on 
the  basis  of  size  and  details  of  morphology. 

Phyllonycterinae. — The  sperm  of  Brachyphylla  cavernarum  was  completely 
unlike  that  of  any  other  phyllonycterines  examined.  Similarity  in  sperm  mor¬ 
phology  does  not  support  placement  of  Brachyphylla  in  the  Phyllonycterinae, 
as  suggested  by  Silva  Taboada  and  Pine  (1969)  from  morphological  and  be¬ 
havioral  investigations  and  Baker  and  Lopez  (1970)  based  on  karyology.  Our 
data  indicate  that  it  would  be  best  to  follow  Miller  (1907)  and  place  Brachyphylla 
in  the  subfamily  Stenoderminae.  Among  the  stenodermines,  the  sperm  of 
Brachyphylla  could  be  distinguished  by  its  long  midpiece. 

The  sperm  head  of  other  phyllonycterine  species  studied  was  more  uniform 
than  that  of  species  within  other  subfamilies;  heads  were  all  relatively  narrow 
and  acrosomes  were  large  and  asymmetrical.  Spermatazoa  from  Erophylla 
hombifrons,  E.  sezekorni,  and  Phyllonycteris  poeyi  were  especially  similar  to 
those  of  Anoura  and  Caro  Ilia. 

The  sperm  of  Phyllonycteris  poeyi  possesses  a  unique  enlargement  in  the  tail 
just  distal  to  its  junction  with  the  midpiece.  This  structure  was  not  seen  in  any 
other  phyllostomatids  examined. 

Desmodontinae. — Sperm  from  the  three  species  of  vampire  bats  were  markedly 
different;  the  only  common  feature  among  the  three  was  a  midpiece  that  proved 
to  be  the  longest  among  the  Phyllostomatidae.  Diphylla  possessed  sperm  heads 
that  were  substantially  broader  and  more  rounded  than  those  of  Desmodus  and 
Diaemus.  The  nuclear  portion  of  the  head  was  similar  in  Desmodus  and  Diaemus\ 
however,  in  Diphylla  the  nucleus  was  broader.  Sperm  from  Diphylla  was  also 
characterized  by  the  neck  and  midpiece  juncture  with  the  head  being  placed 
farther  off  center  than  any  other  phyllostomatid  studied. 

Spermatazoa  of  Desmodus  and  Diphylla  show  great  similarity  in  the  close 
application  of  the  acrosome  to  the  nucleus,  with  little  space  between  the  apices 
of  the  acrosome  and  the  nucleus.  The  acrosome  also  extends  posteriorly  beyond 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


203 


the  midpoint  of  the  nucleus.  Neither  of  these  two  characteristics  appear  in 
Diaemus. 


Literature  Cited 

Baccetti,  B.  1970.  Comparative  spermatology.  Academic  Press,  New  York,  573  pp. 

Baker,  R.  J.  1967.  Karyotypes  of  bats  of  the  family  Phyllostomidae  and  their  taxonomic 
implications.  Southwestern  Nat.,  12:407-428. 

Baker,  R.  J.,  and  H.  H.  Genoways.  1978.  Zoogeography  of  Antillean  bats.  Pp.  53-97,  in 
Zoogeography  in  the  Caribbean  (F.  B.  Gill,  ed.).  Spec.  Publ.  Acad.  Nat.  Sci. 
Philadelphia,  1 3 :  iii  +  1-128. 

Baker,  R.  J.,  and  G.  Lopez.  1970.  Karyotypic  studies  of  the  insular  populations  of 
bats  on  Puerto  Rico.  Caryologia,  23:465-472. 

Bedford,  J.  M.  1967.  Observations  on  the  fine  structure  of  spermatozoa  of  the  bush  baby 
( Galago  senegalensis),  the  African  green  monkey  ( Cercopithecus  aethiops)  and 
man.  Amer.  J.  Anat.,  121:443-459. 

Biggers,  J.  D.  1966.  Reproduction  in  male  marsupials.  Pp.  251-280,  in  Comparative 
biology  of  reproduction  in  mammals  (I.  W.  Rowlands,  ed.),  Academic  Press,  Inc., 
New  York,  xxi  +  559  pp. 

Biggers,  J.  D.,  and  E.  D.  Delamater.  1965.  Marsupial  spermatozoa:  pairing  in  the 
epididymis  of  American  forms.  Nature,  208:402-404. 

Bishop,  M.  W.  H.,  and  C.  R.  Austin.  1957.  Mammalian  spermatozoa.  Endeavour, 
16:137-150. 

Braden,  A.  W.  H.  1959.  Strain  differences  in  the  morphology  of  the  gametes  of  the 
mouse.  Australian  J.  Biol.  Sci.,  ser.  B,  12:65-71. 

Fawcett,  D.  W.,  and  S.  Ito.  1965.  The  fine  structure  of  bat  spermatozoa.  Amer.  J. 
Anat.,  1 16:567-610. 

Forman,  G.  L.  1968.  Comparative  gross  morphology  of  spermatozoa  of  two  families  of 
North  American  bats.  Univ.  Kansas  Sci.  Bull.,  47:901-928. 

Forman,  G.  L.,  R.  J.  Baker,  and  J.  D.  Gerber.  1968.  Comments  on  the  systematic 
status  of  vampire  bats  (family  Desmodontidae).  Syst.  Zool.,  17:417-425. 

Friend,  G.  F.  1936.  The  sperms  of  British  Muridae.  Quart.  J.  Micros.  Sci.,  78:419-443. 

Genoways,  H.  H.  1973.  Systematics  and  evolutionary  relationships  of  spiny  pocket 
mice,  genus  Liomys.  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  5:1-368. 

Gerber,  J.  D.,  and  C.  A.  Leone.  1971.  Immunologic  comparisons  of  the  sera  of  certain 
phyllostomatid  bats.  Syst.  Zool.,  20:160-166. 

Griffiths,  M.  1968.  Echidnas.  Internat.  Ser.  Monogr.  Pure  and  Applied  Biol.,  Zool. 
(G.  A.  Kerkut,  gen.  ed.),  38:ix  +  1-282. 

Helm,  J.  D.,  and  J.  R.  Bowers.  1973.  Spermatozoa  of  Tylomys  and  Ototylomys.  J. 
Mamm.  54:769-772. 

Hirth,  H.  F.  1960.  The  spermatozoa  of  some  North  American  bats  and  rodents.  J. 
Morph.,  106:77-83. 

Hughes,  R.  L.  1964.  Sexual  development  and  spermatozoan  morphology  in  the  male 
macropod  marsupial  Potorotts  tridactylns  ( Ken).  Australian  J.  Zool.,  12:42-51. 

- .  1965.  Comparative  morphology  of  spermatozoa  from  five  marsupial  families. 

Australian  J.  Zool.,  14:533-543. 

Jones,  J.  K.,  Jr.,  and  D.  C.  Carter.  1976.  Annotated  checklist,  with  keys  to  subfamilies 
and  genera.  Pp.  7-38,  in  Biology  of  bats  of  the  New  World  family  Phyllostomatidae, 
Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  D.  C.  Carter,  eds.),  Spec.  Publ.  Mus.,  Texas 
Tech  Univ.,  10:1-218. 

Linzey,  A.  V.,  and  J.  N.  Layne.  1974.  Comparative  morphology  of  spermatozoa  of  the 
rodent  genus  Peromyscus  (Muridae).  Amer.  Mus.  Novitates,  2532:1-20. 

Martin,  D.  E„  K.  G.  Gould,  and  H.  Warner.  1975.  Comparative  morphology  of 
primate  spermatozoa  using  scanning  electron  microscopy.  I.  Families  Hominidae, 
Pongidae,  Cercopithecidae  and  Cebidae.  J.  Human  Evol.,  4:287-292. 


204 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


McFarlane,  R.  W.  1963.  The  taxonomic  significance  of  avian  sperm.  Proc.  Internat. 
Ornith.  Congr.,  8:91-102. 

Miller,  G.  S.,  Jr.  1907.  The  families  and  genera  of  bats.  Bull.  U.S.  Nat.  Mus., 

57:xvii +  1-282. 

Phillips,  C.  J.  1971.  The  dentition  of  glossophagine  bats:  development,  morphological 
characteristics,  variation,  pathology,  and  evolution.  Misc.  Publ.  Mus.  Nat. 
Hist.,  Univ.  Kansas,  54:1-138. 

Silva  Taboada,  G.,  and  R.  H.  Pine.  1969.  Morphological  and  behavioral  evidence  for 
the  relationship  between  the  bat  genus  Brachyphylla  and  the  Phyllonycterinae. 
Biotropica,  1:10-19. 

Slaughter,  B.  H.  1970.  Evolutionary  trends  of  chiropteran  dentitions.  Pp.  51-83, 

in  About  bats  (B.  H.  Slaughter  and  D.  W.  Walton,  eds.),  Southern  Methodist  Univ. 
Press,  Dallas,  Texas,  vii  +  339  pp. 

Wimsatt,  W.  A.,  P.  H.  Krutzsch,  and  L.  Napolitano.  1966.  Studies  on  sperm  survival 
mechanics  in  the  female  reproductive  tract  of  hibernating  bats.  I.  Cytology  and 
ultrastructure  of  intrauterine  spermatozoa  in  Myotis  lucifugus.  Amer.  J.  Anat., 
191:25-59. 

Wooley,  D.  M.,  and  R.  A.  Beaty.  1967.  Inheritance  of  midpiece  length  in  mouse 
spermatozoa.  Nature,  215:94-95. 


ALIMENTARY  TRACT 


G.  Lawrence  Forman,  Carleton  J.  Phillips,  and  C.  Stanley  Rouk 


Bats  of  the  family  Phyllostomatidae  have  extremely  diversified  dietary  habits. 
Although  accurate  and  detailed  dietary  data  often  are  unavailable,  there  never¬ 
theless  are  generalizations  that  can  be  made  and  certain  trends  seem  obvious 
(Gardner,  1977;  Phillips  et  ai,  1977).  In  addition  to  differences  in  diet,  there  also 
are  differences  in  feeding  behavior  and  in  feeding  strategies.  Nonalimentary 
structural  specializations  such  as  reduced  dentitions,  elongate  tongues  (Phillips 
et  al.,  1977),  elaborate  lip  ridges,  and  complex  palatal  topography  also  are 
common  in  leaf-nosed  bats. 

In  view  of  the  great  variability  in  alimentary  function,  it  is  reasonable  to 
hypothesize  that  the  gut  tube  itself  might  be  unusually  variable  within  the 
Phyllostomatidae.  This  is  especially  true  in  comparison  to  other  families  of  bats, 
in  which  the  dietary  habits  are  not  nearly  so  diversified.  Current  data  suggest  that 
at  least  certain  portions  of  the  alimentary  tract  are  in  fact  highly  variable. 

This  account  reviews  what  already  is  known  about  gastrointestinal  structure 
in  phyllostomatids  and  reports  new  information,  particularly  with  regard  to 
histology  and  histochemistry  of  the  stomach.  However,  certain  alimentary  regions, 
such  as  the  intestine  and  esophagus,  still  require  investigation  for  almost  nothing  is 
known  about  them.  A  survey  of  esophageal  structure  could  prove  particularly 
interesting  because  of  the  wide  array  of  food  items  ingested  by  leaf-nosed  bats.  In 
all  likelihood,  the  esophagus  will  reflect  diet-specific  morphological  adaptations. 
Continuing  comparative  analysis  of  digestive  tract  morphology  undoubtedly  will 
prove  important  to  our  understanding  of  systematic  relationships  as  well  as  to  our 
understanding  of  the  evolutionary  process. 

Materials  and  Methods 

Some  information  presented  in  this  chapter  was  extracted  from  a  Ph.D. 
dissertation  by  Rouk  (1973).  In  that  study,  the  following  histological  and  histo- 
chemical  procedures  were  employed:  fixation — 10  per  cent  neutral,  buffered 
formalin;  straining  of  sections — a,  Harris  hematoxylin  and  eosin;  b,  aldehyde- 
fuchsin  for  elastin  and  acid  mucopolysaccharides;  c,  Hale’s  colloidal  iron  followed 
by  acid  fuchsin,  Ponceau  2R,  and  phosphotungstic  acid  sequence  for  acid  muco¬ 
polysaccharides  and  chief  cells;  and  d,  Masson’s  triple  connective  tissue  stain. 

Esophagus 

The  histological  organization  of  the  esophagus  in  phyllostomatids  is  similar 
to  that  of  other  bats  and  other  kinds  of  mammals  as  well.  As  is  typical  for  the 
Chiroptera,  the  phyllostomatid  esophagus  in  preserved  specimens  appears 
to  be  unusually  narrow.  The  luminal  surface  is  characterized  by  protruding 


205 


206 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


longitudinal  folds  of  stratified  squamous  epithelium.  The  esophagus  of  large¬ 
sized  phyllostomatids  can  be  relatively  narrower  than  that  of  smaller  species; 
for  example,  Robin  (1881)  found  that  the  esophagus  of  one  species  of  Artibeus 
was  only  slightly  broader  than  that  of  a  species  of  Glossophaga,  even  though  the 
body  of  the  former  was  three  times  that  of  the  latter. 

Kolb  (1954),  who  reviewed  esophageal  structure  in  bats,  found  some  specific 
variation  in  the  amount  of  cornification  (keratinization)  of  the  esophageal 
epithelium.  He  (Kolb,  1954)  thought  that  such  variation  could  reflect  adaptations 
for  the  ingestion  of  particular  foods.  A  similar  finding  was  reported  for  the  oral 
cavity  (Phillips  et  al.,  1977),  and  it  also  was  suggested  that  the  degree  of 
cornification  could  be  a  local  response  to  a  given  amount  of  surface  stress  rather 
than  a  specific,  inherited  feature.  The  most  complete  histological  study  of  the 
esophagus  of  a  phyllostomatid  is  that  by  Moller  (1932),  who  investigated 
Glossophaga  soricina.  As  might  be  predicted,  he  found  that  the  esophagus  of 
G.  soricina  lacked  significant  corneum,  particularly  in  the  lower  abdominal 
portion.  Cells  lining  the  esophageal  lumen  had  ovoid  nuclei,  unlike  those 
characteristic  of  dead,  cornified  cells.  This  feature  probably  is  reflective  of  the 
general  absence  of  abrasive  food  in  the  diet  of  Glossophaga  and  certainly  is  in 
contrast  to  the  histology  of  insectivorus  species  in  which  the  esophogeal  surface 
is  cornified. 


Stomach 

Comparative  gastrointestinal  structure  and  function  is  of  particular  interest 
because  of  the  variability  in  diet  among  phyllostomatid  species.  It  is  because  of 
this  diversity  in  diet  that  the  phyllostomatids  have  been  subjects  of  more  detailed 
studies  of  alimentary  structure  (especially  the  stomach)  than  have  other 
families  of  bats.  The  following  account,  therefore,  deals  predominantly  with 
morphology  of  the  stomach  because  knowledge  of  variability  in  this  structure  in 
leaf-nosed  bats  even  exceeds  that  for  most  other  groups  of  small  mammals. 
Comments  on  the  small  intestine,  insofar  as  data  are  available,  also  are  included. 

In  most  cases,  stomachs  of  phyllostomatids  can  be  described  in  terminology 
that  has  been  applied  to  other  mammals.  In  those  instances  in  this  account 
where  unusual  or  less  familiar  terms  apply,  a  brief  explanation  parenthetically 
follows  the  term. 

In  all  species  thus  far  studied,  the  stomach  has  the  form  of  a  local  dilation 
of  the  enteron.  Torsion  produces  a  saclike  structure  with  a  lesser  curvature 
(anterior)  and  a  greater  curvature  (posterior).  Specific  variability  in  topography, 
therefore,  has  been  accomplished  by  evolutionary  modification  of  this  general 
plan.  Gastric  glands  occur  throughout  the  mucosa  of  all  species  studied.  Squamous 
epithelium,  on  the  other  hand,  has  been  lacking.  The  summary  given  in  the 
following  paragraphs  is  based  predominantly  on  the  works  of  Forman  (1971a, 
1971b,  1972,  1973),  Rouk  and  Glass  (1970),  and  Rouk(1968,  1973). 


BIOLOGY  OF  THE  PHYLLOSTOM ATIDAE 


207 


d 


Fig.  1. — Semidiagramatic  representations  of  the  stomachs  of  selected  phyllostomatines. 
The  hatched  area  indicates  the  region  of  pylofundic  transition  glands:  a,  Micronycteris 
megalot  is;  b,  Macrophylum  macrophylum;  c,  Tonatia  b  ideas,  d,  Phyllostomus  discolor, 
e,  Phylloderma  septentrionalis,  f,  Vampyrum  spectrum.  Scale  is  10  mm.;  upper  scale  is 
for  figs,  a  to  e;  lower  scale,  f. 


Gross  Morphology 
Phyllostomatinae 

The  phyllostomatines  have  the  simplest  and  least  specialized  stomachs. 
This  probably  relates  to  their  somewhat  unspecialized  or  primitive  feeding 
habits  that  include  insectivorous,  carnivorous,  and  omnivorous  diets.  The 
stomach  in  Micronycteris  is  extremely  simple  in  configuration;  a  cardiac 
vestibule  usually  is  lacking.  The  pyloric  tube  (portion  between  the  esophagus 
and  duodenum)  usually  is  short,  with  that  of  M.  nicefori  being  relatively  longer 
than  that  of  M.  hirsuta  or  M.  megalotis  (Fig.  la).  The  fundic  caecum  ( =  cardiac 
caecum)  is  modestly  developed  in  all  three  of  these  species.  The  stomachs  of 
Macrotus  waterhousii  and  Macrophyllum  macrophyllum  (Fig.  lb)  also  are 
simple  and  generally  resemble  those  of  Micronycteris. 


208 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


The  stomach  in  Tonatia  varies  somewhat  from  those  previously  mentioned, 
and  that  of  Tonatia  minuta  varies  intraspecifically.  For  example,  the  stomach  of 
T.  minuta  may  have  a  poorly  developed  fundic  caecum.  Additionally,  the  pyloric 
tube  is  bent  at  a  right  angle  to  the  general  orientation  of  the  stomach,  as  observed 
in  T.  bidens  (Fig.  lc),  or  it  may  more  closely  approximate  the  simple,  symmetrical 
configuration  found  in  species  of  Micronycteris.  The  esophageal  entrance  is  located 
about  midway  along  the  lesser  curvature.  The  pyloric  tube  in  Chrotopterus  auritus 
differs  from  that  in  species  of  Micronycteris  only  in  being  relatively  longer. 

The  stomach  of  Phylloderma  stenops  (Fig.  le)  is  more  globular  than  those 
of  other  phyllostomatines,  but  otherwise  it  does  not  differ  substantially  from  those 
found  in  species  of  Micronycteris.  The  stomach  of  Trachops  is  Micronycteris- like 
but  still  is  more  tubular,  and  the  lesser  and  greater  curvatures  are  nearly  parallel. 

The  stomachs  of  several  other  phyllostomatines  differ  more  distinctively 
from  the  Micronycteris- like  configuration.  For  example,  in  Phyllostomus 
discolor  (Fig.  Id)  and  P.  hastatus  the  fundic  caecum  is  well  developed  and 
often  is  dilated  at  its  terminus.  The  pyloric  portion  is  distinctively  elongated 
and  sometimes  there  is  a  prominent  constriction  in  front  of  the  gastroduodenal 
junction.  A  small,  but  perceptable  cardiac  vestibule  occurs  between  the  lesser 
curvature  and  the  gastroesophageal  junction.  Although  this  vestibule  is  not 
nearly  so  expansive  as  that  in  some  frugivores,  it  nevertheless  is  more 
distinctive  than  that  of  phyllostomatines  described  above.  The  stomach  of  P. 
hastatus  generally  resembles  that  of  P.  discolor ,  except  for  its  considerably  larger 
size.  The  greater  and  lesser  curvatures  are  nearly  parallel  in  both  species. 

The  stomach  of  Vampyrum  spectrum  (Fig.  If),  a  carnivore  that  often  feeds 
on  other  bats  (see  Rouk,  1973),  is  noticeably  pearshaped  with  a  moderately 
developed  fundic  caecum  and  a  long,  well  differentiated  pyloric  tube.  A  cardiac 
vestibule  is  lacking  and  the  lesser  curvature  is  longer  than  in  other  phyllosto¬ 
matines.  This  is  because  the  pyloric  tube  exits  to  the  side  (right  side  of  the  body) 
with  only  very  slight  anterior  recurvature  of  the  terminal  portion  of  the  stomach. 
The  stomach  of  this  species,  with  its  straight  pyloric  tube,  has  a  strong  resemblance 
to  those  of  many  species  of  the  Insectivora  (see  Allison,  1948;  Myrcha,  1967). 

Simplicity  of  stomach  form  is  evident  in  the  Phyllostomatinae.  Some  elongation 
of  the  pyloric  portion,  along  with  some  dilation  of  the  caecum  also,  is  evident 
in  comparison  with  stomachs  of  insectivorous  bats  of  other  families.  These  slight 
modifications  likely  are  associated  with  increased  volume  of  food  ingested. 

Glossophaginae 

The  stomach  of  Glossophaga  soricina  (Fig.  2a)  is  large  and  saccular. 
Although  its  diet  includes  insects  along  with  nectar,  pollen,  and  fruit,  the 
stomach  is  decidedly  more  specialized  than  that  of  any  of  the  Phyllostomatinae, 
including  the  omnivorous  Phyllostomus  discolor. 

The  fundic  caecum  in  G.  soricina  is  dilated  and  bulbar.  The  caecum  can  be 
distinguished  from  the  remainder  of  the  stomach  by  a  distinctive  furrow  or 
sulcus  on  the  dorsal  surface.  The  stomach  is  curved  in  both  frontal  and  transverse 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


209 


FC 


d  e 


Fig.  2. — Semidiagramatic  representations  of  the  stomachs  of  selected  glossophagines 
and  a  carolliine.  The  hatched  area  indicates  the  region  of  pylofundic  transition  glands: 
a,  Glossophaga  soricina;  b,  Hylonycteris  underwoodr,  c,  Lonchophylla  robusta;  d, 
Lichonycteris  obscurer,  e,  Carol! ia  perspicillata.  Scale  is  10  mm.  for  e;  for  all  others, 
8  mm.  Symbols  are  FC,  fundic  caecum;  CV,  cardiac  vestibule. 

planes.  A  small  cardiac  vestibule  has  been  observed  in  some  specimens,  but 
seems  to  be  absent  in  others.  This  variable  feature  possibly  is  an  individual 
response  to  opportunistic  feeding  by  this  species.  Glossophaga  commissarisi 
has  a  stomach  that  is  similar  to  that  of  G.  soricina  except  for  its  even  more 
distinctive  cardiac  vestibule.  The  fundic  caecum  is  relatively  longer  and  narrower 
than  that  of  G.  soricina.  The  pyloric  tube  is  enlongated  and  more  distinctive 
than  in  G.  soricina. 

Even  though  stomachs  of  Hylonycteris  underwoodi  (Fig.  2b),  Lonchophylla 
robusta,  Anoura  geoffroyi,  Choeronycteris  mexicana,  and  Leptonycteris  all 
bear  a  general  resemblance  to  those  in  Glossophaga ,  distinguishing  characteristics 
can  be  observed  in  most.  For  example,  Hylonycteris  has  a  relatively  long,  narrow 
fundic  caecum  (Fig.  2b)  that  is  nearly  tubular  and  is  marked  by  numerous  deep 
sulci.  The  extemely  broad  pyloric  tube  is  short,  but  decidedly  arched  from  left 
to  right.  The  stomachs  of  Anoura  geoffroyi  and  Choeronycteris  mexicana  bear 
striking  resemblance  to  those  of  Glossophaga.  In  comparison  to  the  other 
glossophagines,  Lonchophylla  robusta  has  an  unusual  stomach  (Fig.  2c)  in 
that  both  the  cardiac  vestibule  and  fundic  caecum  are  developed  distinctively. 
The  gross  morphology  of  this  stomach  approaches  that  of  some  fruit-eating 
stenodermines. 

The  stomachs  of  Leptonycteris  nivalis  and  L.  sanborni  are  nearly  identical. 
They  also  are  somewhat  distinctive  because  of  an  unusually  elongated,  extremely 
pointed  fundic  caecum.  Also,  the  terminal  portion  of  the  stomach  (pylorus)  is 
tubular  and  elongated  to  the  point  of  being  recurved  to  lie  juxtaposed  to  the 
cardiac  vestibule.  Therefore,  the  stomach  assumes  a  C-shaped  configuration 


210 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


when  viewed  from  the  front.  This  striking  elongation  and  recurvature  of  the 
pyloric  stomach  in  Leptonycteris  and  in  Lichonycteris  as  well  (and  to  a  lesser 
extent  in  Choeroniscus  and  Glossophaga)  might  represent  an  adaptation  to 
permit  intake  of  an  increased  percentage  of  plant  material  in  the  diet.  Increased 
length  of  the  pyloric  tube  is  one  way  to  increase  gastric  volume. 

The  stomachs  of  Choeroniscus  godmani  and  Lichonycteris  obscura  (Fig.  2d) 
possess  well-developed  cardiac  vestibules  and  broad  terminal  portions  that 
can  be  recurved  sharply  toward  the  gastroesophageal  junction.  The  fundic 
caeca  of  these  two  species  are  shallow;  unlike  the  other  species  of  glossophagines, 
the  caeca  are  not  delineated  by  a  sulcus  ( =  incisura  cardiaca)  from  the  cardiac 
vestibule.  Therefore,  the  vestibule  merges  gradually  into  the  caecum  on  the 
greater  curvature  of  these  two. 


Carolliinae 

The  stomachs  of  two  species  from  this  subfamily  have  been  examined. 
Carol lia  perspicillata  (Fig.  2e)  and  C.  castanea  generally  are  quite  similar 
but  apparently  are  individually  variable  in  gross  morphology.  The  terminal 
( =  pyloric)  portion  is  elongate  and  strongly  recurved  anteriorly.  This  recurvature 
possibly  functions  to  retard  gastric  emptying.  A  cardiac  vestibule  is  present; 
in  some  specimens  it  is  moderately  developed,  whereas  in  others  it  is  quite  small. 
The  caecum  is  baglike  and  dilated  and  is  more  prominent  in  C.  castanea  than 
it  is  in  C.  perspicillata.  Overall,  the  stomachs  of  these  two  species  are  in  many 
ways  intermediate  between  those  of  glossophagines  and  those  of  stenodermines. 
The  Carolliinae  exhibit  the  overall  simplicity  of  most  glossophagine  stomachs 
in  combination  with  some  specialization  of  the  caecum  (especially  the  pyloric 
tube),  which  is  characteristic  of  fruit-eating  stenodermines. 

Stenoderminae 

An  extensive  array  of  stenodermine  species,  most  of  which  are  considered 
to  be  frugivores,  have  been  studied.  The  stomachs  of  stenodermines  are 
substantially  more  complex  and  more  specialized  than  those  of  the  previously 
described  species.  Virtually  all  gross  features  of  the  stomach  are  enlarged  or 
lengthened,  especially  in  comparison  with  the  simpler  stomachs  of  the 
phyllostomatines  and  glossophagines. 

The  stomachs  of  Sturnira  lilium  and  5.  ludovici  (Fig.  3a)  are  similar  to  one 
another.  In  S.  lilium,  which  is  typical,  the  cardiac  vestibule  is  elongate  and  tapers 
so  that  the  gastroesophageal  junction  lies  well  superior  to  the  gastroduodenal 
junction.  The  fundic  caecum  is  saccular  and  thinwalled,  forming  a  spacious 
chamber  with  an  apex  that  varies  from  being  rounded  to  being  tapered.  A  fold 
of  the  stomach  wall  distinguishes  the  cardiac  vestibule  from  the  fundic  caecum. 
The  tubular  (  =  pyloric)  portion  of  the  stomach  is  long  and  narrow  (S.  ludovici 
has  a  shorter  pylorus  and  a  somewhat  larger  cardiac  vestibule  giving  the  stomach 
a  more  robust  appearance  than  that  of  S.  lilium).  The  stomach  from  a  single 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


211 


FC 


Fig.  3. — Semidiagramatic  representations  of  the  stomachs  of  selected  stenodermines. 
The  hatched  area  indicates  the  region  of  pylofundic  transition  glands:  a,  Sturnira  ludovicr, 
b,  Uroderma  magnirostrunr,  c,  Artibeus  lituratus ;  d,  Centurio  senex ;  e,  Vampyrodes 
caraccioli ;  f,  Chiroderma  villosum.  Scale  is  10  mm.  Symbols  are  identified  in  Fig.  2. 

specimen  of  S.  mordax  was  examined  by  Rouk  (1973)  who  found  it  to  have  a 
a  considerably  simpler  gross  morphology  than  those  of  other  species  of 
Sturnira.  Rouk  (1973)  reported  that  the  terminal  portion  was  relatively 
unspecialized  and  that  the  caecum  was  poorly  developed.  However,  the 
stomach  in  S.  mordax  does  possess  a  moderately  large  cardiac  vestibule. 

The  remaining  stenodermines  for  which  stomachs  have  been  examined  show 
increased  specialization  by  way  of  elongation  or  enlargement  of  one  or  more 
portions  of  the  stomach.  The  stomachs  of  seven  species  of  Artibeus  ( aztecus , 
inopinatus,  jamaicensis,  lituratus,  phaeotis,  toltecus,  and  watsoni)  have  been 
studied  (see  Fig.  3c).  These  seven,  along  with  that  of  Centurio  senex  (Fig.  3d), 


212 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


have  tremendously  enlarged  cardiac  vestibules  that  permit  temporary  storage 
of  large  amounts  of  plant  material.  In  Vampyressa,  Vampyrops,  Uroderma, 
Vampyrodes,  and  Chiroderma,  the  cardiac  vestibule  varies  from  small  to 
moderately  large,  with  the  fundic  caecum  being  variously  drawn  out  into  a  baglike 
or  nearly  tubelike  structure. 

Stomachs  of  Uroderma  bilobatum  and  U.  magnirostrum  (Fig.  3b)  share 
gross  characteristics  with  Sturnira,  as  well  as  with  Artibeus,  and  could  be 
said  to  be  intermediate  between  the  two.  The  caecum  is  elongate  and  narrowed 
nearly  to  a  point  at  its  apex.  The  stomachs  of  Vampyrops  helleri  and  V.  vittatus 
differ  from  that  of  Uroderma  only  slightly  in  that  the  caecum  of  V.  helleri 
and  V.  vittatus  is  somewhat  broader. 

The  stomachs  of  Vampyressa  pusilla  and  V.  nymphaea  are  nearly  identical 
to  one  another.  The  cardiac  vestibule  is  small  in  comparison  with  most  of  the 
other  stenodermines.  The  elongate  fundic  caecum  is  recurved  anteriorly,  as 
it  is  in  Uroderma,  Vampyrops,  Vampyrodes,  Chiroderma ,  and  some  Artibeus, 
and  it  is  dilated  at  its  terminus. 

The  remaining  two  species  to  be  discussed  in  this  account,  Vampyrodes 
caraccioli  (Fig.  3e)  and  Chiroderma  villosum  (Fig.  3f),  possess  greatly 
enlarged  fundic  caeca.  The  stomach  of  Vampyrodes  somewhat  resembles  that 
of  Uroderma  except  that  the  cardiac  vestibule  is  much  reduced.  A  distinctive 
narrowing  occurs  between  the  cardiac  vestibule  and  fundic  caecum  of  both 
species  so  that  there  is  only  a  small  region  where  the  two  are  contiguous.  The 
fundic  caecum  of  Vampyrodes  is  about  1.5  times  the  length  of  the  remainder 
of  the  stomach,  and  that  of  Chiroderma  is  in  excess  of  twice  the  length. 

The  stomach  of  C.  villosum,  which  has  a  tubular  caecum,  represents  perhaps  the 
most  extreme  specialization  for  plant  feeding  in  the  Phyllostomatidae.  This  con¬ 
dition  closely  parallels  that  observed  in  some  Old  World  megachiropterans.  The 
caecum  is  marked  externally  by  a  series  of  parallel  constrictions  that  surround  it 
for  nearly  its  entire  length.  The  duodenum  at  the  gastrointestinal  junction  is  unusual 
in  being  grossly  dilated  on  the  lesser  curvature  to  produce  what  amounts  to  a  small 
ampulla  or  caecum.  The  function  of  this  dilation  is  unknown. 

It  would  appear  that  there  are  two  adaptive  trends  within  the  Stenoderminae. 
Each  apparently  represents  a  different  response  to  increased  need  for  stomach 
volume  in  these  frugivores.  One  trend,  which  is  best  illustrated  in  Artibeus  and  in 
Centurio,  was  to  increase  size  of  the  cardiac  vestibule  while  minimizing  the  impor¬ 
tance  of  the  fundic  caecum.  The  other  approach,  seen  so  vividly  in  such  genera  as 
Vampyressa,  Vampyrodes,  and  Chiroderma,  was  to  minimize,  or  even  to  nearly 
eliminate,  the  cardiac  vestibule  while  correspondingly  enlarging  the  caecum  into 
an  obviously  useful  storage  chamber.  Both  trends  would  permit  increased  con¬ 
sumption  or  storage,  or  both,  of  plant  materials  that  presumably  are  difficult  to 
digest. 


Phyllonycterinae 

Rouk  (1973)  examined  the  stomach  of  only  one  member  of  this  subfamily, 
Brachyphylla  cavernarum  (Fig.  4a).  The  esophagus  enters  the  stomach  quite  near 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


213 


the  gastroduodenal  junction.  Therefore,  the  lesser  curvature  between  esophagus 
and  duodenum  is  extremely  short.  The  fundic  caecum  is  extremely  well  developed 
into  a  “bag”  that  appears  to  be  nearly  compartmentalized  into  a  two-chambered 
structure.  The  caecum  bends  abruptly  anteriorly  about  midway  along  its  length. 
At  this  location,  there  is  a  suggestion  of  a  sphincter,  although  this  constriction  in 
the  muscularis  externa  has  not  been  demonstrated  to  have  a  sphincteric  function. 
The  duodenum  is  quite  enlarged  at  its  junction  with  the  stomach,  which  is  separated 
from  the  intestine  by  a  distinctive  constriction.  The  stomach  of  Brachyphylla 
clearly  is  distinctive  among  phyllostomatids.  Other  phyllonycterines  should  be 
examined  to  determine  if  this  distinctive  form  is  consistent  within  the  group. 

Desmodontinae 

The  gastric  morphology  of  Desmodus  rotundus  (Fig.  4b)  has  been  variously 
described  and  illustrated  by  a  number  of  workers  (Huxley,  1865;  Rouk  and 
Glass,  1970;  Hart,  1971;  Forman,  1972).  Its  simple,  tubular  form  is 
predominately  an  elongate  caecum  of  generally  uniform  breadth  that  lacks 
a  cardiac  vestibule  or  demonstrable  pyloric  portion  (although  pyloric  glands 
are  present).  The  terminal-most  part  of  the  caecum  frequently  is  dilated  into 
a  thin-walled  sac;  the  distal  one-half  is  folded  back  upon  the  proximal  one-half. 
There  is  no  conclusive  evidence  of  any  sphincters  within  the  stomach,  except 
for  that  adjacent  to  the  duodenum. 

In  Diaemus  youngii  (Fig.  4c),  the  stomach  bears  strong  resemblance  to  that  of 
Desmodus  except  that  the  caecum  may  be  less  tubular  and  more  conical  in  this 
species.  The  terminal  part  of  the  caecum  is  slightly  dilated.  In  the  stomach  of 
Diphylla  ecaudata  (Fig.  4d),  numerous  semilunar  folds  within  the  distal  one-half 
of  the  caecum  divide  it  into  smaller  compartments.  The  caecum,  with  its  haustra 
coli,  therefore,  bears  strong  resemblance  to  the  colon  of  man.  The  “pouches”  thus 
formed  in  the  caecum  of  Diphylla  would  tend  to  retard  gastric  emptying,  important 
in  vampires  because  the  stomach  is  specialized  for  absorption.  Additionally,  the 
folds  in  the  caecum  would  tend  to  increase  the  surface  area  to  volume  ratio,  thereby 
increasing  the  efficiency  of  absorption  from  the  stomach. 

Gastric  Mucosa 

The  stomachs  of  all  species  of  phyllostomatids  are  completely  lined  with  a 
glandular  mucosa.  There  is  no  uncornified  or  cornified  squamous  epithelium 
in  the  stomach.  A  zone,  usually  narrow,  of  mucuous-producing  cardiac  glands 
is  found  at  the  gastroesophageal  junction.  A  broader  zone  of  pyloric  glands, 
which  also  are  mucuous  producing  and  which  are  similar  in  structure  to  cardiac 
glands,  are  located  at  the  gastroduodenal  junction  in  all  species.  The  remainder 
of  the  mucosa  is  occupied  by  a  broad  region  of  fundic  glands  composed  of 
mucous  cells,  parietal  cells,  and  chief  ( =  zymogenic)  cells.  A  zone  of 
transitional  glands  that  is  extremely  variable  in  length  occurs  between  fundic 
and  pyloric  mucosa.  This  transitional  area  is  rather  broad  in  species  of  the 
Glossophaginae  but  is  relatively  narrow  in  the  Stenoderminae.  Species  of 


214 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  4. — Semidiagrammatic  representations  of  the  stomachs  of  one  phyllonycterine 
and  three  desmodontines.  The  hatched  area  indicates  the  region  of  pylofundic  transition 
glands:  a,  Brachyphylla  cavernarunr,  b,  Desmodus  rotundas, ;  c,  Diaemus  youngii,  d, 
Diphylla  ecaudata.  Scale  is  10  mm.;  upper  scale  is  for  a;  lower  scale,  all  others. 

Artibeus,  along  with  Centurio  and  Vampyrodes,  consistently  have  extremely 
narrow  “transition”  zones.  This  narrowness  of  the  transition  zone  seems  to  be 
due  to  a  relatively  extensive  proximal  advancement  of  pyloric  glands  within 
the  pyloric  tube. 

Depth  of  the  gastric  mucosa  varies  slightly  within  stomachs  and  among 
species.  The  mucosa  is  shallowest  in  the  vampires,  with  fundic  glands  being 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


215 


only  50  to  75  micrometers  in  Desmodus.  The  gastric  glands  of  vampires  are 
reduced  to  shallow  acini  in  comparison  to  the  tubular  form  of  other  species. 
This  is  accompanied  by  a  general  reduction  in  all  cellular  constituents, 
although  zymogenic,  parietal,  argentiffin,  and  mucous  neck  cells  all  are 
present.  Mucous  neck  cells  comprise  the  most  abundant  cellular  component 
of  the  mucosa,  whereas  parietal  ( =  HCl-producing)  cells  are  extremely  sparse. 

The  gastric  mucosa  of  other  species  varies  from  100  to  600  micrometers, 
in  depth,  although  200  to  250  micrometers  is  most  commonplace.  Pyloric 
glands  frequently  are  longer  than  are  the  fundic  glands  within  a  species;  for 
example,  in  Artibeus  they  are  50  to  80  per  cent  longer.  In  many  species,  the 
fundic  glands  are  somewhat  longer  at  the  apices  of  rugae  than  on  the  stomach 
wall  proper.  In  striking  contrast  is  the  fundic  portion  of  the  mucosa  of 
stenodermines,  such  as  Artibeus  and  Centurio,  in  which  the  glands  are  of 
extremely  uniform  depth.  Relative  constancy  of  cell  frequency  accounts  for  the 
uniformity  of  mucosal  depth.  In  some  phyllostomatines,  especially  Micronycteris 
and  Chrotopterus,  the  fundic  mucosa  is  quite  shallow  at  the  apex  of  the  caecum. 

The  stomach  wall  of  all  species  is  thrown  into  rugae,  which  occur  everywhere 
within  the  stomach.  These  folds  generally  are  oriented  along  the  longitudinal 
axis  and  are  arranged  in  parallel  rows  in  the  terminal,  tubular  stomach.  They 
occur  in  wavy,  parallel  rows  throughout  the  remainder  of  the  stomach  in  many 
other  species.  In  stenodermines,  all  species  that  have  been  examined  with  respect 
to  rugal  organization  reveal  some  degree  of  “complication”  or  interdigitation 
of  folds.  In  Vampyressa,  V ampyrops,  Chiroderma,  and  Sturnira,  they  are 
distributed  diagonally  (toward  the  pyloric  sphincter),  but  only  within  the  caecum. 
Rugae  are  slanted  only  within  the  midregion  of  Uroderma.  In  most  stenodermines 
that  have  been  studied,  folds  interdigitate  only  to  a  moderate  degree,  but  in 
Artibeus  and  Centurio  an  extremely  complex  interlocking  of  folds  produces 
an  elaborate  maze  because  folds  are  highly  branched.  This  arrangement  likely 
would  be  effective  in  retarding  gastric  emptying,  a  particularly  important 
digestive  adaptation  in  obligate  plant  feeders. 

Histochemistry  of  the  Gastric  Mucosa 

Few  systematic  groupings  of  mammals  have  been  examined  comparatively 
with  respect  to  the  histochemistry  or  cytochemistry  of  the  stomach  lining. 
Phyllostomatids  are  an  exception  to  this  in  that  the  mucous  cells  and  their 
secretory  products  have  been  studied  with  a  variety  of  techniques.  Procedures 
have  been  employed  that  elucidate  acid  as  well  as  neutral  mucopolysaccharides. 

A  positive  periodic  acid-Schiff  (PAS)  reaction  is  thought  to  indicate  an 
abundance  of  mucosubstance  and,  thus,  it  provides  an  overall  estimate  of 
the  quantity  of  mucus  within  or  on  the  surface  of  cells  in  the  stomach  or  intestine 
(see  Lillie,  1965).  In  all  examined  species  phyllostomatids,  there  is  a  moderate 
to  intense  coloration  of  mucous  material  in  the  apical  portion  of  the  cytoplasm 
of  surface  columnar  cells.  In  Desmodus  rotundus  (the  only  desmodontine  exam¬ 
ined  to  date),  the  intensity  of  this  reaction  in  surface  mucus  is  somewhat  reduced 


216 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


in  comparison  with  that  of  other  phyllostomatids.  In  many  species  having  well- 
developed  fundic  caeca,  the  staining  is  stronger  in  the  foveolae  of  the  fundic 
glands  of  the  caecum  than  elsewhere  in  the  fundus.  Mucus  possibly  accumulates 
to  a  greater  extent  in  the  caecum  than  elsewhere  in  phyllostomatids. 

Mucous  cells  beneath  the  surface  (the  so-called  mucous  neck  cells),  which  are 
scattered  among  the  parietal  cells,  react  much  more  variably  to  the  PAS  reaction 
than  do  the  surface  columnar  cells.  Mucous  neck  cells  of  frugivorous  species 
generally  are  less  reactive  than  are  those  of  carnivorous  and  omnivorous  kinds. 
Those  of  Desmodus  (and  perhaps  the  other  desmodontines)  react  only  weakly. 

The  upper  portions  of  the  tubules  of  cardiac  and  pyloric  glands  stain  intensely 
with  PAS.  There  is  only  slight  variability  among  species.  As  in  the  case  of  fundic 
glands,  reactivity  in  these  upper  portions  is  somewhat  reduced  in  frugivorous 
species.  Among  studied  species,  the  most  intense  reaction  has  been  found  in  an 
omnivore,  Phyllostomus  discolor.  The  quantity  of  gastric  mucus  in  this  species 
exceeds  that  of  frugivorous  phyllostomatids.  On  the  other  hand,  in  Desmodus 
the  reactivity  is  weak  in  comparison  with  nondesmodontine  phyllostomatids. 

Two  procedures,  or  their  variants,  have  been  employed  in  an  effort  to  elaborate 
the  relatively  acidic  components  of  gastric  mucus  in  phyllostomatids.  Forman 
(1972)  employed  Alcian  blue  8GX,  and  Rouk  (1973)  and  Forman  (19716) 
used  Hale’s  colloidal  iron  procedure  in  efforts  to  categorize  acid 
mucopolysaccharides  in  stomachs  of  selected  species  of  phyllostomatids. 
A  summary  of  their  results  is  presented  here. 

Acid  mucopolysaccharides  are  found  most  consistently  in  the  cardiac  glands 
(those  at  the  gastroesophageal  junction)  and  within  the  few  transitional  and 
fundic  glands  adjacent  to  the  cardiac  glands.  Nearly  all  species  of  phyllostomatids 
studied  to  date  showed  some  positive  staining  of  cardiac  glands.  The  only 
exceptions  are  species  of  Sturnira  (including  S.  lilium,  S.  ludovici,  and  S. 
mordax ).  In  these  species,  the  cardiac  glands  are  either  weakly  reactive  or 
non  reactive  to  procedures  intended  to  demonstrate  the  presence  of  acid 
mucopolysaccharides.  Present  evidence  also  suggests  that  Centurio  and  Desmodus 
have  reduced  amounts  of  acid  mucopolysaccharides  in  their  cardiac  glands. 
The  reaction  of  the  pyloric  glands  to  Hale’s  colloidal  iron  and  Alcian  blue  is 
similar  to  that  of  the  cardiac  glands.  There  is,  however,  less  consistency  among 
species,  less  uniformity  within  the  zone  of  pyloric  glands,  and  often  less  intensity 
in  comparison  to  the  histologically  similar  cardiac  glands. 

In  most  species  of  phyllostomatines,  the  pyloric  glands  are  nonreactive; 
the  exception  is  Vampyrum  spectrum,  in  which  these  glands  are  weakly 
reactive  with  Hale’s  colloidal  iron. 

In  the  glossophagines,  there  are  two  general  conditions  of  stainability  of  the 
pyloric  glands  with  Alcian  blue  and  Hale’s  colloidal  iron.  With  Hale’s  iron 
(as  employed  by  Rouk,  1973)  pyloric  glands  stain  intensely  within  the  basal 
one-third  of  the  tubules  in  Glossophaga  soricina  and  Lonchophylla  robusta. 
Forman  (19716)  studied  glossophagine  cardiac  glands  with  Alcian  blue.  In  his 
study  of  five  species  of  glossophagines,  the  lower  portion  of  each  pyloric  gland 
tubule  was  Alcian  blue  positive  in  three  ( Glossophaga  soricina,  G.  commissarisi, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


217 


and  Anoura  geoffroyi)  but  negative  in  two  others  ( Choeroniscus  godmani  and 
Lichonycteris  obscura). 

Among  the  phyllostomatids,  the  most  widespread  and  distinctive  reactivity 
to  procedures  for  acid  mucopolysacchardies  in  the  stomach  are  found  in  certain 
of  the  carolliines  and  stenodermines.  For  example,  pyloric  glands  in  Vampyrodes, 
Vampyressa,  Chiroderma,  Centurio,  and  in  seven  species  of  Artibeus  that  have 
been  studied,  react  intensively  with  Hale’s  colloidal  iron  either  throughout  or 
nearly  throughout  the  length  of  the  tubule.  Rouk  (1973)  determined  that  nearly 
all  glands  in  the  stomach  of  Vampyressa  pusilla  contain  noteworthy  amounts 
of  Hale  positive  mucin.  In  these  same  stenodermines,  as  well  as  in  Uroderma, 
V ampyrops,  and  Sturnira  mordax,  the  mucous  neck  cells  within  the  upper 
portions  of  fundic  gland  tubules  also  react  moderately  or  strongly  with  Hale’s 
iron.  Reactivity  in  these  cells  rarely  has  been  observed  in  nonstenodermines. 

These  results  suggest  that  a  relationship  might  exist  between  gastric  acid 
mucopolysaccharides  and  plant  feeding  in  phyllostomatids.  Whether  their 
function  is  protective,  digestive,  or  both  remains  to  be  determined. 

Pyloric  Sphincter 

The  muscular  portion  of  the  sphincter  at  the  gastroduodenal  junction  is 
unusually  variable  in  form  in  phyllosotmatids.  Numerous  variations  in 
the  form  of  this  circular  muscle  mass  have  been  observed  in  leaf-nosed  bats, 
and  at  least  part  of  this  variability  appears  to  be  related  to  diet.  The  sphincter 
is  in  some  way  asymmetrical  in  the  majority  of  species  that  have  been  examined. 
In  kinds  where  asymmetry  is  present,  the  valve  on  the  greater  curvature  is  larger 
than  that  portion  on  the  lesser  curvature.  This  condition  always  prevails  in 
insectivorous  and  carnivorous  species.  The  valves  of  Macrotus,  Micronycteris, 
Tonatia  minuta ,  and  Glossophaga  are  short  to  moderate  in  length  and  generally 
are  robust  with  broadly  rounded  apices.  In  Centurio,  the  valve  of  the  greater 
curvature  is  fully  three  times  the  mass  of  the  “lesser”  valve.  This  form  of  valvular 
asymmetry  is  maximized  in  Tonatia  minuta  in  which  the  greater  valve  is  long  and 
extremely  thick,  whereas  the  lesser  valve  is  absent,  or  nearly  so.  Two  noteworthy 
instances  in  which  the  valve  is  greatest  in  mass  on  the  lesser  curvature  are  found  in 
Uroderma  bilobatum  and  in  Chiroderma  villosum.  This  asymmetry  might  result 
in  some  sort  of  “milking”  action  that  permits  slow  release  of  stomach  contents  into 
the  duodenum. 

Two  trends  in  pyloric  sphincter  morphology  are  evident  in  frugivorous 
species  as  well  as  in  some  pollenivorous  and  nectarivorous  kinds.  One  trend 
involves  increased  symmetry,  whereas  the  other  involves  the  amount  of  muscular 
contribution  to  the  valve. 

First,  the  pyloric  valve  of  some  fruit-eating  stenodermines  and  carolliines, 
including  Artibeus,  Sturnira,  Vampyressa,  Carollia  perspicillata  and  perhaps 
others,  is  of  nearly  uniform  length  throughout  its  circumference.  It  would 
appear  that  increased  symmetry  of  the  valve  in  these  species  is  related  to 
consumption  of  plant  material.  None  of  the  insectivorous  or  carnivorous  kinds 
has  a  symmetrical  valve;  indeed,  the  most  pronounced  asymmetry  always  is 


218 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


observed  in  these  species.  The  pyloric  valve  of  Desmodus  rotundus  is  reduced 
in  bulk,  as  compared  with  other  phyllostomatids,  but  it  also  is  nearly  symmetrical. 
It  is  possible  that  symmetry  may  be  related  to  passage  of  liquid  food  into  the 
duodenum,  both  in  vampires  and  in  plant  feeders. 

Second,  bats  that  consume  plant  material  including  fruit,  nectar,  and 
pollen  have  a  valve  flap  that  nearly  always  is  longer  and  thinner  than  valves 
of  bats  that  eat  animal  material.  This  feature  is  particularly  well  developed  in 
stenodermines  and  in  Brachyphylla  cavernarum.  In  species  of  Artibeus, 
Centurio,  Chiroderma,  Uroderma,  Vampyressa,  Vampyrops,  and  Vampyrodes, 
the  flap  achieves  such  length  that  its  apex  is  directed  up  into  the  duodenum. 
This  results  in  valve  flaps  that  are  parallel  with  the  intestinal  wall.  In  addition, 
the  apex  of  the  muscular  flap  is  quite  pointed  in  species  of  Artibeus.  Most 
glossophagines  that  have  been  examined,  including  species  of  Lonchophylla, 
Lichonycteris,  Choeronycteris,  and  Hylonycteris,  but  excluding  Glossophaga, 
have  thin  valves  that  are  similar  to  those  of  stenodermines.  Anoura  and 
Leptonycteris  are  intermediate  between  the  Glossophaga- type  and  stenodermine- 
type  valve,  but  most  similar  to  the  latter.  It  is  reasonable  to  hypothesize  that 
these  longer,  thinner,  often  symmetrical  valve  flaps  might  improve  the 
efficiency  of  gastric  closure,  thus  delaying  gastric  emptying  and  improving 
digestion  (by  increasing  time)  in  these  plant  feeders. 

The  pyloric  sphincters  of  Sturnira  lilium  and  S.  ludovici,  although  symmetrical, 
are  unique  in  that  identifiable  muscular  flaps  either  are  absent  or  nearly  so  as 
barely  to  be  perceptable.  The  functional  significance  of  this  apparent  degeneracy 
is  unknown. 


Tunica  Muscularis 

All  stomachs  of  phyllostomatids  possess  two  layers  in  the  tunica  muscularis, 
an  outer  longitudinal  and  an  inner  circular  one.  An  extremely  thin  muscularis 
mucosae  occurs  just  inside  the  external  tunic.  It  is  separated  from  the  outer 
musculature  by  an  extremely  sparse  complement  of  loose  submucosa.  Both 
external  muscle  layers  often  are  variably  thicker  on  the  greater  curvature  than 
on  the  lesser  curvature.  The  musculature  generally  is  thicker  in  phyllostomatines 
and  phyllonycterines  than  in  the  other  subfamilies. 

Considerable  variability  in  the  relative  thickness  of  the  two  outer  layers  has 
been  observed  within  the  stomachs  of  phyllostomatids.  In  most  species,  the 
layers  are  subequal,  with  the  circular  layer  being  the  more  robust  of  the  two. 
The  circular  layer  is  not  infrequently  organized  into  bundles,  cross-sections  of 
which  are  easily  viewed  in  longitudinal  stomach  sections.  This  “bundling”  is 
most  pronounced  in  the  caecum  (when  present)  where  it  is  prominent  in  the 
greater  curvature  in  the  majority  of  stenodermines  that  have  been  examined. 
In  a  variety  of  leaf-nosed  bats,  particularly  glossophagines  and  stenodermines, 
these  bundles  are  particularly  thick  just  beneath  the  folds  ( =  rugae)  in  the  stomach 
lining.  In  Chiroderma  villosum,  circumferential,  parallel,  external  constrictions 
occur  in  the  elongate  caecum  as  a  result  of  the  distinctively  thickened  circular 
bands  beneath  the  rugae. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


219 


The  circular  layer  clearly  is  the  dominant  portion  within  the  aboral  pyloric 
tube  of  nearly  all  species.  Macrophyllum  macrophyllum  is  a  noteworthy 
exception  because  in  this  species  the  aboral  circular  layer  is  thinner  than 
elsewhere  in  the  stomach.  In  stenodermines,  the  pyloric  circular  layer  thickens 
progressively  from  cardiac  vestibule  to  pyloric  sphincter. 

Species  that  feed  predominantly  or  exclusively  on  plant  material  have 
enlarged  cardiac  vestibules  and  fundic  caeca.  This  development  of  “sub¬ 
compartments”  is  accompanied  by  a  progressive  reduction  in  the  thickness 
of  the  muscularis  extemis  in  the  enlarged  areas.  In  species  that  apparently 
are  omnivorous  (for  example,  Glossophaga  soricina,  Phyllostomus  discolor , 
and  species  of  Micronycteris ),  the  muscularis  externis  is  reduced  in  thickness 
in  the  apex  of  the  caecum.  Such  a  reduction  could  be  regarded  as  an  intermediate 
condition  or  as  reflective  of  a  trend  toward  a  frugivorous  diet. 

Intestine 

Bats  most  often  have  short,  small  intestines  in  comparison  to  other  kinds  of 
small  mammals.  Most  comparative  measurements  of  intestinal  lengths  in 
bats  (see  Eisentraut,  1950;  Robin,  1881)  have  revealed  that  frugivores  usually 
have  relatively  long  intestines  (in  relation  to  body  length)  when  compared  to 
insectivorous,  carnivorous,  or  nectarivorous  species.  This  finding  applies  to 
Phyllostomatidae  as  well  as  to  the  Microchiroptera  in  general. 

Eisentraut  (1950)  noted  that  of  numerous  species  of  bats  with  a  variety  of 
feeding  habits,  those  with  an  intestinal  length  greater  than  four  times  the  body 
length  always  were  fruit-eating  phyllostomatids,  and  that  others  had  intestines 
of  relatively  lesser  length.  Among  species  with  the  longest  intestines  (relative 
to  body  length)  are  Chiroderma  villosum,  Vampyrops  vittatus,  and  several 
species  of  Artibeus  and  Brachyphylla  (Forman,  unpublished  data).  Vampires 
have  intestines  of  moderate  length.  Based  on  only  scattered  and  incomplete 
data,  those  few  glossophagines  for  which  measurements  are  available  generally 
have  relatively  short  intestinal  tubes. 

In  general  morphological  features,  the  intestine  differs  little  from  that  of 
most  other  groups  of  small  mammals.  Both  “small”  and  “large”  intestinal 
segments  are  present  and  a  short  duodenum  is  distinguished  by  noteworthy 
breadth.  One  noteable  feature,  shared  with  other  groups  of  bats,  is  the  lack  of 
an  ascending  or  transverse  colon  so  that  the  large  intestine  is  restricted  to  a 
relatively  short  descending  colon. 

A  caecum  always  is  lacking.  However,  at  the  junction  of  small  and  large 
intestines  there  frequently  is  a  small  ampulla  formed  as  a  result  of  a  hypertrophic 
dilation  of  the  muscularis  externa.  Abundant  lymphoid  tissue  (nodules  of 
Peyer’s  patches)  always  are  present  within  the  ampulla,  which  is  displaced 
well  away  from  mesenteric  attachment  to  the  gut  (Forman,  1974a,  19746). 
This  ampulla  first  was  observed  in  Carollia  perspicillata  (Schultz,  1965). 
Schultz  likened  this  “protrusion,”  in  size  and  location,  to  the  abbreviated 
ileocolonic  caecum  in  species  of  the  Old  World  microchiropterans  Rhinopoma 
and  Megaderma. 


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Fig.  5. — Scanning  electron  micrograph  of  intestinal  villi  in  the  middle  portion  of  the 
small  intestine  of  Phyllonycteris  aphylla.  The  arrow  indicates  a  plane  of  orientation  of 
rows  of  villi  that  is  diagonal  with  respect  to  the  intestinal  wall.  Note  the  generally  pyramidal 
shape  of  each  villous  and  that  villi  in  one  row  lie  between  villi  in  an  adjacent  row  when 
they  are  viewed  directly  from  left  to  right.  Scale  is  0.25  mm. 


Although  an  ileocaecal  valve  is  lacking  in  all  species  that  have  been  examined, 
Schultz  (1965)  noted  the  presence  of  a  valvelike  flap  within  the  middle  portion 
of  the  intestine  in  Diphylla  ecaudata.  Whether  this  structure  functions  as  a  valve 
is  not  known. 

The  complete  gastrointestinal  tracts  of  six  species  of  phyllostomatids  were 
figured  by  Schultz  (1965)  in  his  monograph  on  blood  vessel  supply  to  the  digestive 
tract  of  bats.  Several  of  his  figures  of  the  gut  of  Glossophaga  soricina  reveal  an 
extremely  complex  “looping”  of  the  intestine  in  this  species.  The  first  loop  of 
the  intestine  is  joined  to  the  terminal  portion  of  the  ileum  by  a  mesenteric 
ligament.  The  intestine  then  proceeds  into  considerable  looping,  the  extent  of 
which  is  a  function  of  intestinal  length.  The  attachment  of  the  first  intestinal 
loop  to  the  terminal  ileum  by  a  ligament  also  was  illustrated  by  Schultz  in  a 
figure  of  the  gut  of  Carollia  perspicillata. 

Torsion  is  extensive  in  the  intestine  of  most  phyllostomatids.  In  Carollia  and 
Glossophaga,  it  is  as  much  as  270°  (Schultz,  1965).  In  most  phyllostomatids, 
the  intestine  is  considerably  displaced  to  the  right  within  the  abdominal  cavity. 
One  exception  is  Macrotus  californicus,  in  which  the  intestine  is  not  displaced. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


221 


Fig.  6. — Surface  view  of  intestinal  folds  (  =  villi)  in  one  specimen  of  Artibeus 
jamaicensis.  Note  the  complex  interdigitation  and  maze-like  organization  of  ridges.  Also, 
note  the  narrow  channels  (C)  within  the  intestinal  epithelium.  These  are  most  distinctive, 
in  breadth  and  depth,  at  the  angles  or  bends  of  the  intestinal  folds.  Scale  is  0.5  mm. 


The  small  intestines  of  vampires  ( Desmodus  and  Diphylla )  are  not  grossly 
different  from  those  of  other  phyllostomatids.  However,  twisting  is  only  slight, 
and  most  of  the  intestine  is  folded  back  upon  itself  in  a  series  of  numerous  compact 
winding  folds. 

The  topography  of  the  mucous  membrane  of  the  large  intestine  generally 
is  uniform  among  the  few  species  that  have  been  examined.  Folds  are  longitudinal 
and  usually  have  smooth  surfaces  with  an  abundance  of  goblet  cells. 

Considerable  variation  in  the  topography  of  the  mucous  membrane  of  the 
small  intestine  occurs  within  and  among  species  of  phyllostomatids.  Projections 
of  the  membrane  into  the  lumen  can  be  in  the  form  of  fingerlike  villi,  nearly 
continuous  transverse  folds,  or  projections  of  a  form  to  some  degree  intermediate 
between  the  other  two  extremes.  Although  variation  is  extensive,  a  review  of 
the  literature,  along  with  some  observations  of  gut  morphology  in  phyllostomatids 
by  the  senior  author,  reveals  one  apparent  pattern  of  villous  distribution  within 
the  family.  This  pattern  occurs  most  consistently  within  fruit-eating  species. 
Fingerlike  villi,  if  present,  usually  are  located  within  the  distal-most  portion 
or  ileum.  As  one  progresses  upward  toward  the  gastroduodenal  junction, 
“pyramid-shaped”  projections,  which  are  oriented  in  transverse  rows,  become 


222 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


L 


Fig.  7. — Surface  view  of  intestinal  folds  (  =  villi)  in  one  specimen  of  Carollia 
perspicillata.  Scale  is  0.5  mm. 


L 


i 


Fig.  8. — Surface  view  of  intestinal  folds  (  =  villi)  in  one  specimen  of  Chrotopterus 
auritus.  Note  the  simplicity  of  folds  as  compared  with  those  of  Artibeus  jamaicensis  and 
Carollia  perspicillata.  Scale  is  0.5  mm. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


223 


Fig.  9. — Several  forms  of  villi  observed  within  the  small  intestine  (middle  portion) 
of  Mcicrotus  waterhousii.  Arrows  indicate  the  presence  of  a  groove  on  the  surface  of 
some  villi.  All  villi  are  drawn  to  scale. 

abundant  and  increase  in  lateral  dimension.  These  projections  are  distributed 
in  rows  that  assume  a  zig-zag  configuration  when  viewed  from  the  top.  The 
zig-zags  in  most  kinds  become  progressively  more  flattened  from  the  middle 
portion  of  the  intestine  through  the  duodenum.  Also,  the  transverse  folds  or 
“pennant-shaped  villi”  (after  Schultz,  1965),  which  interdigitate  with  and  are 
interrupted  by  one  another  within  the  lower  portions  of  their  distribution, 
often  loose  much  of  this  complexity  in  the  upper  portions  of  the  small 
intestine. 

The  most  detailed  descriptions  of  intestinal  mucous  membrane  topography 
of  phyllostomatids  are  those  of  Mathis  (1928)  and  Schultz  (1965).  Mathis 


224 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


described  the  villous  pattern  in  Phyllostomus  hastatus  and  Glossophaga 
soricina  and  reported  that  in  his  view  villi,  as  such,  were  lacking  in  portions 
of  the  intestine  in  Phyllostomus.  Also,  the  broad  villi  in  the  uppermost 
intestine  were  set  in  oblique  rows.  This  latter  pattern  also  occurs  in  other  species 
within  the  family  (see  Fig.  5).  Mathis  reported  that  the  villi  of  G.  soricina 
in  some  areas  can  be  tightly  compact  without  any  arrangement  into  rows. 
Digitate  or  club-shaped  villi  may  be  interspersed  among  “transverse  folds” 
and  be  of  somewhat  greater  height  than  the  folds.  Schultz’s  (1965)  description 
of  villous  morphology  in  G.  soricina  generally  agrees  with  that  of  Mathis 
(1928);  Schultz  further  stated  that  the  configuration  in  Anoura  geoffroyi  is 
“just  as  with  G.  soricina .”  The  extent  to  which  the  pattern  as  observed  in  these 
two  species  can  be  applied  to  other  glossophagines  is  unknown. 

Intestinal  villi  of  the  fruit-eating  stenodermines  frequently  are  arranged  in 
extremely  elaborate  interdigitating  networks  (see  Fig.  6).  This  complicated 
arrangement  likely  helps  to  impede  transport  of  food.  Other  fruit-eating 
species  have  less  elaborate  villous  arrangements  (Fig.  7).  One  carnivorous 
kind  (Fig.  8)  has  villi  uncomplicated  in  cross-sectional  configuration  and 
nearly  fingerlike  in  their  appearance. 

Villi  often  are  arched  from  side  to  side.  This  feature  in  combination  with 
staggered  arrangement  of  villi  in  oblique  rows  produces  a  mechanism  for 
entrapment  of  food  material  between  villi  at  their  bases.  This  likely  results 
in  improved  food  assimilation  or  absorption  inasmuch  as  food  would  tend 
to  be  retained  in  the  small  intestine  for  longer  periods  of  time. 

Some  variability  in  the  structure  of  villi  occurs  within  localized  portions 
of  the  small  intestine  in  phyllostomatids.  Villi  within  the  middle  portion  of  the 
intestine  of  Mac  rot  us  waterhousii  can  have  narrowly  rounded  (Fig.  9  a)  or 
relatively  broadly  rounded  apices  (Fig.  9c).  The  “arching”  of  villi,  with 
subsequent  entrapment  of  food  material,  might  be  augmented  by  an  apparent 
groove  on  the  superior  surface  of  some  villi  (see  Figs.  9c,  h).  Food  could  become 
trapped  at  the  base  of  these  folds. 

The  intestinal  topography  of  Desmodus  is  not  known  to  be  particularly 
specialized.  Villi  are  known  to  be  present  in  the  intestines  of  both  Desmodus 
and  Diphylla  (Schultz,  1965)  but  generally  are  not  fingerlike,  and  they  are 
arranged  in  a  pattern  of  interdigitation.  Rouk  and  Lane  (1970)  reported  that 
the  crypts  of  Lieberkuhn  appear  to  be  reduced  in  comparison  to  other  species. 

The  types  of  cells  present  within  the  small  intestine  of  phyllostomatids 
essentially  are  the  same  as  those  of  other  groups  of  bats  and  other  eutherians. 
The  Paneth  cells  of  bats  have  been  examined  by  Schaaf  (1970)  in  relation  to 
food  habits.  Schaaf s  study  group  included  three  insectivorous  species  as 
well  as  Artibeus  jamaicensis,  Bachyphylla  nana,  Phyllonycteris  poeyi,  and 
Monophyllus  redmani.  The  results  of  selected  histochemical  tests  were 
uniform  for  prosecretion  granules  and  mucopolysaccharides  in  all  species. 
Strong  acidophilia  was  present  in  the  cells  indicating  the  probable  presence 
of  lysosomes.  Secretion  granules  contained  a  mixture  of  protein  and  carbohydrates. 
The  results  agree  well  with  those  for  other  species  of  mammals.  Therefore, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


225 


Paneth  cells  presently  are  not  known  to  be  specialized  to  permit  the  assimilation 
of  large  quantities  of  any  particular  food  material  by  phyllostomatids,  for 
which  food  habits  are  highly  varied  but  generally  obligate. 

The  glands  of  Brunner  are  mucus  producing  and  generally  restricted  in 
distribution  to  an  extremely  narrow  submucosal  ring  at  the  gastroduodenal 
junction.  Several  unusual  conditions  with  respect  to  Brunner’s  glands  occur 
within  the  Phyllostomatidae.  These  conditions  might  relate  to  the  varied 
food  habits  that  occur  within  the  family. 

The  stomachs  of  Sturnira  lilium  and  S.  ludovici  have  cells  within  the  bases 
of  the  pyloric  glands  that  are  histologically  identical  to  the  submucosal  glands 
of  Brunner  within  the  uppermost  duodenum.  Several  species  of  Artibeus 
(Forman,  1972;  Rouk,  1973)  have  similar  cells  within  their  pyloric  stomachs. 
Cells  of  Brunner’s  glands  in  the  duodenum  and  those  cells  at  the  base  of 
pyloric  glands  stain  identically  with  the  periodic  acid  Schiff  reaction  for  neutral 
mucopolysaccharides.  This  staining  is  considerably  different  from  that 
within  remaining  cells  of  the  pyloric  glands.  Cells  such  as  those  of  Brunner’s 
glands  may  provide  for  better  protection  of  the  pyloric  mucosa  from  large 
amounts  of  hydochloric  acid  that  likely  are  produced  by  the  considerable 
number  of  parietal  cells  in  some  fruit-eating  phyllostomatids. 

Of  those  studied,  the  Brunner’s  glands  of  Phyllostomus  hastatus  and  P. 
discolor  are  best  developed.  Other  species  of  phyllostomatines  (those  of 
Tonatia,  Micronycteris ,  and  Chrotopterus )  have  relatively  numerous  Brunner’s 
glands  but  they  nevertheless  are  less  distinctive  than  are  those  of  Phyllostomus. 

The  numerous  species  of  stenodermines,  carolliines,  and  some  species  of 
glossophagines  are  in  marked  contrast  to  the  phyllostomatines.  Although 
only  a  few  species  of  Artibeus  have  been  examined,  it  is  known  that  the 
Brunner’s  glands  of  A.  lituratus  and  A.  jamaicensis  are  extremely  sparse  in  the 
most  proximal  portion  of  the  duodenum  and  that  they  are  absent  in  at  least 
some  specimens  of  Artibeus  phaeotis  and  in  A.  inopinatus.  It  is  reasonable  to 
hypothesize  that  other  species  of  Artibeus  harbor  few  of  these  glands.  In  addition 
to  species  of  Artibeus,  the  following  bats  have  been  reported  to  lack  Brunner’s 
glands  at  the  gastroduodenal  junction:  Centurio  senex,  Chiroderma  villosum, 
Uroderma  bilobatum,  Vampyrodes  caraccioli,  Vampyressa  pusilla,  V.  nymphaea, 
and  V ampyrops  helleri.  Artibeus  toltecus  and  Vampyrops  vittatus  are  reported  to 
have  numerous  Brunner’s  glands  at  the  gastrointestinal  junction.  The  basal  cells 
of  the  pyloric  glands  in  Centurio  senex  are  histologically  similar  to  the  Brunner’s 
glands  of  Artibeus  lituratus.  Also,  it  is  noteworthy  that  all  species  of  stenodermines 
that  lack  Brunner’s  glands  in  the  upper  duodenum,  except  for  Chiroderma,  have 
relatively  extensive  zones  of  pyloric  mucosa  in  the  stomach.  It  is  reasonable  at  this 
point  to  suggest  that  the  pyloric  mucosa  in  these  animals  may  be  performing  the 
“neutralization”  action  on  the  food  bolus  that  ordinarily  is  believed  to  be  performed 
by  the  glands  of  Brunner  in  other  species  of  mammals. 

Additionally,  several  species  of  nectar-feeding  glossophagines  ( Lichonycteris 
obscura  and  Choeroniscus  godmani )  have  been  observed  to  have  few  Brunner’s 
glands  (Forman,  1971a).  The  only  phyllonycterine  that  has  been  examined, 


226 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Brachyphylla  cavernarum,  has  no  glands  of  Brunner.  These  observations, 
along  with  those  on  stenodermines,  clearly  indicate  that  the  conditions  in 
certain  phyllostomatids  do  not  support  the  widely  held  view  that  mammals 
consuming  plant  material  have  more  abundant  glands  of  Brunner  than  do 
animals  eating  animal  material. 

The  connective  tissue  of  the  intestine  of  bats  generally  is  extremely 
sparse.  The  intestine  of  Desmodus  rotundus  (and  perhaps  the  other  two 
sanguivorous  species)  is  a  noteworthy  exception.  Both  the  submucosa  and  the 
lamina  propria  of  the  villi  are  unusually  thick  and  dense.  They  are  highly 
vascularized  and  harbor  a  considerable  lymphatic  network. 

Studies  of  organized  gut-associated  lymphoid  tissue  (Peyer’s  patches)  in 
New  World  bats  (Forman,  1974a,  19746)  have  revealed  differences  in 
abundance,  distribution,  and  morphology  of  this  tissue  within  the 
Phyllostomatidae.  These  differences  possibly  relate  to  diet.  For  example,  fruit¬ 
eating  species  usually  have  the  most  patches  when  compared  with  nectarivorous 
or  with  carnivorous  and  insectivorous  kinds.  Also,  the  patches  can  occur 
almost  anywhere  along  the  length  of  the  small  intestine  in  fruit  eaters,  frequently 
including  the  duodenum.  These  patches  have  relatively  large  nodules  with 
extremely  large  geminal  centers.  The  patches  and  nodules  of  insect  eaters 
and  carnivores,  in  contrast,  are  relatively  small  with  small  germinal  centers 
typically  indicating  a  low  state  of  activity.  Patches  in  these  species  usually  are 
restricted  to  the  submucosa  of  the  ileum. 

These  observations  suggest  that  at  least  within  the  family  Phyllostomatidae 
organized  lymphoid  tissue  within  the  gut  might  be  differentially  responsive  to 
intestinal  contents  including  food  material  and  associated  microbial  populations 
as  well. 


Literature  Cited 

Allison,  A.  C.  1948.  The  stomach  in  South  African  Insectivora,  with  notes  on  the 
organization  of  mammalian  gastric  glands.  J.  Anat.,  82:249-261. 

Eisentraut,  M.  1950.  Die  Ernahrung  de  Fledermausen  (Microchiroptera).  Zool. 
Jahrb.,  Jena.,  79: 1 14-177. 

Forman,  G.  L.  1971a.  Gastric  morphology  in  selected  mormoopid  and  glossophagine 
bats  as  related  to  systematic  problems.  Trans.  Illinois  Acad.  Sci.,  64:273-282. 

- .  19716.  Histochemical  differences  in  gastric  mucus  of  bats.  J.  Mamm., 

52:191-193. 

- .  1972.  Comparative  morphological  and  histochemical  studies  of  stomachs 

of  selected  American  bats.  Univ.  Kansas  Sci.  Bull.,  49:591-729. 

- .  1973.  Studies  of  gastric  morphology  in  North  American  Chiroptera 

(Emballonuridae,  Noctilionidae,  and  Phyllostomatidae).  J.  Mamm.,  54:909-923. 

- .  1974a.  Comparative  studies  of  organized  gut-associated  lymphoid  tissue  in 

mammals  with  diverse  food  habits.  Distribution,  size,  and  organization  of 
Peyer’s  patches  in  New  World  bats.  Trans.  Illinois  Acad.  Sci.,  67:152-156. 

- .  19746.  The  structure  of  Peyer's  patches  and  their  associated  nodules  in  New 

World  bats  in  relation  to  food  habits.  J.  Mamm.,  55:738-746. 

Hart,  L.  A.  1971.  Structure  of  the  gastric  mucosa  as  related  to  feeding  habits  in 
selected  species  of  New  World  bats.  Unpublished  Ph.D.  dissertation, 
Virginia  Polytechnic  Inst.,  65  pp. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


227 


Gardner,  A.  L.  1977.  Feeding  habits.  Pp.  293-350,  in  Biology  of  bats  of  the  New 
World  family  Phyllostomatidae.  Part  II  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C. 
Carter,  eds.).  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  13:1-364. 

Huxley,  T.  H.  1865.  On  the  structure  of  the  stomach  in  Desmodus  rufus.  Proc. 

Zool.  Soc.,  London,  pp.  386-390. 

Kolb,  A.  1954.  Biological  Beobachtungen  an  Fledermausen.  Saugetiekundl.  Mitt., 
2:15-26. 

Lillie,  R.  D.  1965.  Histopathologic  technic  and  practical  histochemistry.  McGraw- 
Hill,  New  York,  xii  +  775  pp. 

Mathis,  J.  1928.  Beitrag  zur  Kenntnis  des  Fledermausdarmes.  Z.  Mik.  Anat. 

Forsch.,  12:594-647. 

Moller,  W.  1932.  Das  Epithel  der  Speiserohrenschleimhaut  der  blutenbesuchenden 

Fledermaus  Glossophaga  soricina  im  Vergleich  zu  insektenfressenden 
Chiropteren.  Zeit.  Mikr.  Anat.,  29:637-653. 

Myrcha,  A.  1967.  Comparative  studies  on  the  morphology  of  the  stomach  in  the 

Insectivora.  Acta  Theriol.  12:223-244. 

Phillips,  C.  J.,  G.  W.  Grimes,  and  G.  L.  Forman.  1977.  Oral  biology.  Pp.  121-246, 
in  Biology  of  bats  of  the  New  World  family  Phyllostomatidae.  Part  II  (R.  J. 
Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.).  Spec.  Publ.  Mus.,  Texas  Tech 
Univ.,  13:1-364. 

Robin,  H.  A.  1881.  Recherches  anatomique  sur  les  mammiferes  de  l’ordre  des  Chi- 
ropteres.  Ann.  Sci.  Nat.  Zool.,  12:1-180. 

Rouk,  C.  S.  1968.  Comparative  gastric  histology  of  selected  American  bats.  Unpublished 
M.S.  thesis,  Oklahoma  State  Univ.,  51  pp. 

- .  1973.  Gastric  morphology  and  adaptive  radiation  in  the  Phyllostomatidae. 

Unpublished  Ph.D.  dissertation,  Texas  Tech  University,  85  pp. 

Rouk,  C.  S.,  and  B.  P.  Glass.  1970.  Comparative  gastric  histology  of  American  bats. 
J.  Mamm.,  51:455-472. 

Rouk,  C.  S„  and  W.  L.  Lane.  1970.  Comparative  histology  of  intestines  of  selected 
American  bats.  Abstracts  of  Papers  Presented  at  the  50th  Annual  Meeting, 
American  Society  of  Mammalogists,  College  Station,  Texas,  June. 

Schaaf,  V.  P.  1970.  Untersuchungen  uber  das  histochemische  Verhalten  der  Panethschen 
Kornerzellen  bei  mittelamerikanischan  Fledermausarten  mit  unterschiedlichen 
Ernahrungsweisen.  Anat.  Anz.  Bd.,  126:275-277. 

Schultz,  W.  1965.  Studien  uber  den  Magen-Darm-Kanal  der  Chiropteren.  Ein 
Beitrag  zum  Problem  der  Homolgisierung  von  abschnitten  des  Saugetierdams.  Ziet. 
Wissenschaft.  Zool.,  171:241-391. 


MORPHOMETRIC  ANALYSIS  OF 
CHIROPTERAN  WINGS 


James  Dale  Smith  and  Andrew  Starrett 


Bats  are  unique  among  mammals  in  their  possession  of  wings.  The  evolution 
and  adaptation  of  these  anatomically  complex  structures  along  with  the  develop¬ 
ment  of  an  acute  ability  to  orient  acoustically  has  contributed  markedly  to  one 
of  the  most  interesting  examples  of  adaptive  radiation  in  vertebrate  history.  Yet 
the  morphometric  properties  of  bat  wings  have  remained  poorly  understood. 
Biologists  have  described  chiropteran  diversity  and  faunal  complexity  throughout 
the  world,  but  the  flight  behavior  of  only  a  few  species  has  been  reported  (see 
Eisentraut,  1936;  Dwyer,  1965;  Kulzer,  1968;  Norberg,  1970,  1976a,  1976b; 
Pennycuick,  1 97 1 ;  Schnitzler,  1971). 

Revilliod  (1916)  was  the  first  to  attempt  to  describe  the  morphometries  of 
chiropteran  wings.  In  this  much  overlooked  paper,  he  utilized  several  indices 
to  demonstrate  the  degree  of  adaptation  to  flight  by  several  families  of  bats.  Poole 
(1936)  was  among  the  earliest  investigators  to  report  wing  loading  values  for  bats, 
and  Struhsaker  (1961)  was  the  first  to  calculate  aspect  ratios  of  bat  wings.  Bader 
and  Hall  (1960)  were  the  first  investigators  to  use  computer  techniques  to  analyze 
the  osteometric  variation  of  bat  wings.  In  this  study,  they  employed  correlation 
coefficients  to  assess  the  interrelationships  among  the  skeletal  elements  of  the  wing 
and  foot  of  Myotis  lucifugus  and  M.  sodalis. 

Other  studies,  although  important  contributions,  have  been  limited  in  their  scope 
and  coverage.  Among  these  are  Vaughan’s  (1959)  detailed  anatomical  analysis 
of  three  bat  species  from  North  America;  a  more  recent  survey  of  the  skeletal 
and  muscular  system  and  aerodynamics  appears  in  Vaughan  (1970a,  1970b, 
1970c).  Hartman  (1963),  Gaisler  (1964),  Farney  and  Fleharty  (1969),  and 
Jones  and  Suttkus  (1971)  have  reported  wing  loading  and  aspect  ratios  for  num¬ 
erous  species  of  bats.  Pearson  et  al.  (1952),  Orr  (1954),  Short  (1961),  and  Jones 
(1967)  have  contributed  important  information  relative  to  the  growth  and  de¬ 
velopment  of  chiropteran  wings.  Seasonal  changes  in  wing  loading  of  several 
North  American  species  were  examined  by  Davis  (1969)  and  O’Farrell  and 
Studier  (1976),  and  Norbert  (1969,  1972)  reported  on  functional  osteology  and 
myology  of  the  wings  of  several  bats. 

By  far,  the  most  extensive  analysis  of  the  morphometric  properties  of  bat  wings 
is  that  by  Findley  et  al.  (1972).  In  this  study,  they  relied  on  regression  and 
correlation  procedures  as  well  as  factor  analysis  to  examine  the  wings  of 
approximately  1 35  species.  Our  initial  goal  was  to  expand  on  this  study  with  our 
primary  focus  on  the  bats  of  the  family  Phyllostomatidae.  However,  it  soon  became 
apparent  to  us  that  a  meaningful  interpretation  of  the  morphometries  of  phyllosto- 
matid  wings  required  a  broader  understanding  of  the  overall  variation  in  size 
and  shape  of  wings  in  the  Chiroptera. 


229 


230 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Methods 

Methods  of  deriving  the  form  and  extent  of  chiropteran  wings  for  the  purpose 
of  studying  size  and  shape  have  been  variable.  For  example,  some  workers  have 
traced  the  outline  of  the  extended  wing  of  freshly  killed  bats  or  individuals  pre¬ 
served  in  alcohol.  From  such  tracings,  they  have  derived  the  area  of  the  wing  and 
other  aerodynamic  parameters  by  using  a  planimeter  or  by  some  other  time- 
consuming  procedure.  While  these  efforts  are  to  be  commended,  such  tech¬ 
niques  do  not  readily  permit  an  overall  consideration  of  the  diversity  of  the 
chiropteran  fauna  of  the  world. 

In  addition,  most  past  studies  of  wing  morphology  have  neglected  to  consider 
the  influence  of  the  fourth  digit  in  determining  the  size  and  shape  of  the  wing. 
Typically,  the  lengths  of  the  forearm  and  digit  III  are  taken  to  describe  the  span 
of  the  wing,  and  the  length  of  digit  V,  its  width.  These  measurements  have  been 
used  to  derive  the  aspect  ratio  and  wing  loading  of  chiropteran  wings,  which, 
characterized  in  this  manner,  are  assumed  to  be  rectangular  in  shape.  For  deter¬ 
mination  of  wing  loading,  such  calculations  tend  to  result  in  over-estimates  of 
area  due  to  the  inclusion  of  an  intrinsic  portion  of  the  rectangular  shape  that, 
in  fact,  does  not  exist  in  the  real  wing  (Fig.  1).  These  calculations  also  may  lead 
to  mistaken  estimates  of  similarity  between  markedly  different  wings  and  may 
mask  subtle  differences  between  similarly  shaped  wings.  Furthermore,  most 
past  studies  have  considered  only  the  total  lengths  of  digits  rather  than  examining 
the  variability  of  digital  composition  and  its  influence  on  wing  size  and  shape. 

In  this  study,  1 1  wing  measurements,  length  of  the  head  and  body,  and  weight, 
were  obtained  from  1456  museum  specimens,  which  comprised  433  species  and 
147  genera  from  17  families  of  bats.  Most  of  these  specimens  were  conventional 
study  skins,  although  in  some  cases  only  specimens  preserved  in  alcohol  were 
available.  The  wing  measurements  included  the  length  of  the  forearm  (as  described 
by  Smith,  1972)  and  the  individual  lengths  of  the  metacarpal  and  phalangeal 
elements  of  digits  III,  IV,  and  V.  The  length  of  the  often  curved  and  cartilaginous 
portion  of  the  terminal  phalange  of  the  third  digit  was  recorded  as  the  greatest 
radius  of  the  arc.  When  available,  the  length  of  the  head  and  body  and  the  weight 
of  the  specimens  were  recorded  from  the  specimen  label.  Head  and  body  length 
was  measured  directly  on  specimens  preserved  in  alcohol.  The  weights  of  many 
specimens,  especially  those  in  alcohol,  were  not  recorded  at  the  time  of  capture. 
In  these  cases,  weights  were  estimated  (see  below).  All  measurements  were  re¬ 
corded  in  millimeters  (by  means  of  dial  calipers,  calibrated  in  twentieths  of  a 
millimeter)  or  grams. 


Derived  Variables 

At  the  outset  of  our  analysis,  we,  like  many  others  before  us,  converted  our  raw 
variables,  a  priori ,  into  a  number  of  derived  variables  such  as  aspect  ratio, 
wing  loading,  tip  index,  and  so  forth.  The  subsequent  analysis  of  these  derived 
variables  was  beset  with  a  number  of  problems.  Foremost  among  these  were 
inflated  correlations,  which  resulted  from  linear  dependence  of  the  derived 
variables.  This  resulted  in  obscuring  the  sources  of  dependency.  Atchley  (1978), 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


231 


Fig.  1. — Diagrammatic  comparison  of  an  actual  wing  and  the  construct  of  the  wing 
(stippled  area)  used  in  this  study.  The  dotted  line  indicates  the  assumed  shape  of  the  wing 
if  only  the  length  (forearm  plus  digit  III)  and  width  (digit  V)  are  considered. 


Atchley  and  Anderson  (1978),  Atchley  et  al.  (1976),  and  Pimentel  (1978)  re¬ 
cently  presented  discussions  regarding  the  statistical  properties  of  derived  vari¬ 
ables  such  as  ratios  and  indices.  Although  derived  variables  can  be  useful  in  some 
cases,  they  should  be  scrutinized  closely  and  avoided  when  possible.  Because  the 
goal  of  our  investigation  was  to  examine,  insofar  as  possible,  the  interactions 
among  wing  components  and  because  these  interactions  were  largely  masked  by 
the  difficulties  noted  above,  we  chose  to  analyze  only  our  original  raw  variables. 
However,  after  these  analyses  were  completed  ( a  posteriori),  we  found  that  some 
of  our  derived  variables  could  be  used  in  a  generalized  descriptive  sense.  Those 
which  were  found  to  be  most  useful  are  presented  in  the  Appendix  (Tables  Al- 
A21)  and  are  described  below. 

Weight. — This  variable  was  essential  to  the  computation  of  wing  loading.  To 
circumvent  the  problem  of  missing  data,  Findley  et  u/.(1972,  table  3)  utilized 
the  predicting  qualities  of  a  simple  linear  regression  to  derive  estimated  weight 
from  head  and  body  length.  We  also  examined  this  relationship  for  1082  specimens 
using  a  similar  regression  model  on  known  head  and  body  length  (20  and  weight 
(  Y)  and  found  that  the  residuals  (  Y-  Y')  were  lowest  at  the  small-sized  end  of  the 
variation.  However,  the  residuals  increased  markedly  at  the  large-sized  end  of 
the  spectrum.  In  an  attempt  to  reduce  these  overestimates,  we  computed  a  second 
degree  polynomial  regression.  This  reduced  the  magnitude  of  the  residuals  in 
the  upper  range  of  variation,  but  the  analysis  did  not  provide,  in  our  opinion, 
totally  satisfactory  results.  As  did  Findley  et  al.  (1972),  we  partitioned  our  data 
into  recognized  taxonomic  groups  corresponding  to  familial  and  subfamilial 
categories  and  obtained  different  functions  for  nearly  every  grouping  (Table  1). 


232 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1 . — Results  of  the  second  degree  polynomial  regression  analysis  of  head  and  body  length 
(X-axis)  and  weight  (Y-axis).  Symbols  are:  correlation  coefficient,  r;  Y-intercept,  A;  linear 
regression  coefficient,  Bl;  and  quadratic  regression  coefficient,  B2.  Significant  correlation 
coefficients  and  F -values  are  indicated  with  an  asterisk. 


Taxon 

N 

r 

A 

Bl 

B2 

F 

Pteropodidae 

119 

0.967* 

26.53 

-0.892  +  0.256 

0.011+0.001 

928.24* 

Emballonuridae 

36 

0.877* 

26.89 

-1.104  +  0.720 

0.014  +  0.005 

60.35* 

Rhinolophidae 

153 

0.978* 

2.37 

-0.375  +  0.162 

0.010  +  0.001 

1680.94* 

Nycteridae 

34 

0.651* 

-45.59 

1.789+1.508 

-0.013  +  0.014 

12.66 

Megadermatidae 

18 

0.573* 

-239.29 

6.235  +  2.784 

-0.035  +  0.017 

7.46 

Noctilionidae 

26 

0.925* 

-218.56 

5.229+1.340 

-0.023  +  0.008 

121.54* 

Phyllostomatinae 

168 

0.943* 

-12.33 

0.024  +  0.184 

0.007  +  0.001 

659.23* 

Glossophaginae 

85 

0.865* 

24.61 

-0.813  +  0.297 

0.010  +  0.002 

136.63* 

Carolliinae 

12 

0.967* 

20.76 

-0.830  +  0.658 

0.012  +  0.006 

76.95* 

Stenoderminae 

128 

0.955* 

26.59 

-1.196  +  0.476 

0.017  +  0.003 

678.55* 

Desmodontinae 

42 

0.833* 

-10.85 

-0.123  +  1.898 

0.008  +  0.012 

44.91* 

Phyllostomatidae' 

391 

0.912* 

-12.78 

0.017  +  0.130 

0.007  +  0.008 

1212.65* 

Vespertilionidae 

157 

0.933* 

0.45 

-0.116  +  0.120 

0.005  +  0.001 

524.80* 

Molossidae 

120 

0.979* 

1.54 

-0.227+0.092 

0.006  +  0.001 

1442.02* 

All  bats 

1108 

0.961* 

6.52 

-0.422  +  0.045 

0.009  +  0.001 

1990.05* 

'Combined  sample  of  the  family  Phyllostomatidae. 


The  results  of  our  linear  regression  model  (not  shown)  agreed,  for  the  most 
part,  with  those  presented  by  Findley  et  al.  (1972).  We  found  in  our  regression 
analyses  that  the  regression  coefficients  (Bl  or  B2)  had  relatively  little  effect 
on  the  slope  of  the  line.  More  importantly,  the  F-intercept  values  ( A )  varied 
greatly,  in  both  our  analysis  and  theirs,  and  in  the  majority  of  cases  these  intercept 
values  departed,  negatively,  from  zero  (the  theoretical  intercept  in  these  analyses). 
Therefore,  these  models  predicted  extremely  low  or  even  negative  weights  for 
bats  of  extremely  small  body  size.  In  those  cases  where  the  departure  of  the  F- 
intercept  was  positive,  weight  would  be  given  to  a  bat  that  had  zero  head  and 
body  length.  An  a  priori  manipulation  of  the  regression  model  certainly  might 
improve  the  “fit”  of  the  line,  but  we  suspect  biologic  reality  is  quickly  obscured 
by  such  practice;  biological  meaning  is  not  automatically  ascribed  by  statistical 
significance.  Furthermore,  we  suspect  that  the  complexity  of  the  relationships 
of  weight  to  head  and  body  length  and  other  meristic  parameters  is  more  com¬ 
plicated  than  can  be  measured  precisely  with  regression/correlation  statistics, 
and  we  strongly  caution  other  investigators  against  placing  much  faith  in  such 
predictions.  With  an  awareness  of  these  difficulties  in  mind,  we  utilized  the 
predictions  of  weights  generated  by  our  polynomial  regression  model.  However, 
the  weight  values  obtained  in  this  manner  were  used  only  to  compute  wing  loading 
for  comparative  purposes  and  these  were  not  used  in  any  further  rigorous 
analyses.  In  those  groups  where  there  were  insufficient  numbers  to  compute  a 
regression  function,  we  utilized  the  function  of  the  most  closely  related  group 
for  which  there  was  a  function.  All  weights  (actual  or  estimated)  were  converted 
to  Newtons  (Nt). 

Wing  areas. — The  computation  of  the  area  of  the  wings  was  necessary  for  the 
calculations  of  both  aspect  ratio  and  wing  loading.  The  area  of  the  plagiopatagium 
was  calculated  as  the  area  of  a  rectangle  (length  of  forearm  X  length  of  digit  V). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


233 


In  deriving  the  area  of  the  wing  tip,  we  attempted  to  consider  an  attenuated 
(polygonal)  tip  rather  than  a  simplistic,  rectangular  tip  as  has  been  the  practice. 
To  accomplish  this,  using  measurements  from  museum  material,  we  considered 
a  construct  of  the  wing  (Fig.  1 )  in  which  the  fourth  digit  was  an  integral  component. 
We  noted  from  empirical  observations  that  the  posture  of  this  digit  varied  among 
species  and  that  estimates  of  the  tip  area  varied  with  this  posture.  In  addition, 
we  found  that  in  most  instances,  when  the  wing  was  fully  extended,  the  fifth  digit 
projected  at  approximately  a  right  angle  from  the  leading  edge  (forearm  and 
digit  III).  Although  our  testing  of  empirical  data  was  limited,  we  found  that  we 
could  geometrically  estimate  the  angle  of  projection  of  digit  IV  (alpha  angle),  with 
90  per  cent  confidence,  when  the  panel  areas  A1  and  A2  (Fig.  2)  were  considered 
to  be  equal  or  nearly  equal.  More  precisely,  the  alpha  angle  equals  the  arc 
tangent  of  (length  of  digit  V/length  of  digit  III).  Alpha  angles  are  given  in 
degrees  of  rotation  from  digit  III.  Bats  with  relatively  long  fifth  digits  tended  to 
possess  large  alpha  angles,  whereas  those  with  relatively  long  third  digits  had 
lower  alpha  angles  (Table  Al). 

Once  the  alpha  angle  was  determined,  calculating  the  area  of  the  two  triangles 
Ai  and  A2  (Fig.  2)  was  simply:  area  of  the  wing  panel  between  digits  III  and  IV 
equals  (cosine  alpha  angle  X  (length  of  digit  III  X  digit  IV)  and  area  of  the 
wing  panel  between  digit  IV  and  V  equals  sine  alpha  angle  X  (length  of  digit 
IV  X  digit  V).  The  total  area  of  the  wing,  or  any  portion  thereof,  was  derived  by 
summing  the  respective  areas  and  multiplying  by  2.  All  areas  were  converted  into 
square  meters  (m2). 

Wing  loading. — This  variable  was  obtained  by  weight  (Nt)/total  area  of  the 
wing  (m2).  Wing  loads  are  reported  as  Newtons  per  square  meter  (Nt/m2)  (Table 
A6). 

Aspect  ratio. — We  followed  Hartman  (1963)  in  computing  this  variable:  over¬ 
all  aspect  ratio — 2  (length  of  forearm  plus  length  of  digit  III)2/total  area  of  the 
wing.  We  partitioned  the  aspect  ratio  into  two  additional  ratios  as  follows:  1) 
aspect  ratio  of  the  plagiopatagium — (length  of  the  forearm  X  2)2/area  of 
the  plagiopatagium,  and  2)  aspect  ratio  of  the  wing  tip — (length  of  digit  III  X 
2)2/area  of  the  wing  tip.  These  ratios  are  presented  in  Tables  A3-A5. 

Tip  index. — The  tip  of  the  chiropatagium  is  the  principal  propulsive  portion 
of  the  chiropteran  wing  (Vaughan,  1970c).  The  tip  index  (Findley  et  al.,  1972) 
is  the  ratio  of  length  of  digit  Ill/length  of  forearm.  A  high  tip  index  (2.00)  indicates 
a  proportionately  long  third  digit,  whereas  a  low  index  (1.00)  reflects  a  relatively 
short  wing  tip  (Table  A2). 

Relative  lengths  of  the  wing  elements. — We  followed  Findley  et  al.  (1972)  in 
computing  the  relative  length  of  the  wing,  which  is  (length  of  forearm  plus  length 
of  digit  III)Aength  of  the  head  and  body.  In  similar  fashion,  we  computed  the 
relative  lengths  of  the  forearm  and  digits  III-V  (Tables  A7-A1 1). 

Percentage  of  digital  composition. — In  an  a  priori  effort  to  characterize  the 
varying  composition  of  digits  III-V,  we  computed  the  percentage  that  each  digital 
element  contributed  to  the  total  length  of  its  respective  digit.  These  values  proved 
a  posteriori  to  be  useful  guidelines  in  the  interpretation  of  the  discriminant 
analysis  (Tables  A 1 2-A2 1 ). 


234 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


a 


V 

Fig.  2. — Diagrammatic  representation  of  the  derivation  of  the  alpha  angle.  See  text 
(methods)  for  discussion. 


Statistical  Procedure 

Past  studies  of  the  morphometries  of  chiropteran  wings  have  been  rather 
limited  in  the  sophistication  of  their  statistical  analysis.  Most  report  only  simple 
descriptive  statistics  such  as  mean,  range,  standard  deviation,  and  in  some  cases, 
coefficient  of  variation.  As  noted  earlier,  Bader  and  Hall  (1960)  and  Findley 
et  al.  (1972)  have  applied  more  detailed  statistical  procedures;  the  latter  employed 
both  correlation  and  regression  statistics  as  well  as  factor  analysis. 

In  our  initial  statistical  analysis  of  the  morphometric  properties  of  bat  wings, 
we  computed  such  simple  statistics  as  mean,  range,  one  standard  error  of  the  mean, 
and  coefficient  of  variation  for  all  variables.  As  noted  above,  these  descriptive 
statistics  for  selected  derived  variables  are  presented  in  the  Appendix  (Tables 
A1-A21).  In  these  tables,  taxonomic  groups  are  ranked  by  the  magnitude  of 
their  variable  means  (largest  to  smallest)  rather  than  in  phylogenetic  order. 
Within  the  family  Phyllostomatidae,  subfamilies  were  allowed  to  rank  in  this 
fashion  as  were  genera  within  subfamilies.  The  mean  for  “all  bats”  also  was  al¬ 
lowed  to  take  its  appropriate  position  within  the  familial  ranking. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


235 


We  used  regression  and  correlation  analyses  from  BIOMED  (Dixon,  1973) 
and  SPSS  (Nie  et  al.,  1975)  in  our  examination  of  the  relationships  between  head 
and  body  length  and  weight.  However,  in  the  main  portion  of  our  study,  we 
employed  the  multivariate  procedures  of  principal  components  (PCA)  and 
discriminant  analyses  to  assess  the  morphometric  interactions  among  the  twelve 
original  variables  and  their  effects  on  size  and  shape  of  chiropteran  wings. 
Descriptions  of  these  multivariate  procedures  may  be  found  in  Koons  (1962), 
Cooley  and  Lohnes  (1971),  and  Pimentel  (1978).  The  computations  of  these 
procedures  were  accomplished  in  the  Computer  Center,  California  Polytechnic 
University,  San  Luis  Obispo,  using  an  unpublished  program  (DISANAL) 
written  by  Richard  A.  Pimentel. 

Interpretation  of  the  component  graphs  and  variable  vectors. — We  suspect 
that  many  readers  might  not  be  completely  familiar  with  the  graphical  repre¬ 
sentations  that  we  have  employed  in  this  study.  It  is  difficult  to  portray  visually 
the  multidimensional  patterns  of  variation  computed  by  the  multivariate  statistical 
procedures  used  in  this  study,  which  assess  variation  among  all  ^-variables  in 
p-dimensional  space.  We  have  used  component  graphs  that  are  two  dimensional 
views  of  portions  of  these  multidimensional  spaces.  In  the  figures  beyond,  we 
have  plotted  the  first  and  second  (1X2)  axes  to  show  the  length  /width  character 
of  the  dispersion.  Height  of  the  dispersion  is  shown  in  the  graphs  in  which  axes 
one  and  three  (1x3)  are  plotted.  Viewed  together,  each  set  of  component  graphs 
depicts  the  dispersion  of  centroids  in  three  dimensions.  The  coordinates  used 
to  plot  these  graphs  (Figs.  3,  5,  6)  are  given  in  Tables  3  and  5,  respectively. 

In  Fig.  4,  we  have  plotted  the  direction  cosines  (PCA)  and  canonical  vectors 
for  the  twelve  original  variables  in  much  the  same  manner  as  described  for  the 
component  graphs.  The  coordinates  used  to  plot  these  vectors  are  given  in  Tables 
2  and  4,  respectively.  To  avoid  confusion,  only  the  positive  end  of  each  vector 
is  shown.  The  tail  or  negative  end  of  a  vector  passes  through  the  ordinate  of  each 
graph  for  an  equal  length  in  the  opposite  direction.  The  influence  that  any  one 
vector  has  on  the  location  of  the  group  centroids  is  determined  by  the  magnitude 
or  length  of  that  vector  and  the  proximity  of  its  point  (positive  end)  or  tail  (negative 
end)  to  the  various  centroids.  Long  vectors  exert  a  strong  influence  on  the  location, 
whereas  shorter  vectors  exhibit  weaker  effects.  In  these  analyses,  an  association 
with  the  positive  end  of  a  vector  implies  large  size  (longness)  and  proximity  to 
the  tail  of  a  vector  indicates  small  size  (shortness). 

It  is  important  to  bear  in  mind  continually  the  fact  that  the  overall  ordination 
of  groups  (Figs.  3,  5,  6)  is  the  result  of  synergistic  interplay  among  variables  (Fig.  4) 
and  not  the  result  of  any  one  or  two  of  these.  We  have  attempted  to  illustrate  and 
set  these  figures  in  such  a  way  as  to  facilitate  the  reader’s  perception  of  the 
dimensionality  of  the  variation  on  the  dispersion  of  groups.  To  facilitate  further 
an  interpretation  of  the  component  graphs,  the  reader  may  wish  to  make  a  xerox 
transparency  of  Fig.  4  and  overlay  this  on  the  corresponding  component  graphs. 
In  addition,  this  overlay  may  be  used  to  interpret  Figs.  11  to  16. 


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For  those  readers  who  wish  to  see  the  finer  aspects  of  the  ordination,  we  strongly 
encourage  the  construction  of  three-dimensional  models.  This  may  be  ac¬ 
complished  easily  by  xeroxing  the  1X2  component  graphs  and  attaching  these 
to  a  styrofoam  base.  Sticks  may  be  cut  to  an  appropriate  length  by  using  the  1  X  3 
component  graphs  to  determine  the  height  of  particular  centroids.  Leave  a 
sufficient  excess  on  these  sticks  to  allow  placement  in  the  styrofoam  base  at  their 
respective  (1X2)  centroid  positions.  A  three-dimensional  model  of  the  variable 
vectors  may  be  constructed  by  pushing  wires  through  a  styrofoam  ball  and 
using  Fig.  4  for  proper  orientation. 

Specimens  Examined 

In  the  following  list  of  specimens  examined,  a  total  of  1456,  bold-faced  letters 
preceding  the  familial  or  subfamilial  name  will  be  used  to  identify  the  respective 
group  centroids  in  the  component  graphs  (Figs.  5,  6,  10).  Within  the  Phyllosto- 
matidae,  bold-faced  numbers  indicate  the  species  identity  in  the  component 
graphs  (Figs.  11-16).  Any  variations  from  this  scheme  will  be  noted  in  the 
respective  legends.  Numbers  following  scientific  names  indicate  sample  size. 
The  mnemonic  acronyms  (for  example,  PTEROP  for  Pteropodidae  and 
PHYNYC  for  Phyllonycterinae)  used  in  the  figures  are  sufficiently  phonetic  to 
provide  easy  interpretation. 

A.  Pteropodidae  (172);  Aethalops  alecto,  5;  Chironax  melanocephalus,  4;  Cynopterus 
archipelagus,  4;  C.  brachyotis ,  5;  C.  sphinx ,  4;  Dobsonia  inermis,  2;  D.  minor,  2;  D. 
moluccensis,  1;  D.  praedatrix,  1;  Dyacopterus  spadiceus,  3;  Eidolon  helvum,  2;  Epomophorus 
labiatus,  1;  E.  minor,  4;  E.  wahlbergi,  1;  Balionycteris  maculata,  5;  Epomops  dobsoni, 
1;  Haplonycterisfisheri,  3;  Hypsignathus  monstrosus,  1;  Megaerops  ecaudatus,  3;  M.  wetmorei, 
3;  Micropteropus  pusillus,  3;  Myonycteris  torquatu,  2;  Nanonycteris  veldkampi,  1; 
Penthetor  lucasi,  7;  Ptenochirus  jagori,  6;  Pteropus  alecto,  2;  P.  anetianus,  1;  P.  giganteus, 
3;  P.  hypomelanus,  4;  P.  lylei,  1;  P.  melanotus,  1;  P.  rufus,  1;  P.  tonganus,  5;  P.  woodford i, 
7;  Rousettus  amplexicaudatus,  1;  R.  angolensis,  1;  R.  arabicus,  2;  R.  leschenaulti,  3;  R. 
obliviosus,  1;  Scotonycteris  zenkeri,  1;  Eonycteris  spelaea,  3;  Macroglossus  lagochilus,  4; 
M.  minimus,  3;  Megaloglossus  woermanni,  8;  Melonycteris  melanops,  2;  M.  woodfordi, 
4;  Notonycteris  macdonaldi,  11;  Syconycteris  crassa,  4;  Nyctimene  albiventer,  6;  N. 
cephalotes,  3;  N.  major,  6;  N.  robinsoni,  6;  Paranyctimene  raptor,  3;  Harpionycteris 
white  head  i,  1. 

B.  Rhinopomatidae  (9):  Rhinopoma  hardwickei,  5;  R.  microphyllum,  1;  R.  muscatellum, 
3. 

C.  Craseonycteridae  (5):  Craseonycteris  thonglongyai,  5. 

D.  Emballonuridae  (90):  Centronycteris  maximilliani,  1;  Coleura  afra,  5;  Cormura 
brevirostris,  2;  Emballonura  atrata,  1;  E.  beccarii,  6;  E.  monticola,  3;  E.  nigrescens,  3; 
E.  raffrayana,  2;  E.  semicaudata,  5;  Peropteryx  kappleri,  1;  P.  macrotis,  2;  P.  leucopterus,  2; 
Rhynchonycteris  naso,  2;  Saccopteryx  bilineata,  3;  Taphozous  australis,  2;  T.  flaviventris, 
3;  T.  hamiltoni,  2;  T.  hildegardeae,  2;  T.  longimanus,  2;  T.  mauritianus,  3;  7.  melanopogon, 
8;  T.  nudiventris,  4;  T.  peli,  3;  T.  perforatus,  9;  T.  pluto,  5;  T.  saccolaimus,  5;  Cyttarops  alecto, 
1;  Depanycteris  isabella,  1;  Did  id  urns  scutatus,  1;  D.  albus,  1. 

E.  Rhinolophidae  (140):  Rhinolophus  acuminatus,  2;  R.  affinis,  2;  R.  alcyone,  6;  R. 
arcuatus,  2;  R.  blassi,  2;  R.  borneensis,  1;  R.  creaghi,  2;  R.  capensis,  2;  R.  clivosus,  4;  R. 
cornutus,  2;  R.  deckeni,  2;  R.  denti,  2;  R.  euryale,  3;  R.  euryotis,  2;  R.  ferrumequinum,  4; 
R.  fumigatus,  2;  R.  hildebrandti,  2;  R.  hipposideros,  5;  R.  keyensis,  3;  R.  landeri,  3;  R.  lepidus, 
2;  R.  I  act  us,  2;  R.  macrotis,  1;  R.  madurensis,  1;  R.  malayanus,  1;  R.  megaphyllus,  2;  R. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


237 


mehelyi,  1;  R.  pearsoni,  2;  R.  philippinensis,  2;  R.  pusillus,  2;  refulgens,  4;  R.  rouxi,  2; 
R.  shameli,  1;  R.  simulator ,  1;  R.  stheno,  2;  R.  subbad  ins,  2;  R.  swinnyi ,  2;  Hipposideros 
armiger ,  3;  H.  bicolor,  4;  H.  caffer,  8;  H.  camerunensis,  2;  H.  cineraceous,  2;  H.  commersoni, 
4;  H.  cyclops,  4;  H.  diadema,  2;  H.  galeritus,  1;  H.  lankadiva,  3;  H.  larvatus,  3;  H.  lylei,  2; 
H.  pratti,  1;  H.  speoris,  2;  Aselliscus  tricuspidatus,  2;  Asellia  tridens,  3;  Cloeotis  percivali, 
5;  Coelops  frithii,  2;  Triaenops  persicus,  2. 

F.  Nycteridae  (26):  Nycteris  urge,  2;  N.  grandis,  2;  N.  hispida,  4;  N.  javanica,  3;  N. 
macrotis,  6;  N.  thebaica,  5;  N.  tragata,  1;  N.  woodi,  3. 

G.  Megadermatidae  (14):  Cardioderma  cor,  4;  Lavia  frons,  2;  Macroderma  gigas,  1; 
Megaderma  lyra,  5;  M.  spasma,  2. 

PL  Noctilionidae  (6):  Noctilio  albiventris,  4;  N.  leporinus,  2. 

I.  Mormoopidae  (8):  Pteronotus  parnellii,  2;  P.  davyi,  2;  P.  gymnonotus,  2;  Mormoops 
blainvillii,  1;  M.  megalophylla,  1. 

J.  Phyllostomatinae  (183):  1-2  Micronycteris  megalot  is,  14;  3  M.  schmidtorum,  8;  4 
M.  minuta,  8;  5  M.  hirsuta,  4;  6-7  M.  brachyotis,  6;  8  M.  pusilla,  1;  9  M.  nicefori,  8;  10  M. 
sylvestris,  4;  11  M.  behni,  2;  12  M.  daviesi,  5;  13  Macrotus  waterhousii,  4;  14  M.  cali- 
fornicus,  10;  15-16  Lonchorhina  aurita,  13;  17  L.  orinocensis,  1;  18-19  Macrophyllum 
macrophyllum,  8;  20-21  Tonatia  bidens,  7;  22  T.  brasiliensis,  3;  23  T.  carrikeri,  3;  24  T. 
nicaraguae,  5;  25  T.  silvicola,  8;  26  T.  venezuelae,  3;  27  Mimon  bennetti,  1;  28  M.  cozumelae, 
4;  29-30  M.  crenulatum,  12;  31  M.  koepckeae,  1;  32-33  Phyllostomus  discolor,  10;  34  P. 
hastatus,  6;  35  P.  elongatus,  4;  36  P.  latifolius,  2;  37  Phylloderma  stenops,  2;  38  Trachops 
cirrhosus,  10;  39  Chrotopterus  auritus,  3;  40  Vampyrum  spectrum,  5. 

K.  Glossophaginae  (156):  1  Glossophaga  soricina,  6;  2  G.  alticola,  5;  3  G.  commissarisi, 
10;  4  G.  longirostris,  5;  5  Monophyllus  redmani,  6;  6  M.  plethodon,  5;  7  Leptonycteris  nivalis, 
10;  8  L.  sanborni,  3;  9  L.  curasoae,  5;  10  Lonchophylla  Hesperia,  8;  11  L.  mordax,  10;  12  L. 
concava,  2;  13  L.  robusta,  7;  14  L.  thomasi,  10;  15  Lionycteris  spurrelli,  4;  16 
Anoura  geoffroyi,  5;  17  A.  caudifera,  3;  18  A.  cultrata,  5;  19  A.  werckleae,  2;  20  A. 
brevirostrum,  2;  21  Sderonycteris  ega,  1;  22  Lichonycteris  degener,  1;  23-24  L.  obscura, 
7;  25  Hylonycteris  underwoodi,  5;  26  Platalina  genovensium,  6;  27  Choeroniscus  godmani,  3; 
28  C.  minor,  3;  29  C.  intermedins,  6;  30  C.  inca,  3;  31  C.  periosus,  1;  32  Choeronycteris 
mexicana,  10;  33  Musonycteris  harrisoni,  3. 

L.  Carolliinae  (23):  41  Carollia  castanea,  6;  42  C.  subrufa,  2;  43  c.  brevicauda,  4;  44  C. 
perspicillata,  4;  45  RHinophylla  pumilio,  2;  46  R.  alethina,  2;  47  R.  fischerae,  3. 

M.  Stenoderminae  (276):  1  Sturnira  lilium,  5;  2  S.  thomasi,  3;  3  S.  tildae,  5;  4  S.  magna, 
6;  5  S.  mordax,  1;  6  S.  bidens,  6;  7  S.  nana,  5;  8  S.  aratathomasi,  3;  9  S.  ludovici,  10;  10  5. 
erythromos,  6;  11  Uroderma  bilobatum,  10;  12  U.  magnirostrum,  2;  13  Vampyrops  infuscus, 
5;  14  V.  vittatus,  4;  15  V.  dorsalis,  6;  16  V.  aurarius,  6;  17  V.  nigellus,  2;  18  V.  brachycephalus, 
1;  19  V.  helleri,  6;  20  V.  lineatus,  5;  21  V.  recifinus,  2;  22  Vampyrops  sp.  (new  species,  fide 
Gardner  and  Handley),  5;  23  Vampyrodes  caraccioli,  5;  24  Vampyressa  pusilla,  4;  25  V. 
melissa,  6;  26  V.  nymphaea,  3;  27  V.  brocki,  1;  28  V.  bidens,  3;  29  Chiroderma  doriae,  2; 
30  C.  villosum,  6;  31  C.  salvini,  4;  32  C.  trinatatum,  6;  33  C.  improvisum,  1;  34  Ectophylla 
macconnelli,  6;  35  Artibeus  cinereus,  6;  36  A.  glaucus,  2;  37  A.  watsoni,  4;  38  A.  pliaeotis,  6; 
39  A.  toltecus,  5;  40  A.  aztecus,  5;  41  A.  hirsutus,  6;  42  A.  inopinatus,  5;  43  A.  concolor,  5; 
44  A.  jamaicensis,  8;  45  A.  planirostris,  10;  46  A.  lituratus,  8;  47  Artibeus  sp.  (undescribed 
species,  fide  D.  R.  Patten),  10;  48  Enchisthenes  harti,  6;  49  Ardops  nichollsi,  6;  50  Phyllops 
falcatus,  1;  51  P.  haitiensis,  4;  52  Ariteus  flavescens,  6;  53  Stenoderma  rufum,  6;  54 
Pygoderma  bilabiatum,  1;  55  Ametrida  centurio,  8;  56  Sphaeronycteris  toxophyllum,  2; 
57  Centurio  senex,  5. 

N.  Phyllonycterinae  (27):  58  BrachyphyUa  cavernarum,  6;  59  B.  nana,  3;  60  Erophylla 
bombifrons,  3;  61  E.  sezekorni,  5;  62  Phyllonycteris  poeyi,  4;  63  P.  aphylla,  6. 

O.  Desmodontinae  (13):  48-49  Desmodus  rotundas,  5;  50  Diaemus  youngi,  5;  51  Dipliylla 
ecaudata,  3. 

P.  Natalidae  (4):  Natalus  stramineus,  3;  N.  micropus,  1. 


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Q.  Thyropteridae  (3):  Thyroptera  discifera,  2;  T.  tricolor,  1. 

Furipteridae  (1):  Furipterus  horrens,  1;  sample  too  small  for  analysis. 

R  Myzapodidae  (2):  Myzapoda  aurita,  2. 

S.  Vespertilionidae  (178):  Barbastella  barbastellus,  5;  Chcdinolobus  gouldi,  1;  C.  tuber- 
culatus,  2;  C.  variegatus,  2;  Eptesicus  bottae,  6;  E.  hottentotus,  1;  E.  serotinus,  4;  E. 
sonialiscus,  1;  E.  tenuipinnis,  3;  Euderma  maculatum,  2;  Endiscopus  denticulus,  1; 
Hesperoptenus  tickelli,  2;  Histiotus  montanus,  1;  Laephotis  botswanae,  2;  Lasionycteris 
noctivagans,  1;  Lasiurus  borealis,  3;  L.  cinereus,  2;  L.  egregius,  1;  L.  intermedins,  2;  L. 
seminolus,  2;  Minetillus  moloneyi,  3;  Myotis  adversus,  2;  M.  austroriparius,  2;  M.  bech- 
steini,  3;  M.  blythi,  3;  M.  brandti,  1;  M.  capaccinii,  3;  M.  daubentonii,  3;  M.  evotis,  1; 
M.  formosus,  2;  M.  muricola,  3;  M.  myotis,  3;  M.  mystacinus,  4;  M.  nattereri,  2;  M.  emar- 
ginatus,  3;  M.  ricketti,  1;  M.  scotti,  4;  M.  welwitschii,  2;  Nycticeius  humeralis,  2;  N.  schlief- 
feni,  1;  Nycatalus  aviator,  3;  N.  azoreum,  1;  N.  lasiopterus,  4;  N.  leisleri,  2;  N.  noctula,  5; 
Otonycteris  hemprichi,  5;  Scotoecus  hirundo,  2;  Pliiletor  brachypterus,  4;  Pipistrellus  imbri- 
catus,  4;  P.  kuhlii,  1;  P.  nanulus,  I;  P.  pipistrellus,  5;  P.  savii,  1;  P.  subflavus,  3;  Plecotus 
auritus,  5;  P.  phyllotis,  2;  P.  townsendii,  2;  Scotomanes  ornatus,  1;  Scotophilus  gigas,  2;  S. 
heathi,  3;  S.  leucogaster,  2;  Tylonycteris  pachypus,  3;  T.  robustula,  10;  Vespertilio  superans, 
1;  Miniopterus  medius,  2;  M.  schreibersi,  4;  Harpiocephalus  harpia,  1;  Marina  aurata,  3;  M. 
cyclotis,  2;  M.  huttoni,  1;  M.  leucogaster,  1;  Kerivoula  cuprosa,  1;  K.  hardwickei,  2;  K. 
picta,  1;  Nyctophilus  geoffroyi,  1. 

T.  Mystacinidae  (8):  Mystacina  tuberculata,  8. 

U.  Molossidae  (1 12):  Cheiromeles  torquatus,  3;  Eomops  albatus,  1;  Eumops  auripendulus, 
3;  E.  bonariensis,  1;  E.  glaucinus,  1;  E.  hansae,  1;  E.  trumbulli,  1;  E.  underwoodi,  1;  Molossops 
brachymeles,  1;  M.  temmincki,  1;  M.  greenhalli,  1;  Molossus  ater,  4;  M.  bondae,  1;  M. 
crassicaudatus,  1;  M.  molossus,  6;  Otomops  martiensseni,  4;  O.  wroughtoni,  2;  Sauromys 
petrophilus,  3;  Promops  centralis,  1;  P.  davisoni,  1;  P.  nasutus,  6;  Tadarida  aegyptiaca,  2; 
T.  africana,  2;  T.  aloysiisabaudiae,  2;  T.  ansorgei,  3;  T.  aurispinosa,  2;  T.  australis,  2;  T. 
bivittata,  2;  T.  condylura,  3;  T.  congicus,  2;  T.  demonstrator,  2;  T.  doriae,  4;  T.  femorosacca, 
1;  T.  gallagheri,  1;  T.  jobensis,  4;  T.  jugularis,  2;  T.  laticaudata,  3;  T.  leonis,  1;  T.  lobata, 
2;  T.  macrotis,  2;  T.  major,  1;  T.  midas,  2;  T.  nanulus,  2;  T.  nigeriae,  4;  T.  norfolkensis,  2; 
T.  plicata,  1;  T.  pumila,  3;  7.  russata,  2;  T.  sarasinorum,  5;  T.  spurrelli,  2;  T.  teniotus,  3; 
T.  thersites,  1. 


Acknowledgments 

We  are  deeply  indebted  to  the  following  institutions  and  persons  for  making 
available  the  material  examined  by  us:  American  Museum  of  Natural  History, 
Karl  F.  Koopman;  British  Museum  (Natural  History),  John  E.  Hill;  Florida 
State  Museum,  Stephen  Humphrey;  Louisiana  State  University,  Museum  of 
Zoology,  George  H.  Lowery,  Jr.;  Museum  of  Vertebrate  Zoology,  University  of 
California,  Berkeley,  James  L.  Patton;  Museum  of  Southwestern  Biology, 
University  of  New  Mexico,  James  S.  Findley;  Museum  of  Natural  History, 
University  of  Kansas,  Robert  S.  Hoffmann;  The  Museum,  Texas  Tech  University, 
Hugh  H.  Genoways;  Natural  History  Museum  of  Los  Angeles  County,  Lan  Lester 
and  Donald  Patten;  Naturhistorisch  Museum,  Wien,  Kurt  Bauer;  Natur-Museum 
Senckenberg,  Frankfurt,  Heinz  Felten  and  Dieter  Kock;  Royal  Ontario  Museum, 
R.  L.  Peterson  and  Judith  L.  Eger;  United  States  National  Museum,  including  the 
Biological  Surveys  Collection,  Alfred  L.  Gardner,  Don  E.  Wilson,  and  C.  O. 
Handley,  Jr. 

We  also  wish  to  thank  Russell  Benson,  Department  of  Mathematics,  California 
State  University,  Fullerton  (CSUF),  for  his  assistance  in  developing  the  calculation 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


239 


of  geometric  variables.  Steven  Eich,  James  Lamprecht,  Monte  D’Asta,  and  Mark 
Hartman,  Computer  Center,  CSUF,  provided  valuable  aid  and  advice  in  Fortran 
programming  and  computer  processing.  We  are  especially  grateful  to  Richard  A. 
Pimentel,  California  Polytechnic  University,  San  Luis  Obispo,  who  unselfishly 
assisted  us  with  the  multivariate  analyses,  which  included  using  an  unpublished 
program  (DISANAL)  that  he  developed.  He  also  reviewed  the  manuscript  and 
provided  assistance  in  its  preparation. 

Finally,  we  wish  to  express  our  gratitude  to  Susan  E.  Smith  who  sat  for  many 
hours  recording  measurements,  keypunched  data,  helped  with  the  illustrations, 
and  most  of  all  provided  moral  support  and  companionship  to  the  senior  author. 

Results  and  Discussion 

The  mean  (range  in  parentheses),  one  standard  error,  and  coefficient  of 
variation  for  the  raw  variables  and  selected  derived  variables  are  given  in  the 
Appendix  (Tables  A1-A21).  A  pooled  correlation  matrix  for  raw  variables  was 
computed,  and  all  coefficients,  except  those  for  the  third  phalanx  of  digit  III, 
were  strongly  and  positively  correlated  (P<  0.001).  This  was  to  be  expected  owing 
to  the  size/growth  nature  of  these  variables.  The  coefficients  for  the  third  phalanx 
of  digit  III  were  low  because  this  phalanx  is  not  present  in  all  groups  of  bats 
(for  example,  pteropodids,  emballonuroids,  rhinolophoids,  see  Miller,  1907). 
The  largest  coefficients  of  correlation  for  this  phalanx  were  shown  with  the 
metacarpal  and  two  phalanges  of  digit  V,  0.405  (P<0.05)  and  0.325  (P<0.05), 
respectively. 

Principal  components  analysis. — The  results  of  the  principal  components 
analysis  are  given  in  Figs.  3  and  4  and  Tables  2  and  3.  Because  of  the  notorious 
susceptibility  of  the  first  component  axis  to  size  factors,  this  analysis  yields  only 
broad  generalizations  concerning  the  shape  of  bat  wings.  The  first  component, 
usually  designated  the  “size  component,”  exhibits  91.8  per  cent  of  the  total 
variation  (Table  2).  Also,  the  component  correlations  for  all  variables  are  high 
for  this  component.  The  first  three  components  account  for  96.7  per  cent  of  the 
total  variation.  Although  component  loading  extends  to  the  twelfth  component, 
99.1  per  cent  is  accumulated  by  the  sixth.  The  majority  of  the  loading,  past  the 
first  three  components,  is  contributed  by  the  third  phalanx  of  digit  III,  which 
exhibits  high  loading  in  the  fourth  and  seventh  component  (51.36  and  13.75  per 
cent,  respectively). 

As  noted  above,  the  first  component  contains  high  loading  as  the  result  of 
general  size.  This  is  illustrated  by  the  complete  agreement  of  signs  by  all 
coefficients  in  this  component  (Table  2).  The  direction  of  the  sign  (negative,  in 
this  case)  is  irrelevant  and  simply  indicates  that  all  variables  increase  ( + )  or 
decrease  ( — )  in  the  same  direction  (for  example,  length  of  the  head  and  body 
decreases  in  consort  with  length  of  the  forearm  or  any  of  the  other  raw  variables). 
The  fact  that  the  component  scores  for  each  variable  are  of  different  magnitude 
indicates  general  positive  allometry  among  the  variables.  The  effect  of  size  in 
the  first  component  also  can  be  seen  in  Figs.  3  and  4.  In  figure  4A-B,  the  agree- 


240 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  2. — Eigenvectors  (direction  cosines)  of  principal  components  for  lengths  of  head  and 
body  and  selected  wing  elements.  Only  the  first  three  components  are  shown  because  most 
of  the  variation  is  exhibited  in  these  components.  The  numbers  in  parentheses  following  each 
component  score  indicates  the  percentage  of  variance  contributed  by  each  variable  to  a 

particular  component. 


Variable 

Component  Axes 

Cumulative 
per  cent 

l 

2 

3 

Head  and  body 

-0.626(96.07) 

0.701  (  3.53) 

-0.075  (  0.03) 

99.63 

Forearm 

-0.427  (96.77) 

-0.173  (  0.46) 

0.176  (  0.38) 

97.61 

Metacarpal  III 

-0.313  (92.52) 

-0.124  (  0.42) 

0.485  (  5.13) 

98.07 

Digit  III,  phalanx  1 

-0.186  (89.16) 

-0.038  (  0.11) 

—  0.130  (  1.00) 

90.27 

Digit  III,  phalanx  2 

-0.244  (84.53) 

—  0.165  (  1.13) 

-0.525  (  9.03) 

94.32 

Digit  III,  phalanx  3 

-0.022  (  6.03) 

-0.247  (21.63) 

0.014  (  0.05) 

27.71 

Metacarpal  IV 

-0.303  (93.16) 

-0.261  (  2.02) 

0.362  (  3.08) 

98.26 

Digit  IV,  phalanx  1 

-0.150(86.65) 

-0.069  (  0.53) 

-0.139  (  1.70) 

88.88 

Digit  IV,  phalanx  2 

-0.133  (70.89) 

-0.150  (  2.62) 

-0.440  (17.89) 

91.40 

Metacarpal  V 

-0.276  (86.51) 

-0.505  (  8.51) 

-0.064  (  0.11) 

95.13 

Digit  V,  phalanx  1 

-0.116(82.78) 

-0.106  (  2.04) 

-0.094  (  1.26) 

86.08 

Digit  V,  phalanx  2 

-0.107  (75.09) 

-0.1 14  (  2.50) 

-0.280  (11.90) 

89.49 

Per  cent  trace 

91.8 

2.7 

2.1 

Cumulative  per  cent 

91.8 

94.5 

96.7 

ment  among  the  signs  of  the  first  component  scores  is  manifested  by  all  vectors  of 
variables  (direction  cosines)  orienting  toward  the  left.  Likewise,  the  ordination  of 
group  centroids  along  the  first  component  axis  (Fig.  3)  aligns  large-sized  bats 
(Pteropodidae,  A)  to  the  left,  and  small-sized  bats  (Craseonycteridae,  C)  to  the 
right.  Also,  it  should  be  noted  that  the  nature  of  the  ordination  of  groups  (Fig.  3) 
is  greatly  influenced,  especially  in  the  first  two  component  axes,  by  the  magnitude 
of  the  eigenvalues  for  head  and  body  length  (  —  0.626  and  0.701,  Table  2  and 
Fig.  4A-C).  Other  vectors  of  variables  that  markedly  affect  the  ordination  along 
the  first  component  are  the  lengths  of  the  forearm  (B)  and  the  metacarpals  of 
digits  III-V  (C,  G,  J)  (  —  0.427,  —0.313,  and  —0.276,  respectively). 

In  the  second  component,  all  coefficients,  except  that  for  the  length  of  the  head 
and  body,  agree  in  sign  (Table  2).  This  strongly  suggests  that  the  size  and  shape 
of  bat  wings  are  essentially  independent  of  body  size  and,  presumably,  weight. 
The  fact  that  all  of  the  coefficients  for  intrinsic  wing  elements  vary  in  magnitude 
continues  to  indicate  a  level  of  positive  allometry.  Other  than  head  and  body  length, 
the  strongest  eigenvalue  in  this  component  axis  is  that  for  the  fifth  metacarpal 
(  —  0.505).  It  is  difficult  to  evaluate  the  shape  tendencies  in  the  second  component 
because  the  correlation  structure  is  rather  weak  in  both  this  and  the  third  com¬ 
ponent.  In  addition,  a  minor  portion  of  the  variation  is  shown  in  these  two  com¬ 
ponents  compared  to  the  overwhelming  nature  of  the  first.  A  cautious  in¬ 
terpretation  of  the  shape  trends  in  the  second  component  might  be  that  shape  is 
modified  by  a  factor  of  size. 

Influence  attributable  to  shape  are  much  more  distinct,  albeit  weak,  among 
the  coefficients  of  the  third  component.  Body  size,  as  expressed  by  the  length  of 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


241 


Table  3. — Mean  coordinates  of  group  centroids  from  the  principal  components  analysis.  These 

centroids  are  plotted  in  Fig.  3. 


Taxon 

Code 

Component  axes 

l 

2 

3 

Pteropodidae 

A 

-68.73 

9.16 

-21.46 

Rhinopomatidae 

B 

16.06 

1.95 

5.81 

Craseonycteridae 

C 

60.81 

-1.68 

-9.15 

Emballonuridae 

D 

2.23 

1.05 

10.75 

Rhinolophidae 

E 

9.84 

-2.62 

-1.56 

Nycteridae 

F 

13.07 

-4.78 

-5.53 

Megadermatidae 

G 

-17.38 

-6.65 

-11.30 

Noctilionidae 

H 

-24.10 

-17.87 

2.90 

Mormoopidae 

I 

8.07 

-4.62 

10.09 

Phyllostomatinae 

J 

-0.64 

-7.09 

0.27 

Glossophaginae 

K 

19.63 

2.86 

-1.17 

Carolliinae 

L 

24.47 

-4.63 

-4.36 

Stenoderminae 

M 

4.14 

-6.08 

-1.59 

Phyllonycterinae 

N 

-2.26 

-0.75 

3.41 

Desmodontinae 

O 

-15.05 

-5.44 

7.14 

Natal  idae 

P 

32.07 

-9.13 

0.37 

Thyropteridae 

Q 

40.68 

-4.96 

6.70 

Myzapodidae 

R 

13.70 

-9.45 

1.06 

Vespertilionidae 

S 

19.24 

-2.43 

6.15 

Mystacinidae 

T 

18.16 

-0.76 

6.14 

Molossidae 

U 

2.35 

9.66 

9.51 

head  and  body,  has  little  influence  in  this  component,  having  expended  most  of 
its  force  in  the  ordination  of  the  first  and  second  component  axes.  It  will  be  noted 
(Table  2)  that  several  of  the  wing  elements,  notably  the  third  and  fourth  meta- 
carpals  (C,G)  and  the  second  phalanges  of  digits  III-V  (E,I,L),  have  their  largest 
eigenvalues  in  the  third  component.  The  divergence  of  variable  vectors,  caused  by 
differential  signs  in  the  third  component  axis,  further  substantiates  the  shape  trends 
of  this  component  (Fig.  4B-C).  Bearing  in  mind  that  only  a  small  portion  of  the 
variation  is  expressed  and  the  weak  correlation  structure  of  the  third  component, 
we  cautiously  direct  attention  to  several  interesting  associations  among  the 
variables  in  this  component. 

In  Figure  4B-C,  the  vectors  for  variables  of  all  intrinsic  wing  elements  (B-L) 
are  directed  to  the  left;  the  vector  for  head  and  body  length  (A)  projects  to  the 
right  in  the  2X3  graph  (Fig.  4C)  again  indicating  the  independent  nature  of  this 
variable.  As  noted  previously,  the  general  similarity  in  the  direction  of  orientation 
of  all  vectors  for  wing  elements  postulates  a  general  allometric  relationship  among 
wing  components  in  terms  of  size.  However,  in  the  two  graphs  (1X3  and  2  X  3), 
the  vectors  for  wing  components  diverge  into  different  regions  of  the  graphs  (that 
is,  some  orient  upward  and  others  are  directed  downward).  This  signifies  dif¬ 
ferences  in  relative  independence  that  ultimately  are  expressed  as  shape. 

The  vectors  for  the  third  and  fourth  metacarpals  (C,  G)  project  in  the  same 
general  direction  and  are  nearly  equal  in  length  (Fig.  4B-C),  indicating  that  their 


242 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  3. — Component  graph  from  principal  component  analysis.  Group  centroids  are 
plotted  on  the  1  X2  axes  and  1  X3  axes  to  illustrate  their  position  in  Euclidean  three-space. 
Coordinates  for  these  centroids  are  given  in  Table  3.  Stars  represent  phyllostomatid  centroids 
(see  list  of  specimens  examined  or  Table  3  for  key  to  alphabetic  code).  This  figure  may  be 
xeroxed  and  folded  on  the  dotted  line  to  help  visualize  the  three-dimensionality  of  the  dis¬ 
persion  of  centroids. 

variation  is  associated.  Although  somewhat  removed,  the  vector  for  the  forearm 
(B)  tends  to  share  this  same  general  relationship.  It  is  interesting  to  note  that  the 
vector  for  the  fifth  metacarpal  (J)  is  rather  far  removed  from  the  third  and  fourth 
metacarpals  thereby  suggesting  a  marked  divergence  in  its  pattern  of  variation. 
This  suggests  that  the  forearm  and  metacarpals  of  digits  III  and  IV  vary  as  a  unit, 
whereas  the  metacarpal  of  the  fifth  digit  is  somewhat  independent.  Following 
these  examples,  we  can  point  to  several  additional  interesting  sets  of  vectors  that 

Fig.  4. — Positive  eigenvectors  (A-C)  and  variable  vectors  (D-F)  for  the  raw  variables 
computed  in  the  principal  components  analysis  and  discriminant  analysis,  respectively. 
Coordinates  for  these  vectors  are  given  in  Tables  2  and  4,  respectively.  Corresponding  sets 
of  vectors  from  these  two  analyses  are  shown  side-by-side  to  allow  easy  comparison.  The 
negative  portions  of  the  vectors  were  omitted  to  avoid  confusing  the  diagram.  If  shown,  they 
would  project  an  equal  distance  in  the  opposite  direction  past  the  zero-zero  point.  Letters  at 
the  ends  of  vectors  refer  to  the  respective  lengths  of  variables:  A,  head  and  body;  B,  forearm; 
C,  metacarpal  III;  D,  first  phalanx  III;  E,  second  phalanx  III;  F,  third  phalanx  III;  G, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


243 


metacarpal  IV;  H,  first  phalanx  IV;  I,  second  phalanx  IV;  J,  metacarpal  V;  K,  first  phalanx 
V;  L,  second  phalanx  V.  See  text  for  discussion. 


244 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


have  generally  associated  patterns  of  variation.  The  first  phalanges  of  digits  III 
and  IV  (D,  H)  appear  to  have  a  similarly  related  effect  on  wing  shape.  Likewise, 
the  vectors  for  the  second  phalanges  of  digits  III  to  V  (E,  I,  L)  suggest  a  similar 
relationship  among  these  phalangeal  elements.  These  two  sets  of  variables,  to¬ 
gether,  diverge  markedly  from  the  metacarpal  elements  (C,  G,  J)  of  their  respective 
digits.  The  vector  for  the  first  phalanx  of  digit  V  (K)  tends  to  associate  with  the 
fifth  metacarpal  (J).  These  patterns  of  positive  allometry  generally  indicate  the 
complexities  of  wing  shape. 

As  we  have  seen,  size  greatly  influences  the  ordination  of  groups  in  the  principal 
components  analysis.  This  is  exerted  strongly  in  the  first  component  and  hardly 
at  all  in  subsequent  components.  The  overwhelming  effect  of  size  has  led  many 
investigators  to  attempt  to  eliminate  size  as  an  ordinating  factor  and  thereby  in¬ 
crease  the  component  loading  by  the  “inherent”  shaping  qualities  of  their  raw 
variables.  The  product  of  these  efforts  has  been  the  mathematical  adulteration 
of  raw  variables  into  ratios,  indices,  and  proportions,  which  may  appear  to 
eliminate  size,  but  which  actually  obscure  or  otherwise  confound  the  recognition 
of  independent  patterns  of  variation.  Simply  ignoring  the  first  component  and 
considering  components  2-4  is  not  a  satisfactory  means  of  eliminating  size,  be¬ 
cause  the  component  correlations  are  usually  even  weaker  in  the  fourth  compo¬ 
nent.  We  submit  that  in  a  morphometric  analysis  such  as  this,  and  in  fact  in  all 
analyses  based  on  absolute  measures  of  continuous  variables,  size  reflects  the 
essence  of  variation.  By  this,  we  do  not  mean  absolute  size  in  itself,  but  the  allo- 
metric  and  isometric  aspects  of  size  that  ultimately  are  expressed  as  synergistic 
relationships  among  variables.  Therefore,  any  attempt  to  strip  away  the  effects  of 
size  seriously  risks  masking  or  totally  eliminating  the  interactive  relationships 
between  size  and  shape. 

The  centroids  computed  for  each  group  in  the  principal  components  analysis 
are  given  in  Table  3  and  plotted  in  Fig.  3.  The  cigar-shaped  dispersion,  as  noted 
earlier,  is  oriented  with  the  longest  axis  more  or  less  corresponding  to  the  first 
component  axis.  The  shape  of  this  cluster  is  caused  mostly  by  the  effects  of  gross 
size.  Most  taxa,  including  the  six  subfamilies  of  phyllostomatids  (J  to  O),  are 
packed  in  the  midregion  of  the  dispersion.  By  examining  the  vectors  of  variables 
shown  in  Fig.  4A-C  and  the  group  centroids  plotted  in  Fig.  3,  the  reader  can 
begin  to  appreciate  the  ordinating  effects  exerted  by  the  various  characters.  In 
the  lower  diagram  of  Fig.  3  (axes  1  X  2),  the  pteropodids  (A)  are  pushed  to  the  far 
left  and  into  the  upper  quadrant,  primarily  on  the  basis  of  large  head  and  body 
length.  The  noctilionids  (H),  megadermatids  (G),  and,  to  a  lesser  extent,  the 
desmodontines  (O)  also  are  influenced  by  the  positive  force  of  this  vector.  The 
craseonycterids  (C),  on  the  other  hand,  ordinate  into  the  lower  right-hand  quadrant 
by  the  opposite  (negative)  effect  of  the  vector  for  head  and  body  length.  The  taxa 
in  the  lower  left-hand  quadrant  are  ordinated  by  the  positive  (large  size)  effects  of 
all  vectors  of  variables  for  wing  elements;  especially  lengths  of  the  forearm,  second 
phalanx  of  digit  III,  third  and  fourth  metacarpals,  and  second  phalanx  of  digit 
V.  The  taxa  in  the  upper  right-hand  quadrant  ordinate  by  the  negative  (small 
size)  effects  of  these  wing  elements.  Note  that  the  phyllostomatines  (Fig.  3J) 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


245 


are  pushed,  almost  directly,  by  the  vector  for  the  fifth  metacarpal  (Fig.  4J),  where¬ 
as  the  molossids  (U)  and  emballonurids  (D)  lie  along  the  tail  end  of  this  vector. 
The  majority  of  the  taxa  are  ordinated  into  the  lower  right-hand  quadrant,  which 
results  from  a  complex  synergistic  interaction  among  the  intrinsic  elements  of 
the  wing. 

The  effects  of  the  vector  for  variables  in  the  third  component  may  be  seen  in  the 
upper  diagram  of  Fig.  3  (axes  1  X  3).  In  this  component  graph,  the  pteropodids 
(A)  and  megadermatids  (G)  are  ordinated  into  the  lower  left-hand  quadrant  by 
large-sized,  distal  phalangeal  elements  (E,  I,  L).  In  these  two  groups,  the  meta- 
carpals  constitute  a  relatively  smaller  portion  of  the  total  length  of  the  various 
digits  (Fig.  7;  Tables  A12,  A15,  A16).  On  the  other  hand,  noctilionids  (H), 
desmodontines  (O),  and,  to  a  lesser  extent,  carolliines  (N)  are  characterized  by  a 
generally  long  forearm  (B),  third  and  fourth  metacarpal  (C,  G),  and  third  phalanx 
of  digit  III  (F).  The  taxa  positioned  in  the  upper  right-hand  quadrant  generally 
reflect  a  complex  synergism  among  variables. 

In  summary  to  this  point,  principal  components  analysis  is  an  effective  screening 
procedure  that  allows  some  general  insights  into  the  interactive  relationships  of 
size  and  shape  exhibited  by  the  wings  of  bats.  However,  this  procedure,  because 
of  its  sensitivity  to  gross  size,  is  not  well  suited  to  the  detection  of  subtle  nuances 
in  the  variation  of  wing  shape  among  chiropterans.  It  provides  a  generalized  view 
of  the  tip  of  the  iceberg,  so  to  speak,  but  does  not  give  a  clear  perspective  of  the 
underlying  complexity  of  shape.  With  regard  to  the  phyllostomatids  as  a  group, 
little  can  be  said  other  than  that  they  tend  to  ordinate  amongst  the  medium  to 
large-sized  bats  near  the  grand  centroid. 

Discriminant  analysis. — The  transformation  from  Euclidean  space  into 
discriminant  space  effectively  reduces  the  overwhelming  influence  of  general 
size  on  the  ordination  of  group  centroids  without  otherwise  adulterating  the 
intrinsic  variation  of  the  raw  variables.  In  Table  4,  there  is  a  more  equitable 
dispersal  of  the  variation  across  the  first  six  canonical  axes.  There  is  much  more 
symmetry  shown  by  the  canonical  vectors  of  variables  in  Fig.  4D-F  than  by  vectors 
from  the  component  analysis  (Fig.  4A-C).  In  addition,  the  correlations  of 
canonical  vectors  and  variables  are  more  evenly  dispersed  across  the  various 
canonical  axes  rather  than  being  heavily  focused  in  the  first  axis  as  was  the  case 
in  the  principal  components  analysis. 

It  should  be  pointed  out  that,  although  the  variable  vector  for  the  third  phalanx 
of  digit  III  (F)  is  not  particularly  strong  as  compared  to  other  vectors,  its  influence 
on  the  dispersion  in  the  first  canonical  axis  essentially  segregates  taxa  into  two 
groups — those  that  possess  this  element  and  those  that  do  not.  The  correlation 
coefficient  for  this  variable  with  the  first  canonical  axis  is  comparatively  high 
(0.540).  This  is  equalled  by  the  correlation  coefficients  for  the  fifth  metacapal 
(J)  and  second  phalanx  of  digit  V  (L),  which  have  their  greatest  affinity  with  the 
third  canonical  axis  (0.489  and  0.582,  respectively).  A  more  detailed  discussion 
of  the  effects  of  these  various  variable  vectors  on  the  size  and  shape  of  chiropteran 
wings  will  be  presented  in  the  following  accounts. 


246 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


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BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


247 


Table  5. — Group  centroids  for  the  first  six  canonical  axes.  The  first  three  axes  are  plotted 

in  Figs.  5  and  6. 


Canonical  axes 


Taxon 

Code 

1 

2 

3 

4 

5 

6 

Pteropodidae 

A 

-3.738 

-2.621 

2.456 

0.992 

-0.101 

-0.294 

Rhinopomatidae 

B 

-2.791 

3.379 

-2.446 

4.291 

0.374 

-3.076 

Craseonycteridae 

C 

-1.660 

0.525 

-0.725 

-2.080 

-1.265 

-0.199 

Emballonuridae 

D 

-0.733 

0.376 

-2.779 

2.421 

-1.728 

1.512 

Rhinolophidae 

E 

-3.741 

1.106 

-1.076 

-1.379 

0.010 

-0.547 

Nycteridae 

F 

-3.711 

-0.434 

-0.739 

-3.183 

0.863 

2.374 

Megadermatidae 

G 

-4.678 

1.284 

-0.556 

-4.321 

-2.089 

1.244 

Noctilionidae 

H 

3.106 

1.231 

-3.222 

-2.933 

-3.010 

-3.535 

Mormoopidae 

I 

2.203 

1.986 

-0.166 

1.015 

-2.226 

-1.714 

Phyllostomatinae 

J 

1.364 

1.412 

1.114 

-0.510 

-0.535 

-0.170 

Glossophaginae 

K 

1.778 

0.770 

1.622 

-0.059 

-0.792 

0.518 

Carolliinae 

L 

1.590 

0.298 

1.608 

-0.233 

-0.538 

1.227 

Stenoderminae 

M 

2.752 

1.360 

2.018 

-0.124 

-0.294 

0.345 

Phyllonycterinae 

N 

1.028 

1.885 

1.073 

0.891 

0.758 

-0.912 

Desmodontinae 

O 

2.207 

3.805 

1.199 

-0.187 

-0.700 

-1.650 

Natal  idae 

P 

-2.076 

1.060 

-1.075 

-0.762 

0.158 

2.415 

Thyropteridae 

Q 

-0.024 

0.323 

-0.695 

-0.636 

2.004 

1.694 

Myzapodidae 

R 

1.146 

0.288 

0.051 

-0.416 

1.367 

-1.053 

Vespertilionidae 

S 

0.781 

0.480 

-0.843 

0.423 

1.813 

0.291 

Mystacinidae 

T 

2.315 

2.132 

1.447 

0.774 

1.658 

-1.224 

Molossidae 

U 

3.194 

-3.510 

-1.523 

-0.585 

-0.120 

-0.474 

Pteropodidae 

Fruit  bats  are  generally  the  largest  chiropterans  in  terms  of  absolute  size  of  all 
raw  variables.  We  have  observed  that  overall  large  size  greatly  effects  the 
ordination  in  the  principal  components  analysis.  However,  in  discriminant  space, 
these  overwhelming  effects  of  size  are  much  reduced.  Because  of  their  large  size, 
the  pteropodids  are  especially  well  suited  to  illustrate  the  moderation  of  size  in 
the  discriminant  analysis.  The  strongest  vector  in  the  principal  component  analysis 
was  that  for  head  and  body  length  (A) — see  Fig.  4A-C.  This  feature  in  the  dis¬ 
criminant  analysis  is  one  of  the  least  powerful  (Fig.  4D-F;  Tables  4  and  6).  Not 
only  is  the  length  of  the  vector  short  as  compared  to  others  such  as  those  for  the 
lengths  of  the  forearm  (B),  and  third  and  fifth  metacarpal  (C,  J),  for  example, 
but  it  is  directed  away  (approximately  90  degrees)  from  the  group  centroid  for 
pteropodids.  The  canonical  coefficient  (0.225)  for  head  and  body  length  in  the 
first  canonical  axis  (Table  4)  is  positive  and  near  zero,  suggesting  the  denial  of 
large  body  size  by  pteropodids  relative  to  this  axis.  Although  comparatively  mi¬ 
nor,  the  greatest  influence  by  this  variable  on  the  ordination  of  bats  in  discrimi¬ 
nant  space  occurs  in  the  second  and  third  axes,  but  here  too  the  vector  generally 
orients  away  from  the  pteropodids.  Furthermore,  the  contribution  of  this  variable 
to  the  overall  discriminant  functions  of  the  various  centroids  appears  to  be  minor 
(Table  6). 


248 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  5. — Canonical  graph  from  discriminant  analysis.  Group  centroids  are  plotted  with 
their  respective  confidence  circles  (95  per  cent)  on  1X3  canonical  axes  of  discriminant 
space  and  the  coordinates  for  these  are  given  in  Table  5.  Stars  and  stippled  area  indicate 
phyllostomatid  centroids  and  confidence  circles.  The  confidence  circle  for  the  Myzapodi- 
dae  (R)  is  too  large  to  plot.  See  list  of  specimens  examined  or  Table  5  for  key  to  alphabetic 
code  and  text  for  discussion. 

The  length  of  the  forearm  (B)  is  one  of  the  more  powerful  forces  in  the  overall 
ordination  of  groups  (Tables  4,  and  6).  In  the  first  two  canonical  axes,  the  vector 
for  this  variable  lies  approximately  perpendicular  to  the  pteropodid  centroid. 
Relative  to  the  molossid  centroid  (U),  this  vector  may  be  interpreted  as  exerting 
a  positive  force  on  the  pteropodid  centroid.  However,  the  peripheral  position  of 
this  centroid  to  the  vector  suggests  a  weak  influence  by  this  variable.  In  the  third 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


249 


Fig.  6. — Canonical  graph  from  discriminant  analysis.  Group  centroids  are  plotted  with 
their  respective  confidence  circles  (95  per  cent)  on  1X2  canonical  axes  of  discriminant 
space  and  the  coordinates  for  these  are  given  in  Table  5.  Stars  and  stippled  area  indicate 
phyllostomatid  centroids  and  confidence  circles.  The  confidence  circle  for  the  Myzapodidae 
(R)  is  too  large  to  plot.  See  list  of  specimens  examined  or  Table  5  for  key  to  alphabetic 
code  and  text  for  discussion. 

canonical  axis  (Figs.  4E,  5),  the  effect  of  this  variable  is  somewhat  more  direct 
(negative).  A  heuristic  interpretation  of  this  variable  vector  would  suggest  a 
medium  to  short  forearm  for  the  pteropodids. 

An  examination  of  the  relative  lengths  of  the  wing  and  forearm  (Tables  A7, 
A8)  clarify  and  substantiate  this  interpretation.  Although  the  absolute  lengths  of 
all  wing  elements  are  large,  the  wings  of  pteropodids  average  proportionately 


250 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


shorter  than  those  of  other  chiropterans  (1.80).  The  mean  relative  length  of  the 
forearm  is  second  smallest  for  the  order  (0.65);  only  that  of  molossids  is  smaller 
(0.63).  In  terms  of  the  composition  of  wing  span,  the  forearm  of  pteropodids 
contributes  an  average  of  35.98  per  cent  (range,  32.69-38.84)  to  the  length  of  the 
wing. 

The  canonical  coefficients  for  the  metacarpals  of  digits  III-V  (C,  G,  J) 
illustrate  the  simultaneous  nature  of  the  interactive  relationships  among  these 
variables.  By  comparing  Fig.  4D  with  Fig.  6,  it  will  be  noted  that  the  negative 
end  (smallness)  of  the  variable  vector  for  the  third  metacarpal  (C)  passes  in 
proximity  to  the  centroid  of  pteropodids.  This  indicates  a  rather  strong  tendency 
in  the  direction  of  small  size,  especially  in  the  first  and  second  canonical  axes. 
The  variable  vectors  for  the  fourth  and  fifth  metacarpals  (G,  J)  orient  in  nearly 
opposite  directions  from  each  other,  and  both  orient  almost  perpendicularly  to  the 
centroid  for  pteropodids.  It  will  be  noted  that  the  latter  vector  (J)  is  the  stronger 
of  the  two  (Table  4)  and  it  is  oriented  directly  toward  the  rhinopomatids.  The 
vector  for  the  fourth  metacarpal  (G)  is  of  a  lesser  magnitude  and  is  oriented 
generally  toward  the  centroid  of  the  Molossidae.  The  influences  of  these  two 
variable  vectors  on  these  two  centroids  will  be  discussed  beyond  and  are 
mentioned  here  only  for  orientation  by  the  reader.  It  is  difficult  to  assess  the  nature 
of  the  effect  these  two  vectors  have  on  the  pteropodid  centroid  in  the  first  and 
second  canonical  axes;  suffice  it  to  say  that  it  is  synergistic.  In  the  third  canonical 
axis  (Figs.  4E,  5),  the  interaction  of  these  three  vectors  is  somewhat  clearer.  The 
vector  for  the  third  metacarpal  (C)  continues  in  its  implication  of  small  size.  The 
vectors  for  the  fourth  and  fifth  metacarpals  (C,  J)  maintain  their  opposite  orien¬ 
tation,  but  their  negative  (smallness)  ends  are  closer  to  the  centroid  of  the 
Pteropodidae  than  before.  The  net  effect  of  all  three  of  these  vectors  is  to  carry 
the  centroid  in  an  upward  direction  in  three-dimensional  space  and,  because  it  is 
the  tail  end  of  these  vectors  that  effects  this  lifting,  the  implication  is  small  size 
for  all  three  metacarpals.  An  examination  of  Tables  A12,  A16,  and  A19  reveals 
that  these  manal  elements  of  the  pteropodid  wing  contribute  the  smallest  per¬ 
centage  to  the  overall  lengths  of  digits  III  to  V  as  compared  to  other  chiropteran 
taxa.  Norberg  (1972)  also  noted  the  general  shortness  of  the  metacarpals  of  the 
megachiropterans. 

A  long  first  phalanx  of  digits  III  and  V  (D,  K)  is  strongly  implicated  in  the 
discrimination  of  pteropodidis  in  all  three  canonical  axes.  The  vector  for  this 
phalanx  in  the  fourth  digit  (H)  is  most  influential  in  the  third  canonical  axis  (Figs. 
4E-F,  5,6;  Table  4)  and  here  also  suggests  relatively  long  length.  The  vectors  for 
the  second  phalanx  of  digits  IV  and  V  (I,  L)  share  a  similar  orientation  as 
described  above  for  the  fourth  and  fifth  metacarpals  (G,  J)  except  that  the  positive 
ends  of  these  variable  vectors,  rather  than  the  negative  ends,  carry  the  centroid 
aloft.  An  examination  of  the  percentages  contributed  to  the  discrimination  of  each 
group  (Table  6)  generally  substantiates  the  characteristically  long  phalangeal 
elements  of  pteropodids.  In  addition,  Tables  A 13,  A 17,  A 18,  A20,  and  A21  show 
the  mean  percentages  contributed  by  these  phalanges  to  the  overall  lengths  of 
digits  III  to  V,  respectively,  and  further  support  the  above  interpretations  by 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


251 


ranking  the  pteropodids  as  the  largest,  or  nearly  so,  with  respect  to  these  wing 
elements. 

The  vector  for  the  second  phalanx  of  digit  III  (E)  presents  an  interesting  paradox 
in  that  it  nearly  parallels,  in  both  direction  and  sign,  the  orientation  of  the  variable 
vector  for  the  third  metacarpal  (C).  This  seems  to  suggest  short  length  of  this 
feature  in  the  first  two  canonical  axes.  However,  there  is  a  slight  elevating  quality 
by  the  point  of  this  vector  on  the  centroid  in  the  third  dimension  of  discriminant 
space.  Those  familiar  with  pteropodid  wings  should  be  duly  impressed  by  the 
extraordinary  length  and  massive  structural  nature  of  this  phalanx.  However, 
though  this  wing  element  is  outwardly  large-sized  in  appearance,  pteropodids 
rank  fourth  largest  in  terms  of  the  average  percentage  contributed  by  this  element 
to  the  overall  length  of  digit  III,  as  compared  to  the  Craseonycteridae,  Megader- 
matidae,  and  Furipteridae  (Table  A 14).  (The  latter  group  was  not  included  in  the 
multivariate  analyses  because  the  sample  size  was  too  small.) 

Therefore,  pteropodids  are  characterized  by  having  a  relatively  short  wing  as 
the  combined  result  of  a  relatively  short  forearm  and  third  metacarpal  (Fig.  7). 
Although  the  two  phalangeal  elements  of  the  third  digit  are  long,  the  shortness 
of  the  metacarpal  tends  to  suppress  the  overall  length  of  the  digit.  The  total  length 
of  digit  III  contributes  between  61  and  67  per  cent  to  the  wingspan,  shown  by  a 
mean  tip  index  of  1.78  (2.06-1.57),  which  ranks  in  the  middle  to  upper  range  of 
all  bats  (Table  A2).  The  wings  of  pteropodids  are  further  characterized  by  their 
generally  broad  aspect  (Fig.  10;  Tables  A3-A5).  Although  the  shortness  of  the 
fourth  and  fifth  metacarpals  would  tend  to  cause  a  narrow  wing,  apparently  the 
combined  lengthening  of  the  phalangeal  elements  of  digits  III  to  V  maintains  the 
proportional  breadth. 

Contrary  to  Findley  et  al.  (1972),  such  a  wing  should  have  an  excellent  lift 
potential  at  slow  speeds.  In  addition,  the  relatively  long  phalanges  of  all  three 
digits,  especially  those  of  digits  IV  and  V,  should  facilitate  increased  camberability 
with  relatively  little  digital  flexion  and  thereby  further  augment  lift  potential  at 
slow  speeds.  Whereas  the  nearly  equal  (isometric)  partitioning  of  the  respective 
digits  may  contribute,  in  a  crude  sort  of  way,  to  the  slow-flight  characteristics 
of  pteropodid  wings,  the  fine  adjustments  necessary  for  maneuverability  in  slow 
flight,  such  as  hovering,  apparently  are  not  possible.  In  the  following  accounts 
we  will  show  that  all  other  chiropteran  families  depart  from  the  general  isometric 
construction  of  the  wing  as  exhibited  by  the  Pteropodidae. 

Rhinopomatidae 

As  in  the  pteropodids,  the  mouse-tailed  bats  possess  a  morphometrically  unique 
and  interesting  wing  (Fig.  7).  Whereas  the  pteropodids  are  in  a  generally  peripheral 
location  relative  to  the  variable  vector  for  head  and  body  length  (A),  the  rhinopo- 
matids  receive  nearly  the  full  negative  (shortness)  force  of  this  vector  in  the 
ordination  of  their  centroid.  This  vector  is  discriminatory  in  all  three  canonical 
axes  (Figs.  4D-E,  5,  6).  The  variable  vector  for  the  length  of  the  forearm  (B)  is 
closely  aligned  with  that  for  head  and  body  length,  but  the  ordinating  effect  of  this 
vector  is  more  direct  and  positive  (large-sized)  rather  than  negative,  and  its 


252 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


influence  in  discrimination  is  more  important  (Table  6).  The  relative  length  of 
the  forearm  approaches  unity  (0.94;  Table  A8),  which  may  or  may  not  reflect 
an  interactive  relationship  between  these  two  variables.  It  is  interesting  to  note 
that  this  relationship  is  maintained  through  the  sixth  canonical  axis  (Table  4), 
although  the  position  of  the  centroid  shifts  slightly  into  a  more  peripheral 
location. 

A  much  more  complex,  extra-dimensional  interaction  exists  for  the  combined 
variation  of  the  metacarpal  elements  of  digits  III-V  (C,  G,  J)  (Figs.  4D-E,  5, 
6;  Table  4).  In  the  first  three  canonical  axes,  and  similar  to  the  vector  for  the 
forearm  (B),  the  vector  for  the  fifth  metacarpal  (J)  is  a  strong  positive  discriminator 
(Table  6).  This  manal  element  comprises  68.10  per  cent  of  the  total  length  of 
the  fifth  digit  (Table  A 19),  and  this  percentage  is  exceeded  only  by  some  ves- 
pertilionids  and  Noctilio.  The  tail  (shortness)  of  the  variable  vector  for  the  length 
of  the  fourth  metacarpal  (G)  also  is  directed  toward  the  centroid  for  rhinopomatids 
in  the  first  two  canonical  axes.  However,  in  the  third  canonical  axis,  the  tail  of  this 
vector  is  directed  upward,  and,  although  it  seems  to  interact  synergistically  with 
other  vectors  to  carry  some  centroids  aloft,  its  effect  seems  minimal  in  this  regard 
to  the  rhinopomatids.  This  is  interesting  in  light  of  the  apparent  importance  of 
this  variable  in  the  discrimination  of  the  group  (Table  6).  In  Table  4,  it  will  be 
noted  that  the  cumulative  percentage  of  the  variance  contributed  by  the  fourth 
metacarpal  to  the  first  three  canonical  axes  is  low — 20.57  (1.66,  18.52,  and  0.39, 
respectively),  whereas  the  percentage  contributed  in  the  fourth  axis  of  discriminant 
space  is  markedly  increased — 51.55  (72.12  cumulative  per  cent).  In  addition, 
this  vector  becomes  a  strong  discriminator  for  shortness  of  the  fourth  metacarpal 
and  is  again  oriented  more  directly  toward  the  centroid  of  the  rhinopomatids. 
This  metacarpal  contributes  59.77  per  cent  to  the  total  length  of  the  fourth  digit 
(Table  A16).  The  length  of  the  third  metacarpal  of  rhinopomatids  is  particularly 
striking  (61 .70  per  cent  of  the  length  of  digit  III)  compared  to  that  of  other  bats. 
In  Table  A12,  it  is  exceeded  only  by  the  emballonurid  Depanycteris  (63.21). 
However,  the  influence  of  this  variable  on  the  dispersion  in  the  first  three  canonical 
axes  is  not  readily  apparent  (Figs.  4D-E,  5,  6).  An  examination  of  Table  4  will 
show  that  there  exists  an  extradimensional  effect  similar  to  that  described  for  the 
fourth  metacarpal.  Whereas  the  orientation  of  this  variable  vector  is  oblique 
to  the  rhinopomatid  centroid  in  the  first  three  canonical  axes,  its  point  (longness) 
directly  ordinates  this  group  in  the  fourth  through  sixth  dimensions  of  dis¬ 
criminant  space. 

The  vectors  for  the  lengths  of  the  first  and  second  phalanges  of  digit  III  (D,  E) 
generally  indicate  small  size,  although  the  interaction  between  these  variables 
results  in  vectors  that  tangentially  effect  the  centroid  for  rhinopomatids.  Again, 
this  effect  becomes  more  direct  in  extradimensional  space.  The  variable  vectors 
for  these  two  phalanges  of  the  fourth  digit  (H,  I)  directly  indicate  shortness  in 
the  first  three  canonical  axes.  The  effect  is  strongest  for  the  distalmost  member  (I) 
of  this  pair  of  phalanges.  Table  6  indicates  a  rather  minor  role  for  the  first  and 
second  phalanges  of  digit  V  (K,  L)  in  the  discrimination  of  the  rhinopomatids. 
Nonetheless,  the  vector  for  the  proximal  member  of  this  pair  (K)  is  oriented 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


253 


24  9 

20.2  22.0  22.0 
6.6  3  6  5  3 

196  10. 1  6.3 


Fig.  7. — Diagrammatic  representation  of  the  wing  construct  based  on  the  mean  lengths  of 
variables  for  the  Pteropodidae,  Emballonuroidea,  and  Rhinolophoidea.  Columns  of  numbers 
associated  with  each  construct  are,  from  left  to  right:  length  of  forearm,  metacarpal,  and 
phalanges  of  digit  III;  length  of  metacarpal  and  phalanges  of  digit  IV;  and  length  of  meta¬ 
carpal  and  phalanges  of  digit  V.  Digit  IV  is  projected  at  the  mean  alpha  angle  computed  for 
each  taxon  (see  Table  Al). 


positively  toward  the  group  centroid  in  the  first  two  canonical  axes  (Figs.  4D,  6). 
In  the  third  canonical  axis,  the  effect  of  this  variable  is  reduced.  Similarly,  the 
vector  for  the  terminal  phalanx  of  digit  V  (L)  is  oriented  toward  the  centroid  for 
rhinopomatids,  but  in  the  third  dimension  this  vector  stands  far  above  the 
centroid.  This  further  indicates  the  complexity  of  the  variation  and  interactive 
associations  among  variables. 

The  overall  effect  of  the  interplay  among  the  wing  elements  of  rhinopomatids 
is  to  produce  a  wing  with  a  below  average  overall  aspect  ratio  of  5.57  (Figs.  7, 
10;  Table  A3).  Findley  et  al.  (1972)  noted  the  shortness  of  the  wing  tip  and 
indicated  that  rhinopomatids  had  the  lowest  tip  index  of  all  bats  examined  by 
them.  We  computed  an  average  tip  index  of  1 .09,  which  agrees  with  1.19  reported 
by  these  authors  for  Rhinopoma  hardwickei.  In  addition,  they  commented  on 
the  relatively  short  wings  possessed  by  these  bats.  Although  these  indices  and 
ratios  provide  a  vague  impression  of  the  rhinopomatid  wing,  they  do  not  clearly 
delineate  the  uniqueness  of  its  shape  or  the  causative  aspects  of  this  shape. 

The  shortness  of  digits  III  to  V  is  most  greatly  effected  by  short  phalangeal 
elements  and  a  relatively  short  fourth  metacarpal;  the  third  and  fifth  metacarpals 
are  among  the  longest  for  all  bats  (Tables  A 16,  A 19).  As  noted  above, 


254 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


rhinopomatids  possess  nearly  the  longest  forearm  relative  to  their  head  and  body 
length  (the  relationship  is  almost  1:1).  Whereas  pteropodids,  and  to  a  greater 
extent  molossids,  have  much  higher  tip  indices,  the  relative  length  of  digit  III 
for  both  is  only  slightly  higher  than  that  shown  for  rhinopomatids  (Tables  A2, 
A9).  The  long  forearm,  in  combination  with  a  relatively  short  fifth  digit  (0.92 
as  compared  to  1.00  for  molossids)  produces  a  plagiopatagium  with  an  extremely 
high  aspect  ratio  (2.16)  (Figs.  7,  10;  Table  A5).  This  mean  value  is  the  largest 
among  bats  and  is  exceeded  in  range  only  by  molossids  and  emballonurids. 

According  to  Harrison  (1964:62),  the  flight  of  Rhinopoma  hardwickei  is 
peculiar  and  distinctive,  consisting  of  a  “series  of  alternating  flutters  and  glides, 
with  a  rising  and  falling  motion.  .  .  .”  Dr.  Gamal  Madkour,  who  is  familiar 
with  R.  microphylum  of  Egypt,  indicated  to  us  (personal  communication)  that 
these  bats  are  rather  swift-flyers  that  forage  in  open  country.  In  view  of  these 
apparent  conflicting  observations,  we  hesitate  to  comment  on  the  functionality 
of  the  wing  of  rhinopomatids  other  than  to  say  that  it  should  be  capable  of 
producing  moderate  speed  as  well  as  maneuverability.  We  see  little  basis  for  a 
close  functional  relationship  between  the  Rhinopomatidae  and  the  rhinolophoids, 
the  wings  of  which  are  constructed  differently.  Finally,  mouse-tailed  bats  share 
the  closest  resemblance  with  the  family  Emballonuridae — generalized  (not 
taxonomic)  distance  6.94.  This  resemblance  is  founded  on  similarity  of  variable 
vectors  for  the  lengths  of  the  forearm,  first  phalanx  of  digit  IV,  second  phalanx 
of  digits  IV  and  V,  and  the  fifth  metacarpal. 

Craseonycteridae 

Craseonycteris  thonglongyai  represents  the  small  extreme  in  the  size  variation 
among  the  Chiroptera.  These  bats,  recently  described  as  a  monotypic  family 
(Hill,  1974),  can  truly  be  thought  of  as  “bumble-bee  bats,”  as  they  are  scarcely 
larger  than  their  hymenopteran  namesake.  Because  of  their  extreme  small  size, 
we  can  reemphasize  the  rather  minor  effect  that  general  size  has  on  ordination 
in  discriminant  space.  In  the  principal  components  analysis  (Fig.  3),  this  family 
was  strongly  ordinated  along  the  first  component  axis  by  the  tail  (smallness)  of  the 
variable  vector  for  head  and  body  length  (A).  As  we  noted,  the  pteropodids  were 
ordinated  in  the  opposite  direction  and  the  remaining  taxa  disperse  between 
these  two  extremes.  In  terms  of  distance  coefficients,  this  spread  (PCA)  constitutes 
a  taxonomic  (Euclidean)  distance  value  of  131.18.  Similarly  large  taxonomic 
distances  were  computed  between  pteropodids  and  other  small-sized  taxa  such 
as  the  Natalidae  and  Thyropteridae  (105.1 1  and  1 14. 12,  respectively).  However, 
in  discriminant  space,  these  general  size  effects  are  markedly  moderated  and  the 
generalized  distances  between  pteropodids  and  these  three  small-sized  taxa 
(9.23,  6.43,  and  6.72,  respectively)  are  suggestive  of  shape  rather  than  size 
differences.  Whereas  the  vectors  for  nearly  all  variables  effect  the  ordination  of 
the  pteropodids  by  pushing  them  away  from  the  centroid  of  the  craseonycterids, 
that  for  head  and  body  length  contributes  the  least  percentage  to  the  group  dis¬ 
crimination  vector  of  the  latter  (0.36,  Table  6).  Therefore,  again  we  see  that  in 


Table  6. — Percentage  contributed  by  each  direction  cosine  to  the  discriminant  function  of  each  group.  These  values  may  be  compared  with  the 
canonical  coefficients  (Table  4)  in  an  interpretation  of  the  overall  effect  a  variable  or  set  of  variables  has  on  the  ordination  of  a  group(s). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


255 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


discriminant  space  the  quantitative  effects  of  general  size  are  much  reduced  in 
lieu  of  the  more  complicated  synergistic  interactions  among  variables  that  reflect 
the  qualitative  aspects  of  size. 

The  variable  vector  for  the  length  of  the  forearm  (B)  appears  to  have  a  mod¬ 
erately  strong  influence  on  the  ordination  of  the  craseonycterids  (Figs.  4D-E,  5, 
6;  Tables  4,  6).  Although  this  variable  is  the  shortest,  in  terms  of  absolute  length, 
among  all  bats,  the  vector  suggests  longness  of  the  forearm.  In  Table  A8,  the 
mean  relative  length  of  the  forearm  (0.82)  is  somewhat  larger  than  that  of  all  bats 
(0.73),  further  substantiating  the  interpretation  of  this  vector.  The  combined 
effect  of  the  lengths  of  digit  III  and  the  forearm  is  the  production  of  a  relatively 
long  wing  for  the  craseonycterids  (Tables  A2,  A7). 

Hill  (1974),  in  his  detailed  comparison  of  the  structure  of  the  wing  of 
Craseonycteris  with  those  of  other  bats,  noted  a  rather  peculiar  variation  among 
the  metacarpal  elements.  The  third  metacarpal  of  Craseonycteris  is  relatively 
short  as  compared  to  the  fourth  and  fifth,  which  are  somewhat  longer  and  approxi¬ 
mately  equal  in  length.  The  relationship  of  the  vector  for  the  length  of  the  third 
metacarpal  (C)  to  the  centroid  of  craseonycterids  is  similar  to  that  discussed  for 
pteropodids.  The  contribution  of  this  element  to  the  length  of  digit  III  (43.44, 
Table  A 12)  is  below  the  average  of  other  bats.  The  qualitative  shortness  of  the 
fourth  metacarpal  is  suggested  by  the  vector  for  this  variable  (G)  in  the  first 
and  second  canonical  axes  (Figs.  4D,  6).  In  the  third  axis  (Figs.  4E,  5),  the  group 
centroid  is  located  somewhat  to  the  side  of  this  variable  vector,  although  the 
implication  of  shortness  persists.  The  relationship  of  the  variable  vector  for 
the  fifth  metacarpal  (J),  in  all  three  axes,  implies  longness.  The  contribution  of 
the  fourth  and  fifth  metacarpal  elements  to  the  lengths  of  their  respective  digits 
is  above  average  for  all  bats  (Tables  A 16,  A 19). 

Perhaps  the  most  striking  feature  of  the  third  digit  is  the  relatively  long  second 
phalanx  (Fig.  7).  This  phalanx  is  nearly  equal  to  the  metacarpal  in  length  (Tables 
A 12,  A 14)  and  its  contribution  to  the  length  of  the  digit  is  largest  among  all  bats. 
Although  the  percentage  contributed  to  the  discrimination  vector  of  the  group 
(12.42,  Table  6)  is  relatively  high,  the  implication  of  this  variable  vector  (E)  in 
the  first  three  canonical  axes  (Figs.  4D-E,  5,  6)  is  toward  shortness.  However, 
in  the  fourth  and  fifth  canonical  axes,  the  positive  end  (longness)  of  the  vector 
is  strongly  oriented  toward  the  centroid  of  craseonycterids.  Again,  this  em¬ 
phasizes  the  multidimensional  and  synergistic  nature  of  the  interaction  among 
variables  on  alar  shape. 

A  similar  relationship  for  the  distal  phalanx  of  digit  III  exists  for  the 
rhinolophids,  megadermatids,  and,  to  a  lesser  extent,  nycterids.  The  actual 
structure  of  the  tip  portion  of  the  wing  in  these  bats  is  rather  curious  and  is  not 
found  in  any  other  group.  The  middle  and  distal  portion  of  the  shaft  of  the  second 
phalanx  of  digit  III  is  arched  in  such  a  way  as  to  trap,  and  maintain  taut,  a  small 
section  of  the  alar  membrane  in  much  the  same  fashion  as  the  string  of  a  bow. 
The  joint  between  the  distal  phalanx  and  the  first  phalanx  of  digit  III  is  broad, 
and  there  appears  to  be  a  great  deal  of  mobility  at  this  joint,  judging  from  specimens 
preserved  in  alcohol.  Although  we  are  not  prepared  to  discuss  the  functional 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


257 


Fig.  8. — Diagrammatic  representation  of  the  wing  construct  based  on  the  mean  lengths  of 
variables  for  the  Phyllostomatoidea.  Columns  of  numbers  associated  with  each  construct  are, 
from  left  to  right:  length  of  forearm,  metacarpal,  and  phalanges  of  digit  III;  length  of  meta¬ 
carpal  and  phalanges  of  digit  IV;  and  length  of  metacarpal  and  phalanges  of  digit  V.  Digit  IV 
is  projected  at  the  mean  alpha  angle  computed  for  each  taxon  (see  Table  A1 ). 


ramifications  of  this  anatomical  configuration,  we  suggest  that  the  apparent 
emphasis  in  the  ordination  of  these  families  by  this  feature  implies  not  only 
similarity  in  shape,  but  also  functional  similarity.  Perhaps  it  is  employed  during 
the  “flick  phase”  of  the  wing  beat  cycle,  or  it  simply  may  be  a  device  for  furling 
this  long  wing  element.  Although  the  phylogenetic  sources  of  shape  are  not  our 
primary  goal  in  this  paper,  we  would  point  out  that  this  feature  suggests  a  close 
relationship  among  these  families.  The  emballonurids  possess  a  slightly  different 
folding  device  in  this  distal  region  of  their  wings,  and  the  rhinopomatids,  which 
lack  this  feature,  might  represent  the  underived  (primitive)  condition  for  this 
characteristic. 

Of  all  the  variables  employed  in  this  study,  the  length  of  the  first  phalanx  of 
digit  IV  appears  to  be  the  most  distinctive  of  Craseonycteris  (Table  6).  This  wing 
element  is  extremely  short  and  constitutes  only  10.2  per  cent  of  the  total  length 
of  the  fourth  digit  and,  in  a  relative  sense,  is  the  shortest  observed  in  all  bats  (Table 
A 17).  The  shortness  of  this  wing  element  is  emphasized  in  the  discriminant 
analysis  by  the  variable  vector  (H)  in  the  first  three  canonical  axes  (Figs.  4D-E, 


258 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


5,  6;  Table  4).  This  vector  is  involved  similarly,  but  to  a  slightly  lesser  degree, 
in  the  ordination  of  the  Rhinolophidae,  Nycteridae,  Megadermatidae,  Noctilioni- 
dae,  and  Natalidae.  The  second  phalanx  of  digit  IV  is  long  and  is  second  in  size 
only  to  that  of  the  Noctilionidae  (Table  A 18).  However,  the  interactive  re¬ 
lationship  of  this  variable  is  obscured  by  the  synergistic  complexity  among  all 
variables. 

The  variable  vectors  for  both  phalangeal  elements  of  the  fifth  digit  (K,  L) 
also  are  difficult  to  interpret,  although  they  indicate  longness  in  the  first  three 
canonical  axes.  Of  the  two  variables,  the  length  of  the  first  phalanx  of  this  digit 
appears  to  be  the  most  influential  in  the  discrimination  of  the  group  (Table  6). 
The  precentages  contributed  to  the  length  of  digit  V  by  the  first  and  second 
phalanges  (15.79  and  18.64,  respectively)  are  below  the  average  for  all  bats 
(Tables  A20,  A21). 

The  overall  aspect  ratio  of  the  wing  of  Craseonycteris  thonglongyai  is  slightly 
below  the  mean  for  all  bats  (5.64,  Table  A3  and  Fig.  10).  The  aspect  of  the 
plagiopatagial  portion  is  not  particularly  distinctive  (1.48)  and  falls  in  the  middle 
to  lower  range  for  all  bats  (Table  A5).  In  addition,  the  aspect  ratio  of  the  tip  portion 
of  the  wing  is  approximately  equal  to  the  average  for  all  bats  (Table  A4  and  Fig. 
10).  On  the  other  hand,  the  third  digit  is  1.86  times  as  long  as  the  forearm,  which 
is  generally  high  compared  to  that  of  other  bats  (Table  A2;  Fig.  10). 

The  overall  shape  of  the  craseonycterid  wing  is  the  result  of  a  rather  unusual 
combination  of  interactions  among  the  various  wing  elements.  The  length  of 
the  third  digit  appears  to  be  most  strongly  influenced  by  the  length  of  the  distal 
phalanx,  which  tends  to  offset  the  shortness  of  the  metacarpal.  In  the  fourth  digit, 
the  relatively  long  metacarpal  and  distal  phalanx  appear  to  compensate  for  the 
markedly  shortened  first  phalanx.  The  fifth  digit  is  relatively  long,  owing  to  a 
generally  isometric  association  with  the  metacarpal  and  second  phalanx  of 
digit  IV,  and  tends  to  offset  the  length  of  the  third  digit.  These  interactions  thereby 
contribute  to  the  generally  broad  aspect  of  the  wing  tip. 

Prompted  by  comments  made  by  Findley  et  al.  (1972)  concerning  an  average 
or  below  average  aspect  ratio  coupled  with  a  high  tip  index,  Hill  (1974)  suggested 
a  hovering  ability  for  these  small  bats.  We  agree  that  Craseonycteris  may  possess 
this  flight  potential,  but  our  basis  for  this  assumption  lies  more  with  the  structural 
nature  of  the  third  digit,  especially  the  long  distal  phalanx,  rather  than  with  the 
relationship  between  aspect  ratio  and  tip  index. 

Emballonurid.ae 

From  the  standpoint  of  wing  diversity,  the  emballonurids  represent  one  of  the 
most  intriguing  families  of  bats.  In  terms  of  aspect  ratios,  they  range  from  slightly 
above  average  (6.05)  for  the  order  to  extremely  high  aspect  ratios  (7.93).  Their 
forearms  may  be  relatively  short  to  long  and,  as  a  consequence,  the  tip  indices 
for  members  of  the  family  also  vary  from  low  to  high.  In  these  general  descriptive 
terms,  the  wings  of  emballonurids  most  closely  resemble  those  of  bats  of  the  family 
Molossidae  and,  in  some  respects  the  Noctilionidae  and  Mormoopidae.  However, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


259 


this  resemblance  is  merely  superficial  as  these  families  acquire  their  extreme 
wing  shapes  through  different  morphometric  modes.  To  draw  attention  to  this 
misleading  resemblance,  we  will  draw  comparisons  between  the  emballonurids, 
molossids,  and  noctilionids  in  this  account.  The  group  centroids  of  these  three 
families  are  located  in  separate  regions  of  discriminant  space  (Figs.  5,  6). 

Emballonurids  are  about  average  for  bats  in  length  of  the  head  and  body.  The 
vector  for  this  variable  (A)  is  a  minor  force  in  the  overall  discrimination  of  the 
group  (Table  6).  Head  and  body  length  has  a  slightly  stronger  effect  in  the 
ordination  of  the  Molossidae;  this  is  particularly  true  in  the  first  and  second 
canonical  axes  (Figs.  4D,  6). 

Length  of  forearm  appears  to  be  a  moderately  important  variable  in  the  ordi¬ 
nation  of  the  emballonurid  centroid.  This  appears  to  be  a  general  feature  of 
those  bats  referred  to  the  superfamilies  Emballonuroidea  and  Rhinolophoidea, 
which  are  generally  characterized  by  possessing  relatively  long  forearms  (Table 
A8).  Within  the  Emballonuridae,  the  mean,  relative  length  of  the  forearm  ap¬ 
proaches  unity  (0.93).  Although  most  species  range  below  this  value, 
the  exceptions  are  notable:  Centronycteris  maximiliani  (1.14);  Cyttarops 
alecto  (1.10);  Emballonura  solomonis  (1.11);  E.  beccarii  (1.06);  and  Cormura 
brevirostris  (1.04).  The  vector  for  the  length  of  the  forearm  (B)  contributes  a 
moderately  low  percentage  (3.15)  to  the  discrimination  vector  of  the  emballonurids 
(Table  6).  By  comparison,  the  ordination  of  the  molossids  is  more  strongly  in¬ 
fluenced  by  the  tail  (shortness)  end  of  this  vector.  This  emphasis  on  short 
length  of  the  forearm  is  reflected  in  the  higher  percentage  contributed  by  this 
vector  (9.28)  to  the  group  discrimination  vector  of  molossids  (Table  6).  There¬ 
fore,  although  the  absolute  length  of  the  forearm  in  these  two  groups  is  outwardly 
similar,  there  is  a  fundamental  difference  in  their  respective  contribution  to  the 
shape  of  the  wing  (Table  A8). 

The  variation  of  the  dactylopatagial  portion  depicts  even  more  striking 
differences  in  the  wing  construction  of  emballonurids  and  molossids  (Figs.  7,  9). 
On  the  whole,  the  length  of  the  third  digit  of  emballonurids  is  not  particularly 
impressive.  The  mean  tip  index  (1.61)  is  well  below  the  average  for  all  bats 
(Table  A2).  Centronycteris,  Saccopteryx,  and  several  species  of  Taphozous, 
especially  T.  peli,  have  unusually  large  tip  indices  (1.70-1.90).  On  the  contrary, 
molossids  generally  are  characterized  by  larger  than  average  tip  length  (Table 
A2). 

The  vectors  for  the  various  elements  of  digit  III  (C,  D,  E)  are  involved  in  the 
overall  complex  synergism  among  variables  and  their  effect  is  not  easily  inter¬ 
preted.  In  the  first  canonical  axis  (Figs.  4D-E,  5,  6),  only  the  vector  for  the  first 
phalanx  (D)  exerts  a  positive  force  on  the  ordination  of  emballonurids  (Tables  4, 
5);  shortness  is  emphasized  by  the  other  vectors  for  this  digit.  The  converse  of 
these  actions  is  implied  for  the  ordination  of  the  Molossidae  with  respect  to  the 
vectors  associated  with  digit  III.  Also,  the  centroid  for  the  Noctilionidae  is 
closely  associated  with  that  of  the  Molossidae  in  this  canonical  axis. 

Ordination  along  the  second  canonical  axis  illustrates  a  somewhat  different 
picture  (Tables  4,  5).  Here  the  vectors  for  the  metacarpal  and  second  phalanx 


260 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


39.2 

35.9  35.4  34.2 
154  9.3  9.5 
20.6  9.4  10.0 


34.8 

33.7  33. 1  30.0 
14.6  9.3  7.9 
9.5  6.0  7.5 
5.0 


42.4 

39.6  38  5  36.6 
15.0  12.0  9.4 
144  10.2  7.6 

6.9 


42  9 

40.1  38.9  35.2 
108  9.9  6.0 

13.2  13.0  I  I.  I 
I  I.  I 


Fig.  9. — Diagrammatic  representation  of  the  wing  construct  based  on  the  mean  lengths  of 
variables  for  the  Vespertilionoidea.  Columns  of  numbers  associated  with  each  construct  are, 
from  left  to  right:  length  of  forearm,  metacarpal,  and  phalanges  of  digit  III;  length  of 
metacarpal  and  phalanges  of  digit  IV;  and  length  of  metacarpal  and  phalanges  of  digit  V. 
Digit  IV  is  projected  at  the  mean  alpha  angle  computed  for  each  taxon  (see  Table  Al). 


(C,  E)  exert  a  positive  force  and  that  for  the  first  phalanx  a  negative  effect  on  the 
ordination  of  emballonurids.  Again,  the  molossids  are  ordinated  in  an  opposite 
manner.  Interestingly,  the  centroid  for  the  noctilionids  is  not  carried  in  association 
with  the  molossids,  but  is  maintained  in  its  same  relative  position  in  discriminant 
space.  As  will  be  noted  later,  the  vectors  for  elements  of  the  third  digit  are  more 
directly  involved  in  the  ordination  of  noctilionids. 

In  the  third  canonical  axis,  the  vectors  for  elements  of  digit  III  appear  to 
be  less  important  in  the  overall  ordination  of  these  three  centroids.  In  this  axis, 
vectors  for  the  fourth  and  fifth  digits  are  emphasized  in  a  relative  sense. 

As  stated  above,  variation  of  the  third  digit  is  difficult  to  describe  because  of  its 
involvement  in  the  complex  synergistic  interactions  among  variables.  However, 
the  net  effect  is  a  relatively  long  digit  (Table  A9).  The  metacarpal  is  particularly 
important  in  this  regard,  judging  from  the  high  percentage  contributed  to  the 
discrimination  vector  of  emballonurids  (22.96,  Table  6).  The  combined  effect 
of  a  long  digit  III  and  forearm  is  the  production  of  a  relatively  long  wing  as  can  be 
seen  in  Table  A7.  In  fact,  the  high  extremes  in  the  range  of  variation  are  note¬ 
worthy.  The  relative  length  of  the  wing  of  Centronycteris  is  nearly  three  and  a 
half  times  (3.34)  longer  than  the  head  and  body  length,  which  greatly  exceeds 
that  for  all  bats.  Likewise,  Cyttarops  exhibits  an  unusually  long  wing  (2.91)  as 
compared  to  other  chiropterans.  These  two  species  also  fall  at  the  high  extreme 
for  relative  length  of  digit  III  (Table  A9). 

Whereas  the  length  of  the  third  digit  is  important  in  the  overall  length  of  the 
wing,  the  lengths  of  the  fourth  and  fifth  digits  combine  to  determine  the  overall 
aspect  of  the  dactylopatagium.  We  have  noted  that  in  the  rhinopomatids  and 
craseonycterids  the  length  of  digit  III  is  generally  offset  by  a  relatively  long 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


261 


fourth  and  fifth  digit  for  the  overall  production  of  a  short,  broad  tip.  In  the  for¬ 
mation  of  high  aspect  tips,  the  trend  is  toward  a  relatively  long  fourth  digit  and 
a  shortened  fifth  digit.  The  emballonurids,  noctilionids,  and  molossids  generally 
follow  this  trend,  although  the  manner  in  which  each  responds  is  somethat  dif¬ 
ferent. 

The  vectors  for  elements  of  the  fourth  digit  (G,  H,  I)  of  emballonurids  defy 
easy  interpretation  because  of  their  overall  interaction  with  other  variables. 
In  the  first  two  canonical  axes,  these  vectors  imply  shortness  of  the  fourth  digit. 
However,  in  the  third  axis,  a  longish  fourth  metacarpal  is  suggested.  The  reader 
will  recall  that  the  elements  of  the  fourth  digit  are  not  particularly  strong  factors 
in  the  ordination  in  the  first  three  canonical  axes,  but  that  they  gain  strength  in 
the  extradimensional  fourth  through  sixth  axes.  In  the  fourth  and  fifth  axes 
(Table  4),  the  vector  for  the  fourth  metacarpal  (G)  is  strong  in  its  effect  on  the 
ordination  of  the  emballonurids  and  suggests  a  relatively  long  length  for  this 
element.  The  contribution  by  this  vector  to  the  discrimination  of  the  group  also 
is  high  (17.86,  Table  6).  A  similar  implication  applies  to  the  molossids,  but  to  a 
lesser  extent — 9.90  per  cent  contributed  to  the  function.  This  variable  appears 
to  have  only  a  minor  role  in  the  discrimination  of  noctilionids. 

The  vectors  for  the  respective  lengths  of  the  two  phalanges  of  digit  IV  do  not 
appear  to  be  important  in  the  overall  ordination  of  the  emballonurids.  The  general 
implication  is  toward  small  size  (Figs.  4D-E,  5,  6).  However,  the  position  of  the 
centroid  relative  to  these  two  canonical  vectors  suggests  a  null  effect,  or  at  least 
no  significant  elongation,  when  compared  to  the  grand  centroid  for  all 
bats.  The  ordination  of  both  the  molossids  and  noctilionids  are  effected  by  one  or 
the  other  of  these  vectors.  In  the  case  of  the  molossids,  a  long  first  phalanx  of 
digit  IV  is  emphasized,  whereas  a  long  second  phalanx,  in  combination  with  a 
short  first  phalanx,  is  suggested  for  the  Noctilionidae. 

The  length  of  the  fifth  digit  of  emballonurids,  as  well  as  that  of  noctilionids 
and  molossids,  is  relatively  short  as  compared  to  the  total  length  of  digits  III  and 
IV,  forearm,  and  head  and  body  (Tables  A7-A1 1).  In  a  general  sense,  molossids 
represent  the  extreme  of  this  variation.  The  most  striking  differences  among  these 
three  groups  is  in  the  composition  of  this  digit  and  specifically  in  the  relative 
length  of  the  metacarpal  element  (Table  A19).  The  vector  for  this  wing  element 
(J)  is  directly  involved  in  the  ordination  of  the  emballonurids  and  molossids, 
and,  to  a  lesser  extent,  noctilionids  (Figs.  4D-E,  5,  6).  The  percentage 
contributed  by  this  vector  to  the  discrimination  of  each  of  these  groups  is 
9.88,  4.47,  and  4.36,  respectively  (Table  6).  This  vector  implies  large  size  with 
respect  to  this  variable  for  emballonurids  and  noctilionids,  but  suggests  small 
size  for  molossids.  The  most  important  feature  of  the  fifth  digit  of  emballonurids 
is  a  relatively  long  proximal  phalanx  (Table  6).  This  phalanx  contributes  nearly 
a  quarter  of  the  total  length  of  digit  V  (Table  A20).  Similarly,  this  phalanx  is 
distinguished  as  long  in  the  noctilionids,  but  the  importance  in  discrimination 
of  the  group  is  slightly  reduced  (Table  6).  The  molossids,  more  than  either  of 
these  two  groups,  emphasize  the  length  of  the  first  phalanx  of  digit  V  (Table  6). 
On  the  average,  almost  30  per  cent  of  the  total  length  of  the  fifth  digit  is  reflected 


262 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


by  the  first  phalanx,  the  largest  contribution  noted  among  all  bats  (Table  A20). 
Cheiromeles  and  Otomops  (both  molossids)  represent  the  high  extremes  with 
39.13  and  35.30  per  cent,  respectively.  The  distalmost  phalanx  of  the  fifth  digit 
is  markedly  shortened  in  the  emballonurids  and  molossids.  The  significance  of 
this  reduction,  in  the  overall  ordination  of  these  two  families,  is  strongest  for  the 
molossids  compared  to  emballonurids  (Table  6).  On  the  other  hand,  this  wing 
element  is  markedly  elongated  in  the  Noctilionidae  (Fig.  8),  and  the  vector  for 
this  variable  (L)  contributes  31.67  per  cent  to  the  discrimination  vector  of  this 
group.  It  is  noteworthy  to  point  out  that  this  is  the  largest  contribution  by  any 
variable  vector  to  the  group  discrimination  vector  of  any  group. 

Thus  we  have  seen  that  emballonurids  possess  wings  that  may  be  characterized 
as  relatively  long  and  narrow  (Fig.  7).  An  overall  aspect  ratio  generally  would 
reflect  this  shape  (Table  A3),  but  would  reveal  little  in  terms  of  the  composition 
and  interaction  among  the  variables  that  produce  such  a  shape.  Outwardly,  the 
short  tip  index,  relatively  long  wing,  and  low  wing  loading  tend  to  confuse  any 
univariate  or  bivariate  interpretation  of  this  shape  (Findley  et  ai,  1972).  The 
multivariate  approach  does  help  to  clarify  the  issue.  The  wings  of  emballonurids 
are  truly  high  aspect  in  nature.  However,  a  functional  interpretation  of  this  wing 
shape  is  liable  to  be  confounded  if  the  wings  of  emballonurids  are  compared  to 
the  high  aspect  wings  of  molossids.  In  such  a  comparison,  one  is  likely  to  be  biased 
and  misled  by  the  apparent  high  correlation  between  high  aspect  ratio  and 
swiftness  of  flight,  both  attributes  of  molossids.  In  addition,  generally  high  wing 
loading  appears  to  accompany  the  high  aspect  ratio  of  the  molossids  and  not  that 
of  emballonurids  (Table  A6). 

We  have  shown  that  the  construction  of  the  wings  of  emballonurids  differs 
greatly  from  those  of  molossids  and  noctilionids,  albeit  the  end  product  is  vaguely 
similar.  Emballonurids  appear  to  have  modified  a  fundamentally  short  tip  into  a 
long,  high  aspect  tip  by  maintaining  relatively  long  metacarpal  elements  and 
elongating  the  terminal  phalanx  of  digit  III;  the  distalmost  phalanges  of  digits 
IV  and  V  appear  to  be  shortened.  The  development  of  a  high  aspect  wing  in  this 
manner  may  avoid  allometric  complexities  associated  with  the  modification  of 
more  proximal  wing  elements.  In  addition,  to  achieve  a  high  aspect  wing,  such 
modifications  might  allow  greater  versatility.  The  highly  maneuverable  flight  of 
emballonurids  is  suggestive  of  a  wide  range  of  flight  potentials.  Some  species 
(notably  those  of  Taphozous,  Emballonura,  Diclidurinae,  and  perhaps  Centro- 
nycteris )  appear  to  have  capitalized  on  the  speed  qualities  of  high  aspect  wings. 

Rhinolophidae 

Horseshoe  bats  possess  wings  that  average  the  lowest  in  overall  aspect  ratio 
(5.41)  as  compared  to  all  other  bats  (Table  A3,  Fig.  7).  The  length  of  the  third 
digit  averages  only  slightly  longer  than  the  head  and  body  (1.28).  Also,  the  forearm 
nearly  equals  the  length  of  head  and  body  (Table  A8).  These  attributes  combine 
to  produce  a  wing  with  next  to  the  lowest  average  tip  index  (1.39)  for  all  bats 
(Table  A2  and  Fig.  10);  only  the  rhinopomatids  average  lower. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


263 


Fig.  10. — Bivariate  graphs  that  illustrate  the  relationships  between  tip  index  and  three 
aspect  ratios  of  the  wing.  Triangles  represent  phyllostomatid  centroids  and  the  grand  centroid 
for  phyllostomatids  is  indicated  by  a  circle  with  a  plus.  See  list  of  specimens  examined  or 
Table  5  for  key  to  alphabetic  code  and  text  for  discussion. 

The  synergistic  relationships  among  the  raw  variables,  discussed  above  for 
the  rhinopomatids  and  emballonurids,  generally  apply  to  the  rhinolophids.  A 
relatively  long  forearm  is  implied  by  the  vector  for  this  variable  (B).  The  ordination 
of  the  group  centroid  for  the  rhinolophids,  as  well  as  that  of  the  Nycteridae  and 
Megadermatidae,  appear  to  be  more  strongly  effected  by  vectors  associated  with 
elements  of  the  third  digit.  However,  the  relationships  are  difficult  to  characterize 
because  they  are  involved  in  a  complex  interaction  among  all  variables.  In  the 
first  two  canonical  axes,  the  implication  is  toward  shortness,  whereas  in  the  third 
axis  there  is  a  general,  but  weak,  expression  of  large  size.  Our  general  impression 
is  that  these  vectors  describe  the  shortness  of  the  digit  as  a  whole,  but  the  individual 
components  are  either  not  affected  or  show  only  slight  elongation. 

With  regard  to  the  fourth  digit,  all  variable  vectors  for  the  elements  of  this 
digit  (G,  H,  I)  imply  shortness  in  the  first  two  canonical  axes  (Figs.  4D-E,  5,  6). 
In  the  third  axis,  the  vector  for  the  fourth  metacarpal  (G)  further  emphasizes 
shortness.  However,  in  this  third  dimension  of  discriminant  space,  the  vectors 
for  both  phalanges  (H,  I)  of  digit  IV  suggest  large  size.  The  percentage  of  the 
variance  contributed  to  the  discrimination  vector  of  the  group  by  the  proximal 
element  of  this  series  is  exceptionally  high  (22.92,  Table  6). 

The  vectors  for  the  components  of  the  fifth  digit  (J,  K,  L)  are  somewhat  more 
influential  (Table  6)  and  all  imply  large  size  in  the  overall  ordination  of  the 


264 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


rhinolophids.  The  vector  for  the  distalmost  phalanx  (L)  tends  to  elevate  the 
group  centroid  in  the  third  dimension  of  discriminant  space,  but  the  combined 
effect  of  the  vectors  for  the  metacarpal  and  the  proximal  phalanx  (J,  K)  act  to 
suppress  the  elevation  of  the  centroid. 

The  wings  of  rhinolophids,  although  perhaps  not  structurally  as  striking  as 
those  of  the  emballonurids,  craseonycterids,  or  rhinopomatids,  do  agree  in 
general  structure  and  composition  with  wings  of  bats  in  these  families.  The 
generalized  distance  between  rhinolophids  and  these  other  group  centroids  is 
relatively  small — Rhinopomatidae  (7.24),  Craseonycteridae  (4.23),  and 
Emballonuridae  (5.92).  The  most  notable  difference  between  the  wing  of  rhino- 
lophoids  and  that  of  emballonuroids,  and  a  feature  that  appears  to  distinguish 
the  former,  is  the  short  tip  and  generally  broad  aspect.  The  composition  of  the 
wing  in  these  two  superfamilies  appears  to  be  similar  and  to  reveal  a  relatively 
close  common  ancestry. 


Nycteridae  and  Megadermatidae 

Because  of  their  close  association  in  discriminant  space,  (generalized  distance 
4.57),  we  will  discuss  these  two  families  together.  Although  the  megadermatids 
average  somewhat  larger  in  general  size  than  do  nycterids  and  rhinolophids,  all 
three  families  are  similar  in  general  wing  shape  and  composition  (Fig.  7).  The 
ordination  of  these  two  families  is  influenced  by  vectors  of  nearly  the  same  direc¬ 
tion  and  magnitude  as  discussed  in  the  preceding  account  of  the  Rhinolophidae; 
major  differences  are  mostly  quantitative  rather  than  qualitative. 

The  mean  aspect  ratio  of  the  wings  of  nycterids  and  megadermatids  is  only 
slightly  higher  than  that  of  rhinolophids  (Table  A3).  The  relative  lengths  of  digit 
III,  and  consequently  the  tip  indices  also,  are  similar  (Tables  A9,  A2).  The  agree¬ 
ment  among  these  values  further  attests  to  the  qualitative  similarity  of  wing  shape 
in  these  three  families. 

The  major  differences  between  the  wings  of  these  two  families  and  the 
Rhinolophidae  appear  to  involve  the  two  phalanges  of  digit  III.  The  nature  of 
these  quantitative  differences  is  strong  enough  to  produce  a  group  discrimination 
vector  capable  of  consistently  classifying  the  respective  members  of  each  family 
(Fig.  17). 

The  first  phalanx  of  the  third  digit  is  comparatively  longer  in  nycterids  than 
in  either  rhinolophids  or  megadermatids.  The  vector  for  this  variable  contributes 
19.33  per  cent  to  the  discrimination  of  the  group  (Table  6).  The  vectors  for  the 
third  and  fourth  metacarpal  (C,  G)  of  all  three  groups  ordinate  toward  small  size 
as  discussed  in  the  account  of  the  Rhinolophidae.  The  vector  for  the  fifth  meta¬ 
carpal  (J)  is  slightly  stronger  in  the  ordination  of  the  Nycteridae  than  it  is  in  either 
the  Rhinolophidae  or  Megadermatidae  (Table  6). 

In  the  ordination  of  the  Megadermatidae,  the  vectors  for  the  third  metacarpal 
and  second  phalanx  of  this  digit  (C,  E)  are  the  strongest  relative  to  these  three 
families  and  contribute  9.69  and  8. 17  per  cent,  respectively,  to  the  discrimination 
of  the  group.  The  vector  of  the  former  implies  shortness,  whereas  the  latter  in- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


265 


dicates  large  size.  The  combined  effect  appears  to  be  elements  of  nearly  equal 
length.  The  phalanges  of  the  fourth  digit  are  slightly  longer,  in  a  relative  sense,  and 
these  vectors,  likewise,  are  strong  contributors  to  the  discrimination  vector  of  the 
group  (27.23  and  13.12  per  cent,  respectively). 

Noctilionidae 

Many  of  the  distinguishing  features  of  the  wings  of  Noctilio  were  discussed 
in  the  account  of  the  Emballonuridae.  The  wings  of  both  Noctilio  albiventris  and 
N.  leporinus  are  essentially  alike  in  shape  even  though  they  differ  markedly  in  the 
absolute  size  of  all  raw  variables.  The  wing  of  these  two  species  are  nearly  two 
and  a  half  times  the  length  of  the  head  and  body  and  almost  65  per  cent  of  the  span 
is  composed  of  the  third  digit.  As  a  consequence,  the  tip  index  for  the  family  is 
high  for  the  order  (1.92  for  N.  albiventris  and  1.98  for  N.  leporinus).  Although 
the  overall  aspect  ratio  of  the  wing  is  high  and  similar  to  that  of  molossids  and 
emballonurids,  we  have  noted  that  the  acquisition  of  this  aspect  is  achieved  through 
different  independent  interactions  among  the  elements  that  comprise  the  wing 
in  these  three  families  (Figs.  7,  8,  9). 

All  vectors  relating  to  features  of  the  third  digit  (C,  D,  E,  F)  weigh  heavily  in 
the  ordination  of  the  group.  In  addition,  all  but  that  for  the  first  phalanx  indicate 
large  size.  The  vectors  for  the  most  proximal  phalanx  of  the  third  digit  (D)  and 
fifth  digit  (K),  as  well  as  those  for  the  fourth  and  fifth  metacarpal  (G,  J),  imply 
smallness  and  tend  to  suppress  the  ordination  of  the  group  centroid  in  the  third 
canonical  axis  (Figs.  4E,  5). 

Although  the  wings  of  Noctilio  are  high  in  aspect,  we  again  caution  comparisons 
with  the  apparent  swift  flying  ability  of  molossids.  We  have  observed  both  species 
in  the  field  and  would  note  that  N.  leporinus  flies  with  a  constant,  but  relatively 
slow  and  shallow  wing  beat.  It  does  not  appear  to  be  a  particularly  fast  flier.  The 
smaller  species,  N.  albiventris ,  is  an  insectivorous  bat  and  from  our  observations 
is  capable  of  faster  flight  judging  from  the  force  with  which  individuals  strike  a 
mist-net.  N.  albiventris  also  exhibits  a  fair  amount  of  maneuverability  in  close 
quarters  and  is  capable  of  avoiding  obstacles. 

In  our  discussion  of  the  Pteropodidae,  we  suggested  that  the  possession  of  wing 
elements  of  rather  long  span  allowed  for  the  control  of  large  portions  of  the  cam¬ 
bered  surfaces.  Slight  flexion  of  these  elements  might  greatly  affect  the  camber 
of  the  wing,  in  a  manner  similar  to  the  downward  deflection  of  the  hinged  flaps 
on  an  airplane.  This  would  contribute  markedly  to  the  lift  potential  at  low  speeds. 
We  further  suggested  that  the  nearly  equal  lengths  of  the  manal  elements  of 
pteropodids  might  allow  for  rather  crude,  yet  effective,  camber  adjustments. 
We  continue  this  argument  here  and  suggest  that  the  shortening  of  a  proximal 
phalanx,  especially  in  digits  III  and  IV,  would  allow  a  greater  range  of  variation 
as  well  as  finer  dexterous  control  of  the  camber  of  the  wing. 

With  regard  to  Noctilio ,  and  perhaps  mormoopids,  the  shortened  first 
phalanx  in  digits  III  and  IV  not  only  contributes  to  the  high  aspect  construction, 
but  might  account  for  the  apparent  versatility  of  flight  behavior.  Furthermore, 


266 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


in  wings  that  have  three  phalangeal  segments  in  the  third  digit,  this  means  of 
differential  elongation  of  elements  also  may  allow  an  increase  in  dexterity  during 
the  “flick  phase’’  of  the  wing  beat  cycle. 

Mormoopidae 

Bats  of  this  family  possess  a  relatively  long  wing,  63  per  cent  of  which  is 
contributed  by  the  third  digit  (Fig.  8).  As  we  observed  in  the  Emballonuridae,  the 
relatively  long  forearm  may  mask  or  otherwise  offset  the  length  of  the  tip.  The 
tip  index  of  mormoopids  (1.70)  is  only  slightly  higher  than  that  obtained  for 
emballonurids  and  both  values  are  well  below  average  for  all  bats.  Our  data 
suggest  that  mormoopid  wings  are  well  above  average  in  overall  aspect  ratio 
(Table  A3)  and  that  the  tip  can  hardly  be  characterized  as  short.  The  mormoopids 
appear  to  be  closest,  in  wing  morphology,  to  the  Phyllostomatidae;  misclassi- 
fication  occurred  with  the  least  derived  species,  Pteronotus  parnellii,  being 
assigned  to  the  Phyllonycterinae  (Fig.  17). 

The  length  of  head  and  body  is  a  relatively  minor  feature  in  the  discrimination 
of  mormoopids  (Table  6).  Also,  the  length  of  forearm  appears  to  have  little  effect 
on  the  overall  discrimination  of  the  group. 

The  most  important  variable  vectors  in  the  ordination  of  the  mormoopids 
appear  to  be  those  associated  with  elements  of  the  third  digit  (C,  D,  E,  F) — long 
metacarpal,  short  first  phalanx,  and  long  third  phalanx  are  emphasized  (Figs. 
4D-E,  5,  6).  The  former  two  components  of  the  mormoopid  wing  contribute  the 
most  to  the  discrimination  of  the  group  (20.56  and  26.30  per  cent,  respectively). 
Tables  A12-A15  generally  reflect  these  features.  The  percentage  contributed  to  the 
length  of  digit  III  by  the  first  phalanx  is  nearly  the  lowest  for  all  bats  (11.18), 
whereas  that  contributed  by  the  distal  phalanx  is  the  highest  (16.64).  This  appears 
to  be  a  general  phyllostomatoid  feature. 

The  effects  of  the  vectors  for  elements  of  the  fourth  digit  (G,  H,  I)  are  difficult 
to  interpret  because  of  their  apparent  involvement  in  the  overall  synergistic 
interaction  among  all  variables.  In  the  first  and  second  canonical  axes  (Figs.  4D, 
6),  the  vector  for  the  fourth  metacarpal  (G)  is  oriented  away  from  the  group 
centroid  for  the  mormoopids  and  thereby  implies  shortness.  However,  in  the  third 
axis  (Figs.  4E,  5),  this  vector  exerts  a  more  positive  force  in  the  ordination  of 
the  centroid.  Both  vectors  for  the  phalanges  of  digit  IV  (H,  I)  indicate  large  size, 
with  emphasis  on  the  distalmost  phalanx.  This  terminal  phalanx  is  not  nearly  so 
long  or  apparently  so  important  in  the  discrimination  of  the  group  as  was  observed 
in  the  Noctilionidae  (Table  6).  The  vectors  for  the  corresponding  pair  of  phalanges 
in  the  fifth  digit  (K,  L)  also  indicate  large  size  with  emphasis  on  the  proximal 
member.  These  two  phalanges  weigh  heavily  in  the  discrimination  of  the 
group  (Table  6)  and  appear  to  cause  a  lengthening  of  the  fifth  digit,  which  tends 
to  broaden  the  wing. 

Vaughan  and  Bateman  ( 1 970)  presented  an  excellent  discussion  of  the  functional 
myology  of  this  group.  They  noted  the  remarkable  maneuverability  of  these  bats 
and  their  rapid  and  sustained  flight.  Mormoops  megalophylla  is  extreme  in  nearly 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


267 


all  aspects  of  the  wing.  Mormoops  blainvillii  is  rather  curious  in  that  the  aspect 
ratio  of  its  wing  is  nearly  equal  to  that  of  the  larger-sized  species  M.  megalophylla 
(6.32),  whereas  its  wing  loading  is  a  third  lower  (4.99  Nt/m2).  Members  of  the 
genus  Pteronotus,  and  especially  P.  parnellii,  appear  to  be  less  specialized  in  most 
features  of  the  wing. 


Phyllostomatidae 

The  New  World  leaf-nosed  bats,  along  with  the  noctilionids  and  mormoopids, 
tend  to  dominate  the  upper  right-hand  quadrant  of  discriminant  space  (Figs.  5,  6). 
Within  this  portion  of  space,  each  of  the  phyllostomatid  subfamilies  tends  to  occupy 
a  discrete  region  and  group  discrimination  vectors  generally  distinguished  each 
of  their  centroids.  There  is  a  rather  high  percentage  (22.30)  of  “misclassifica- 
tions”  (Fig.  17),  which  reflect  a  considerable  amount  of  variation  within  the 
family.  The  majority  of  these  “misclassifications”  involves  species  that  occupy 
a  position  near  the  grand  centroid.  Misclassifications  outside  of  the  family  limits, 
although  fewer  in  number,  also  tend  to  occur  in  this  region.  Among  phyllosto- 
matids,  the  desmodontines  exhibit  the  most  fidelity  to  their  group  discrimination 
vectors,  whereas  the  carolliines  show  the  least.  We  will  consider  the  general 
nature  of  phyllostomatid  wing  morphology  before  dealing  with  that  of  each  of  the 
subfamilies. 

As  has  been  the  case  in  previous  accounts,  the  length  of  head  and  body  of 
phyllostomatids  is  of  minor  importance  in  the  discrimination  of  the  family 
(Table  6).  The  range  of  variation  of  this  variable  is  large  and  ranges  from  such 
small-sized  species  as  Ametrida  centurio  to  the  large-sized  Vampyrum 
spectrum.  This  variation  nearly  encompasses  the  range  of  variation  observed  for 
the  order. 

The  vector  for  the  length  of  the  forearm  indicates  small  size  with  respect  to  this 
variable  for  all  phyllostomatid  subfamilies  (Figs.  4D-E,  5,  6).  The  absolute  length 
of  the  forearm  averages  slightly  below  the  mean  computed  for  all  bats  as  does 
the  relative  length  of  the  forearm  (Table  A8).  Table  6  indicates  a  rather  strong 
importance  of  the  shortness  of  the  forearm  in  the  discrimination  of  most  sub¬ 
families.  This  is  strongest  for  the  glossophagines,  carolliines,  and  stenodermines, 
but  it  is  rather  minor  with  regard  to  the  phyllonycterines. 

Although  the  dispersion  of  centroids  is  caused  by  the  overall  interaction  among 
all  variables,  the  vectors  that  appear  to  influence  most  directly  the  ordination 
of  phyllostomatid  centroids  are  those  associated  with  features  of  the  third  digit; 
most  imply  large  size.  The  vector  for  length  of  the  third  metacarpal  (C)  apparently 
is  a  strong  factor  in  the  discrimination  of  all  subfamilies  (Table  6).  The  tail  end 
of  the  vector  for  the  first  phalanx  of  digit  III  (D)  is  oriented  toward  the  phyl¬ 
lostomatid  centroids  (Figs.  4D-E,  5,  6)  and  implies  shortness  (see  also  Fig.  8). 
This  vector  is  a  moderately  strong  discriminator  of  the  family  (Table  6),  although 
it  does  not  appear  to  be  so  important  in  the  discrimination  of  the  Carolliinae.  The 
proportionately  long  third  phalanx  (F)  is  a  strong  discriminator  of  nearly  all  phyl¬ 
lostomatid  subfamilies  (Table  6);  phyllonycterines  and  desmodontines  appear  to 
be  less  characterized  by  this  variable. 


268 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


The  overall  effect  of  interplay  among  the  elements  of  the  third  digit  is  the 
production  of  a  span  that  generally  averages  longer  than  that  of  any  other  group 
of  bats  (Table  A9).  The  length  of  this  digit  contributes  nearly  67  per  cent  (range, 
61.31-70.10)  to  the  overall  length  of  the  wing,  which  is  larger  than  in  any  other 
chiropteran  family.  This  is  of  interest  in  that  the  third  digit  of  the  molossid  wing, 
which  is  generally  long-tipped  and  of  high  aspect,  contributes  a  somewhat  lower 
average  of  66.40  percent  (range,  64.36-69.70)  to  the  overall  span  of  the  wing.  The 
combined  effect  of  the  relatively  short  forearm  of  phyllostomatids  and  their  long 
third  digits  results  in  the  highest  tip  indices  of  any  group  of  chiropterans  (Fig.  10; 
Table  A2),  as  also  noted  by  Findley  et  al.  (1972). 

The  vectors  for  elements  of  the  fourth  digit  (G,  H,  I)  pass  tangentially  to  the 
position  of  phyllostomatid  centroids  and  a  precise  interpretation  of  their  effect  on 
alar  shape  is  difficult.  The  vector  for  the  fourth  metacarpal  (G)  suggests  large  size 
in  all  three  canonical  axes  (Figs.  4D-E,  5,  6).  The  vectors  for  the  two  phalangeal 
elements  of  digit  IV  (H,  I)  appear  to  exert  their  greatest  force  on  the  ordination  of 
phyllostomatid  centroids  in  the  third  canonical  axis  and  here  also  imply  large  size. 
Although  there  is  variation  within  the  family,  as  will  be  discussed  below,  the 
second  phalanx  of  the  fourth  digit  tends  to  be  proportionately  longer  than  the  first 
(Tables  A17,  A18).  Relative  to  the  span  of  the  wing,  the  fourth  digit  of  phy- 
lostomatids  averages  longer  (60. 1 1  per  cent  of  span)  than  does  that  of  most  other 
groups  of  bats;  only  the  molossids  are  larger  in  this  respect  (60.28  per  cent  of 
span).  In  addition,  the  total  length  of  this  digit  in  phyllostomatids  averages  nearly 
one  and  a  half  times  the  length  of  the  forearm  (range,  1.23-1.83). 

Whereas  phyllostomatids  and  molossids  exhibit  some  similarities  relative  to 
the  lengths  of  digits  III  and  IV,  these  two  families  are  markedly  dissimilar  with 
regard  to  the  length  of  digit  V.  Indicative  of  the  generally  low  aspect  nature  of 
phyllostomatid  wings,  the  fifth  digit  is  long  and  averages  1.44  (range,  1.26- 
1.68)  times  the  length  of  the  forearm.  The  vector  for  the  second  phalanx  of  this 
digit  (L)  appears  to  be  an  important  feature  in  the  discrimination  of  all  subfamilies 
of  phyllostomatids  (Table  6).  This  variable  has  its  strongest  effect  on  the  ordi¬ 
nation  of  phyllostomatid  centroids  in  the  third  canonical  axis  where  it  implies  large 
size  (Figs.  4E,  5).  The  vector  for  the  fifth  metacarpal  (J),  as  that  of  the  fourth 
metacarpal,  is  difficult  to  interpret  because  it  is  oriented  tangentially  to  the 
phyllostomatid  centroids  (Figs.  4D-E,  5,  6).  In  the  first  and  second  axes,  the 
implication  is  large  size,  but  shortness  is  emphasized  in  the  third  axis.  The  effect 
of  the  first  phalanx  of  digit  V  (K)  on  the  ordination  of  phyllostomatids  is  some¬ 
what  clearer,  and  it  implies  shortness  in  all  three  axes.  The  vector  for  the  second 
phalanx  of  digit  V  (L)  suggests  large  size.  The  relative  importance  of  these  two 
proximal  elements  in  the  discrimination  of  the  phyllostomatid  subfamilies  is 
variable  but  generally  high  (Table  6). 

Finally,  the  structural,  and  perhaps  phylogenetic,  similarity  of  wing  mor¬ 
phology  among  phyllostomatids  may  be  summarized  by  examining  the  angles 
between  the  discrimination  vectors  of  each  subfamily  (Table  7).  In  this  table, 
the  phyllostomatines  are  nearest  the  carolliines  and  glossophagines.  The  latter 
two  subfamilies  are  relatively  close  to  each  other  as  indicated  by  a  23.08  degree 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


269 


Table  7. — Angles  between  the  group  discriminant  functions  for  the  subfamilies  of  the 

Phyllostomatidae. 


Phyllostomatinae 

Glossophaginae 

Carolliinae 

Stenoderminae 

Phyllonycterinae 

Desmodontinae 

Phyllostomatinae 

00.00 

28.80 

26.30 

42.32 

43.47 

48.41 

Glossophaginae 

28.80 

00.00 

23.98 

25.46 

44.30 

37.17 

Carolliinae 

26.30 

23.98 

00.00 

35.02 

48.38 

56.18 

Stenoderminae 

42.32 

25.46 

35.02 

00.00 

45.68 

42.78 

Phyllonycterinae 

43.47 

44.30 

48.38 

45.68 

00.00 

45.48 

Desmodontinae 

48.41 

37.17 

56.18 

42.78 

45.48 

00.00 

divergence  between  their  respective  discrimination  vectors.  The  stenodermines 
fall  nearest  the  discrimination  vectors  of  glossophagines,  carolliines,  and  phyl- 
lostomatines,  respectively.  The  most  divergent  angles  between  group  dis¬ 
crimination  vectors  occur  between  phyllonycterines  and  desmodontines,  and  all 
other  subfamilies.  The  angle  between  the  discrimination  vectors  of  these  two  sub¬ 
families  also  is  rather  large  (45.48  degrees).  These  relationships  suggest  that  the 
phyllostomatines  form  the  nucleus  of  the  family,  which  is  rooted  in  proximity 
to  the  grand  centroid  for  all  bats.  The  glossophagines  and  carolliines  are  positioned 
relatively  close  to  the  phyllostomatines  and  these  three  subfamilies  constitute 
a  core  around  which  the  remaining  subfamilies  are  positioned.  The  stenodermines 
appear  to  be  morphologically  most  similar  to  the  glossophagines,  carolliines,  and 
phyllostomatines,  respectively.  The  phyllonycterines  and  desmodontines  occupy 
widely  separated  positions  from  each  other  as  well  as  from  the  other  subfamilies. 
The  phyllonycterines  appear  to  be  morphologically  nearer  phyllostomatines  and 
glossophagines,  respectively,  than  to  other  subfamilies,  whereas  desmodontines 
appear  to  approach  most  closely  the  glossophagines. 

Phyllostomatinae 

The  phyllostomatines  are  generally  the  largest  bats  of  the  family  in  terms  of 
absolute  size;  Vampyrum  spectrum  (40),  Chrotopterus  auritus  (39),  and 
Phyllostomus  hastatus  (34)  far  exceed  most  New  World  species  in  overall  size. 
However,  aside  from  these  and  several  other  large-sized  species,  the  phyllosto¬ 
matines  are  about  average  or  slightly  below  average  in  size.  Compared  to  other 
phyllostomatids,  their  wings  are  relatively  long  (Table  A7)  and  the  relative  length 
of  the  forearm  averages  longest  of  all  phyllostomatids  (Table  A8).  The  relative 
length  of  the  third  digit  is  average  or  slightly  above  average  for  the  family  (Table 
A9).  As  a  consequence  of  the  interaction  between  these  two  lengths,  the  tip  index 
of  phyllostomatines  is  comparatively  low  for  the  family  (Table  A2).  In  terms  of 
the  overall  aspect  ratios,  the  wings  of  phyllostomatines  are  in  the  middle  of  the 
range  for  the  family  (Tables  A3-A5).  Wing  loading  for  this  subfamily  also  is  near 


270 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  11. — Canonical  graph  of  the  species  of  the  subfamilies  Phyllostomatinae,  Carolliinae, 
and  Desmodontinae  plotted  on  the  first  and  third  canonical  axes.  Stars  represent  the  sub- 
familial  group  centroids:  A,  Phyllostomatinae;  B,  Glossophaginae;  C,  Carolliinae;  D, 
Stenoderminae;  E,  Phyllonycterinae;  F,  Desmodontinae.  Genera  are  encircled  as  follows: 
Phyllostomatinae — Micronycteris  (1-12),  Macrotus  (13-14),  Lonchorhina  (15-17),  Macro- 
phyllum  (18-19),  Tunatia  (20-26),  Mimon  (27-31),  Phyllostomus  (32-36),  Phylloderma 
(37),  Trachops  (38),  Chrotopterus  (39),  Vampyrum  (40);  Carolliinae — Carollia  (41-44), 
Rhinophylla  (45-47);  Desmodontinae — Desmodus  (48-49),  Diaemus  (50),  Diphylla  (51). 
Species  are  identified  by  corresponding  bold-faced  numbers  in  the  list  of  specimens  examined. 

the  median  of  the  family,  although  the  range  of  variation  within  the  subfamily  is 
large  (Table  A6). 

The  centroid  for  the  phyllostomatines  is  located  near  the  grand  centroid  for 
all  bats.  In  the  canonical  graphs  that  show  positions  of  individual  species  (Figs.  1 1, 
12),  it  will  be  noted  that  the  genus  Micronycteris  (1-12)  encompasses  the  grand 
centroid  in  the  first  three  canonical  axes.  It  is  interesting  to  note  here  that  the  five 
classificatory  “misses”  from  this  subfamily  to  the  Vespertilionidae  (Fig.  17) 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


271 


Fig.  12. — Canonical  graph  of  the  species  of  the  subfamilies  Phyllostomatinae,  Carolliinae, 
and  Desmodontinae  plotted  on  the  first  and  second  canonical  axes.  See  legend  of  Fig.  1 1  for 
key  to  group  centroids  (stars)  and  genera  (encircled  dots). 


involve  Micronycteris  megalotis  (1-2),  M.  pusilla  (8),  M.  nicefori  (9),  and  M. 
behni  (11).  Most  of  the  other  species  of  phyllostomatines  cluster  together  around 
the  centroid  for  the  subfamily.  However,  there  are  several  notable  departures 
from  the  group  centroid. 

Two  species,  V ampyrum  spectrum  (40)  and  Chrotopterus  auritus  (39),  are 
most  obvious  in  their  departure  from  the  subfamilial  centroid,  especially  along 
the  third  canonical  axis.  Most  of  this  dispersion  appears  to  be  caused  by  the  vector 
for  length  of  the  head  and  body.  In  addition,  vectors  associated  with  comparatively 
short  wings  appear  to  affect  these  two  species.  In  both,  the  lengths  of  forearm 
and  third  digit  are  short  as  compared  to  other  members  of  the  subfamily  (Tables 
A8,  A9).  The  span  of  the  third  digit  is  most  influenced  by  the  vector  for  the  third 
metacarpal,  which  implies  shortness  of  this  element  in  these  two  species  (Table 


272 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  13. — Canonical  graph  of  the  species  of  the  subfamily  Glossophaginae  plotted  on  the 
first  and  third  canonical  axes.  Stars  represent  subfamilial  group  centroids  (see  legend  of  Fig. 
11  for  key).  Genera  are  encircled  as  follows:  Glossophaga  (1-4),  Monophyllus  (5-6), 
Leptonycteris  (7-9),  Lonchophylla  (10-14),  Lionycteris  (15),  Anoura  ( 16-20),  Scleronycteris 
(21),  Lichonycteris  (22-24),  Hylonycteris  (25),  Platalina  (26),  Choeroniscus  (27-31), 
Choeronycteris  (32),  Musonycteris  (33).  Species  are  identified  by  corresponding  bold-faced 
numbers  in  the  list  of  specimens  examined. 

A 12).  However,  the  lengths  of  the  first  and  third  phalanges  average  the  largest 
in  percentage  contributed  to  the  length  of  digit  III  (Tables  A13,  A15).  The 
metacarpals  of  the  fourth  and  fifth  digit  are  proportionately  short  for  the  sub¬ 
family  (Table  A 16,  A 19),  although  the  phalangeal  elements  of  these  two  digits  are 
generally  long.  The  terminal  phalanx  of  the  fifth  digit  is  comparatively  longer 
than  in  most  other  phyllostomatines  (Table  A21). 

For  the  most  part,  the  genus  Phyllostomus  (32-36)  ordinates  with  the  previous 
two  species  in  the  first  and  second  canonical  axes  (Fig.  12).  However,  Phyl¬ 
lostomus  disassociates  from  this  relationship  in  the  third  dimension  of  dis- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


273 


Fig.  14. — Canonical  graph  of  the  species  of  the  subfamily  Glossophaginae  plotted  on  the 
first  and  second  canonical  axes.  See  legend  of  Fig.  1 1  for  key  to  group  centroids  (stars)  and 
legend  of  Fig.  13  for  key  to  genera  (encircled  dots). 


criminant  space  (Fig.  11).  In  this  axis,  Phyllostomus  tends  to  deny  the  influence 
of  length  of  head  and  body  and  is  aligned  by  vectors  that  imply  a  long  third  digit. 
The  vector  for  the  metacarpal  (C)  is  especially  important  in  this  regard  (Table 
A 12).  The  first  phalanx  is  the  shortest  among  all  members  of  the  subfamily  and 
nearly  the  family  as  a  whole  (Table  A13);  only  the  vampire  bats  have  a  pro¬ 
portionately  shorter  first  phalanx  in  the  third  digit.  Other  features  that  dis¬ 
tinguish  Phyllostomus  from  most  other  phyllostomatines  are  long  fourth  and  fifth 
metacarpals  (Table  A16,  A19),  and  short  distal  phalanx  in  digit  V  (Table  A21). 
These  features  also  are  characteristic  of  the  vampire  bats,  and  it  is  interesting  to 
note  that  all  species  of  Phyllostomus,  except  P.  latifolius  (36)  and  a  close  associate 
Phylloderma  stenops  (37),  “misclassify”  as  desmodontines.  The  species  latifolius 
and  stenops  “misclassify”  as  stenodermines  (Fig.  17). 


274 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


The  genera  Mimon  (27-31)  and  Tonatia  (20-26)  portray  an  interestingly 
antagonistic  relationship  to  each  other  relative  to  the  group  centroid.  This  re¬ 
lationship  is  exaggerated  by  M.  crenulatum  (29-30)  and  M.  koepckeae  (31)  on 
the  one  hand  and  T.  silvicola  (25)  and  T.  venezuelae  (26)  on  the  other.  Generally, 
Minion  has  the  highest  aspect  ratio  as  compared  to  other  phyllostomatines, 
whereas  Tonatia  has  the  lowest  (Tables  A3-A5).  Mimon  has  the  longest  wing, 
in  a  relative  sense,  of  any  phyllostomatid,  whereas  the  wing  of  Tonatia  is  much 
shorter  (Table  A7).  This  relationship  applies  to  most  features  examined  in  this 
study.  Incidently,  the  two  extreme  species  of  Mimon  “misclassify”  as  stenoder- 
mines  (Fig.  17),  which  generally  have  longer,  narrower  wings  as  compared  to 
the  other  phyllostomatids.  Other  phyllostomatines  that  are  “misclassified”  (Fig. 
17)  are  Micronycteris  daviesi  (12)  and  Macrophyllum  macrophyllum  (18-19), 
which  are  aligned  with  the  Glossophaginae. 

Glossophaginae 

The  long-tongued  bats  tend  to  form  a  rather  tightly  packed  cluster  (Figs.  13, 
14),  which  nestles  close  to  the  clusters  of  the  phyllostomatines  and  carolliines 
(Table  7).  As  a  group,  the  glossophagines  have  relatively  short  wings  as  com¬ 
pared  to  other  phyllostomatids  (Table  A7).  The  relative  length  of  the  forearm 
averages  a  little  over  half  (0.63)  the  length  of  head  and  body  (Table  A8).  Com¬ 
paratively  speaking,  the  third  digit  is  relatively  long,  which  produces  a  rather 
large  average  tip  index  (2.06)  for  the  subfamily  (Table  A2).  The  overall  aspect 
ratio  of  the  wings  of  glossophagines  is  highest  for  the  family — notable  extremes  are 
Anoura  (16-20)  6.50,  Musonycteris  ( 33)  6.30,  and  Scleronycteris  { 21)  6.23.  This 
also  applies  to  the  aspect  ratio  of  the  tip  region  (Tables  A3,  A4). 

In  view  of  the  tight  packing  of  the  group,  a  precise  interpretation  of  the 
variable  vectors  on  the  dispersion  of  glossophagines  is  difficult.  Most  of  the 
differences  are  small,  quantitative  shifts  in  the  range  of  variation.  The  vectors 
that  appear  to  affect  most  heavily  the  ordination  of  the  glossophagines  are  those 
for  the  forearm  (B),  third  metacarpal  (C),  and  second  phalanx  in  the  fifth  digit 
(L).  The  vector  for  the  forearm  (B)  implies  shortness  for  most  species.  However, 
Leptonycteris  (7-9),  Lionycteris  (15),  Scleronycteris  (21),  and  Choeronycteris 
(32)  generally  have  longer  forearms  than  other  glossophagines  (Table  A8). 

The  vector  for  the  metacarpal  of  digit  III  (C)  suggests  large  size  and  Leptonyc¬ 
teris  and  Lionycteris  represent  the  large  extremes  relative  to  this  feature  (Table 
A 12).  As  a  group,  the  glossophagines  possess  proportionately  longer  second 
phalanges  of  digit  V  than  do  any  other  bats  except  the  pteropodids  (Table  A21). 

Two  species,  Hylonycteris  underwoodi  (25)  and  Platalina  genovensium  (26), 
“misclassify”  to  the  Stenoderminae  and  are  most  closely  associated  with  Sturnira 
and  Vampyrops.  Also,  Lichonycteris  (23-24)  disperses  among  these  stenoder- 
mine  genera,  although  its  classification  is  mostly  to  the  Glossophaginae. 

Carolliinae 

The  group  discrimination  vectors  for  this  subfamily  are  relatively  weak.  In 
Fig.  17,  two  species,  Rhinophylla  pumilio  (45)  and  R.  fischerae  (47)  are  “mis- 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


275 


classified”  as  glossophagines  and  four  other  species,  Rhinophylla  alethina  (46), 
Carollia  subrufa  (42),  C.  brevicauda  (43),  and  C.  perspicillata  (44)  are  associated 
with  the  Stenoderminae.  This  leaves  only  one  species,  Carollia  castanea  (41), 
which  suggests  that,  in  terms  of  wing  shape,  the  carolliines  are  rather  indistinct 
and  may  bridge  the  gap  between  glossophagines  and  the  stenodermines  (Figs.  1 1, 
12;  Table  7). 

As  a  group,  the  carolliines  have  relatively  long  wings  (Table  A7).  This  results 
from  the  combination  of  a  moderately  long  forearm  and  an  exceptionally  long 
digit  III.  These  features  also  characterize  the  stenodermines.  Carolliines  further 
resemble  stenodermines  in  possessing  a  comparatively  short  digit  IV;  primarily 
the  result  of  a  proportionately  short  fourth  metacarpal  (Table  A 16). 

Stenoderminae 

Stenodermines  represent  the  most  diverse  of  the  phyllostomatid  subfamilies. 
The  dispersion  of  the  various  species  of  this  subfamily  in  discriminant  space  is 
comparable  to  that  seen  in  the  Phyllostomatinae,  although  the  group  generally 
occupies  space  unfilled  by  other  taxa  (Figs.  15,  16).  The  group,  as  a  whole,  is 
generally  displaced  away  from  the  congested  area  nearer  the  grand  centroid. 
However,  two  small-sized  species,  Vampyressa  pusilla  (24)  and  Sphaeronycteris 
toxophyllum  (56),  approach  the  grand  centroid  close  enough  to  be  confused  with 
the  Vespertilionidae  (Fig.  17).  In  addition,  Phyllops  haitiensis  (51)  and  Centurio 
senex  (57)  are  “misclassified”  as  phyllostomatines,  and  Vampyressa  nymphaea 
(26),  Pygoderma  bilabiatum  (54),  and  Ametrida  centurio  (55)  are  confused  with 
glossophagines. 

Unlike  any  other  subfamily  of  phyllostomatids,  which  tend  to  orient  in  uni- 
modal  directions  in  discriminant  space,  the  stenodermines  appear  to  ordinate 
into  two  slightly  different  portions  of  this  space  (Figs.  15,  16).  The  extremes 
of  this  dichotomy  are  Artibeus  (35-47)  on  one  hand  and  Vampyrops  (13-22) 
and  Sturnira  (1-10)  on  the  other.  Although  the  small-sized  species  of  both  groups 
tend  to  congregate  around  the  group  centroid,  the  large-sized  species  of  each 
group  orient  away  from  each  other  (Fig.  16). 

In  the  first  three  canonical  axes  (Figs.  15  and  16),  vectors  that  imply  large  size 
for  the  forearm  (B),  fifth  metacarpal  (J),  and  second  phalanx  of  digit  V  (L) 
ordinate  Artibeus  away  from  Vampyrops  and  Sturnira  (Fig.  16).  These  vectors 
imply  shortness  of  these  variables  in  both  Sturnira  and  Vampyrops.  The  latter 
two  taxa  are  more  directly  ordinated  by  vectors  associated  with  the  third  meta¬ 
carpal  (C),  and  second  and  third  phalanges  of  digit  III  (E,  F).  All  suggest  long 
length. 

The  tip  index  and  aspect  ratio  of  the  tip  are  generally  higher  in  Vampyrops  and 
Sturnira  than  in  Artibeus.  As  might  be  expected,  Artibeus  has  a  somewhat  higher 
aspect  ratio  of  the  plagiopatagial  region,  primarily  as  a  result  of  a  proportionately 
longer  forearm  (Table  A8).  The  composition  of  the  third  digit  is  similar  in  both 
groups,  although  Artibeus  tends  to  have  a  long  metacarpal  and  generally  short 
phalangeal  elements,  whereas  in  Vampyrops  and,  to  a  lesser  extent,  Sturnira , 
construction  of  most  of  the  span  of  this  digit  results  from  long  phalangeal  elements. 


276 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


III 


I  I 


Fig.  15. — Canonical  graph  of  the  species  of  the  subfamilies  Stenoderminae  and  Phyl- 
lonycterinae  plotted  on  the  first  and  third  canonical  axes.  Stars  represent  subfamilial  group 
centroids  (see  legend  of  Fig.  1 1  for  key).  Genera  are  encircled  as  follows:  Stenoderminae — 
Sturnira  (1-10),  Urodermci  (1 1-12),  Vampyrops  (13-22),  Vumpyrodes  { 23),  Vampy ressa  (24- 
28),  Chiroderma  (29-33),  Ectophylla  (34),  Artibeus  (35-47),  Enchisthenes  ( 48),  Ardops  (49), 
Phyllops  (50-51),  Ariteus  (52),  Stenoderma  (53),  Pygoderma  (54),  Ametrida  (55),  Sphae- 
ronycteris  (56),  Centurio  (57);  Phyllonycterinae — Brachyphylla  (58-59),  Erophylla 
(60-61),  Phyllonycteris  (62-63).  Species  are  identified  by  corresponding  bold-faced  numbers 
in  the  list  of  specimens  examined. 


Sturnira  does  not  quite  fit  this  scheme  because  the  second  phalanx  is  propor¬ 
tionately  short  (Table  A 14).  However,  the  proportional  length  of  the  distalmost 
phalanx  of  the  third  digit  appears  to  compensate  for  this  (Table  A15). 

Phyllonycterinae 

This  subfamily,  as  well  as  the  desmodontines,  is  ordinated  into  a  peripheral 
position  of  discriminant  space  relative  to  the  other  phyllostomatid  subfamilies 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


277 


Fig.  16. — Canonical  graph  of  the  species  of  the  subfamilies  Stenoderminae  and 
Phyllonycterinae  plotted  on  the  first  and  second  canonical  axes.  See  legend  of  Fig.  1 1 
for  key  to  group  centroids  (stars)  and  legend  of  Fig.  15  for  key  to  genera  (encircled  dots). 


(Figs.  11-12,  15-16;  Table  7).  The  flower  bats  have  the  shortest  wings,  in  a 
relative  sense,  among  the  Phyllostomatidae  (Table  A7).  They  resemble  phyl- 
lostomatines  in  possessing  relatively  long  forearms  (Table  A8).  The  group  has 
the  shortest  relative  length  of  digit  III  as  compared  to  that  of  other  phyllostomatids. 
This  is  not  particularly  surprising  inasmuch  as  the  vectors  for  elements  of  this 
digit  (C,  D,  E,  F)  are  oriented  away  from  the  group  centroid  (Figs.  4D-E,  15,  16). 
The  length  of  the  third  digit  is  composed  primarily  of  the  phalangeal  elements, 
which  are  equal  or  subequal  in  length  (Fig.  8).  As  might  be  predicted  from  their 
relative  position  in  discriminant  space,  Erophylla  bombifrons  (60)  and  E.  seze- 
korni  (61)  “misclassify”  as  phyllostomatines  (Fig.  17). 


278 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Desmodontinae 

The  vampire  bats  occupy  the  most  peripheral  position  in  discriminant  space 
relative  to  all  other  phyllostomatid  subfamilies.  As  there  is  complete  fidelity  to 
their  discriminant  vectors,  there  are  no  instances  of  “misclassification”  of  members 
of  this  group  (Fig.  17),  which  suggests  the  distinctive  shape  of  the  desmodontine 
wing  (Fig.  8).  This  distinctness  also  is  reflected  in  the  rather  large  generalized 
distance  from  the  other  phyllostomatid  centroids:  Carolliinae,  5.65;  Stenoder- 
minae,  4.46;  Glossophaginae,  4.43;  Phyllostomatinae,  4.28;  and  Phyllonycteri- 
nae,  4.28.  The  most  important  vectors  in  the  ordination  of  the  group  appear  to 
be  those  associated  with  the  third  metacarpal  (C),  which  imply  large  size,  and 
those  for  the  first  phalanx  of  digit  III  (D)  and  first  and  second  phalanx  of  digit  V 
(K,  L),  which  emphasize  shortness  (Tables  A12,  A13,  A20,  and  A21).  Because  of 
the  compensating  effects  of  long  metacarpal  elements  in  the  fourth  and  fifth 
digits,  the  wing  of  vampire  bats  tends  to  be  relatively  short  and  broad  and  of  gen¬ 
erally  low  aspect  ratio.  The  vampire  wing  is  the  most  heavily  loaded  of  all  phyl- 
lostomatids  (Table  A6);  note  that  the  phyllonycterines  follow  the  desmodontines 
in  this  regard. 


Natal  idae 

An  interpretation  of  the  alar  shape  of  natalid  wings  is  difficult.  Part  of  this 
results  from  the  rather  small  sample  size  for  this  family  as  well  as  for  other  families 
with  which  the  natalids  appear  to  be  associated — namely,  the  Thyropteridae  and 
Craseonycteridae.  Also,  these  three  families  appear  to  be  associated  with  the 
Vespertilionidae,  for  which  there  was  a  disproportionately  large  sample  size. 
Finally,  the  centroids  of  all  four  families  as  well  as  that  of  the  Myzapodidae 
lie  in  proximity  to  the  grand  centroid  for  all  bats  (Figs.  5,  6),  tending  to  obscure 
the  precise  relationships  of  one  to  another. 

In  the  principal  component  analysis,  the  natalids,  craseonycterids,  and 
thyropterids  dispersed  together  towards  the  right-hand  portion  of  Euclidean  space 
(Fig.  3),  which,  as  we  have  noted  above,  indicates  their  general  small  size  for  all 
variables.  The  vespertilionid  centroid,  although  ordinated  towards  the  small¬ 
sized  side  of  the  array,  occupies  a  more  central  position  in  the  overall  dispersion. 
On  the  other  hand,  the  position  of  these  four  group  centroids  in  discriminant 
space  is  somewhat  different  (Figs.  5,  6). 

The  natalids  align  most  closely  with  the  craseonycterids  in  the  discriminant 
analysis.  The  shared  absence  of  the  third  phalanx  of  digit  III  appears  overly  to 
bias  this  association.  On  the  basis  of  this  variable  alone,  the  generalized  distances 
between  natalids/craseonycterids,  thyropterids,  and  vespertilionids;  craseonycte- 
rids/thyropterids  and  vespertilionids;  and  thyropterids/vespertilionids  are: 
0.093,  1.466,  and  2.016;  1.466  and  2.016,  and  0.550,  respectively.  The  overall 
generalized  distances  between  these  centroids  are  4.050,  3.540,  and  4.400; 
5.580  and  5.457;  and  2.489,  respectively.  However,  the  generalized  distances 
between  these  four  families,  on  the  basis  of  each  variable,  tend  to  indicate  a  closer 
association  between  natalids,  thropterids,  and  vespertilionids  than  between 
craseonycterids  and  these  three  families. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


279 


Thus,  the  resemblance  of  natalids  and  craseonycterids  might  be  spurious  as 
a  result  of  the  absence  of  the  third  phalanx  of  digit  III  and  a  concomitant  com¬ 
pensation  in  the  length  of  elements  of  this  digit,  especially  that  of  the  second 
phalanx  (Fig.  9).  In  addition,  there  appears  to  be  a  “general”  tendency  for  small¬ 
sized  bats  to  have  similarly  constructed  wings  (that  is,  long  forearm,  long  digit 
III,  and  generally  long  digits  IV  and  V).  Findley  et  al.  (1972)  also  noted  this 
tendency,  but  we  would  caution  the  reader  by  noting  that  some  relatively  large¬ 
sized  bats,  such  as  noctilionids,  emballonurids,  and  nycterids  (among  others),  also 
follow  this  trend  (Tables  A2,  A7,  A8-A1 1).  Hence,  we  reiterate  our  earlier  state¬ 
ment  that  the  relationships  between  general  body  size  and  wing  morphometries 
are  much  more  complicated  than  bivariate  comparisons  would  seem  to  indicate. 

In  the  first  three  canonical  axes  (Figs.  4D-E,  5,  6),  the  ordination  of  the  natalid 
centroid  appears  to  be  affected  by  interactions  among  variables,  similar  to  those 
noted  above  for  the  craseonycterids.  In  the  previous  accounts,  we  have  discussed 
the  apparent  minor  role  of  the  length  of  the  head  and  body  in  the  discrimination 
of  groups.  With  regards  to  the  natalids  as  well  as  the  thyropterids  and  myzapodids 
the  influence  of  this  variable,  albeit  weak,  is  comparatively  stronger  than  noted 
for  other  families  (Table  6).  The  relative  length  of  the  wing  of  natalids  is  2.61  times 
the  length  of  the  head  and  body  and  is  among  the  longest  found  among  all  bats 
(Table  A7).  This  span  is  composed  of  a  relatively  long  forearm  (Table  A8),  and 
digit  III  has  a  mean  relative  length  (1 .69)  that  is  highest  among  all  bats,  Table  A9. 
Similarly,  large  values  for  these  relative  lengths  will  be  noted  for  craseonycterids, 
thyropterids,  and  furipterids. 

The  composition  of  the  third  digit  of  natalids  is  more  like  that  of  thyropterids 
and  vespertilionids  than  that  of  craseonycterids.  The  vector  for  the  third  meta¬ 
carpal  (C)  of  natalids  implies  shortness  as  was  the  case  in  the  Craseonycteridae. 
However,  the  reader  will  recall  that  the  second  phalanx  of  digit  III  offset  the 
proportional  length  of  the  third  metacarpal  in  the  craseonycterids.  In  the 
extradimensional  fourth  and  fifth  canonical  axes,  the  vector  for  the  third  meta¬ 
carpal  more  strongly  implies  longness  of  this  variable  for  natalids,  thyropterids, 
and  vespertilionids.  This  also  is  generally  the  case  for  the  first  and  second  phalanx 
of  digit  III  for  these  three  families. 

The  combined  effect  of  variable  vectors  for  elements  of  the  fourth  digit  (G,  H, 
I)  of  natalids  indicates  longness  of  this  digit  (Table  A 10).  The  most  important 
components  of  length  appear  to  be  the  phalangeal  elements,  although  these  are 
generally  below  the  average  computed  for  all  bats  (Tables  A17,  A18).  The  length 
of  the  first  phalanx  of  digit  IV  contributes  markedly  to  the  discrimination  of  the 
group  (Table  6).  Again,  an  interpretation  of  the  vectors  for  this  digit  is  obscured 
by  the  synergistic  interaction  among  all  variables.  Shortness  of  the  fourth  meta¬ 
carpal  (G)  is  suggested  in  the  first  three  canonical  axes.  However,  in  extradimen¬ 
sional  axes  this  vector  implies  longness  of  this  variable  in  natalids,  thyropterids, 
and  vespertilionids;  shortness  is  indicated  for  that  of  craseonycterids. 

The  relative  length  of  the  fifth  digit  of  natalids  averages  the  longest  among  all 
bats  (Table  All);  the  mean  relative  length  of  this  digit  for  furipterids  and  thy¬ 
ropterids  also  is  high.  The  variable  vectors  for  the  lengths  of  the  metacarpal  (J) 


280 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


and  second  phalanx  (L)  strongly  suggest  longness  in  the  first  three  canonical  axes 
(Figs.  4D-E,  5,  6),  whereas  that  for  the  length  of  the  first  phalanx  of  digit  V 
implies  shortness. 

In  general  appearance  (Fig.  9),  the  wings  of  natalids  are  below  average  in  their 
overall  aspect  ratio  (Table  A3).  This  low  aspect  also  is  reflected  in  the  aspects 
of  the  tip  and  plagiopatagial  portions  of  the  wing  (Fig.  10;  Tables  A4,  A5). 
Craseonycterids,  thyropterids,  furipterids,  myzapodids,  and  to  a  certain  extent, 
vespertilionids  resemble  natalids  in  these  respects.  It  is  interesting  to  note  that, 
with  regard  to  wing  loading,  the  craseonycterids  possess  more  heavily  loaded  wings 
than  do  any  of  the  five  aforementioned  families  (Table  A6). 

Little  is  known  concerning  the  flight  characteristics  of  natalids.  We  concur  with 
Findley  et  al.  (1972)  in  their  suggestion  of  slow,  maneuverable  flight  potential 
for  these  bats;  also,  hovering  may  be  well  within  this  potential. 

Thyropteridae 

The  interpretation  of  wing  morphometries  of  the  disc-winged  bats  is  obscured 
by  the  positioning  of  their  group  centroid  almost  exactly  on  the  grand  centroid  of 
all  bats  (Figs.  5,  6).  This,  in  itself,  reflects  the  average  character  of  the  shape  of 
their  wings.  However,  the  confidence  circle  for  the  group  centroid  is  com¬ 
paratively  large,  possibly  reflecting  the  rather  small  sample  size  utilized  in  this 
study. 

In  the  classificatory  phase  of  the  discriminant  analysis  (Fig.  17),  both  species  of 
thyropterids  “misclassify”  as  vespertilionids.  This  could  reflect  a  correct  assign¬ 
ment  or  it  simply  might  be  an  artifact  of  small  sample  size.  The  generalized  distance 
between  these  two  families  is  comparatively  small  (2.489)  and  the  generalized 
distances,  based  on  each  variable,  likewise  support  this  close  association  of  the 
two. 


Myzapodidae 

Little  can  be  said  concerning  the  shape  of  the  wing  of  Myzapoda  aurita.  The 
group  centroid  is  in  proximity  to  the  grand  centroid  for  all  bats  (Figs.  5,  6);  the 
confidence  circle  exceeds  the  limit  of  the  figures  and  probably  reflects  the  small 
sample  size  of  two  specimens.  In  the  classificatory  phase  of  the  discriminant 
analysis,  these  bats  as  well  as  the  thyropterids  (noted  above)  were  “misclassified” 
as  vespertilionids. 


Vespertilionidae 

The  members  of  this  family  are  extremely  diverse  in  the  shapes  of  their  wings 
and  presumably  in  their  flight  characteristics.  The  group  centroid  is  located 
near  the  grand  centroid  for  all  bats  (Figs.  5,  6),  but  unlike  the  previous  two  groups 
the  confidence  circle  is  small,  and  the  group  discrimination  vector  appears  to  be 
relatively  strong.  The  one  “misclassification”  from  this  family  involved  Eudiscopus 
denticulus,  which  was  confused  with  the  Phyllostomatinae  (Fig.  17).  Several  other 
species  of  vespertilionids  were  associated  with  phyllostomatid  subfamilies,  but 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


281 


only  this  one  was  so  classified.  In  previous  accounts,  it  was  noted  that  several 
phyllostomatids,  as  well  as  thyropterids  and  myzapodids,  were  incorrectly  assigned 
to  the  Vespertilionidae.  Aside  from  possible  errors  associated  with  sample  size, 
we  suspect  that  these  “misclassifications”  reflect  general  similarities  among  these 
species  as  a  result  of  their  proximity  to  the  chiropteran  norm  (grand  centroid). 

Generally,  the  wings  of  the  vespertilionids  are  moderately  long  and  average 
about  twice  (2.07)  the  length  of  head  and  body  (Table  A7).  The  range  in  variation 
is  markedly  large  and  extends  from  Mimetillus  moloneyi,  with  its  peculiarly- 
shaped  wing  (barely  1.4  times  the  length  of  its  head  and  body),  to  Otonycteris, 
Kerivoula,  Miniopterus,  and  Eudiscopus,  wings  of  which  are  nearly  2.5  times 
the  head  and  body  length. 

The  vector  for  length  of  the  forearm  (B)  contributes  moderately  to  the  group 
discrimination  vector  of  the  family  (Table  6).  The  mean  relative  length  of  the 
forearm  is  slightly  above  average  for  all  bats  (0.74),  but  the  range  within  the 
family  includes  nearly  the  total  of  variation  exhibited  by  the  order  (Table  A8). 

The  position  of  the  group  centroid  relative  to  the  vectors  associated  with  the 
elements  of  digit  III  (C,  D,  E,  F)  generally  reflects  the  emphasis  on  the  long  length 
of  this  digit  in  the  composition  of  the  wing  (Figs.  4D-E,  5,  6).  The  mean  tip  index 
for  the  family  (1.81)  is  slightly  below  the  average  for  all  bats  (Table  A2),  but 
the  range  in  variation  includes  values  that  are  twice  the  length  of  the  forearm 
(for  example,  Eudiscopus,  2.17;  Kerivoula,  2.12;  Harpiocephalus,  2.04;  and 
Lasiurus,  2.00).  In  the  first  three  canonical  axes,  the  vector  for  the  meta¬ 
carpal  (C)  implies  large  size.  The  percentage  of  variation  contributed  to  the  group 
discrimination  vector  by  this  vector  is  relatively  high  (18.60,  Table  6).  Table  A12 
shows  that,  on  the  average,  approximately  50  per  cent  of  the  length  of  the  third 
digit  is  accounted  for  by  this  element.  As  has  been  the  case  for  the  majority  of 
the  families  discussed  to  this  point,  the  vector  for  the  first  phalanx  of  digit  III  (D) 
implies  shortness.  On  the  whole,  vespertilionids  fall  just  below  the  average  for  all 
bats  with  respect  to  this  feature  (Table  A13).  The  vector  for  the  length  of  the 
second  phalanx  of  digit  III  (E)  nearly  equals  the  metacarpal  in  its  influence  in 
the  discrimination  of  the  group  centroid  (Table  6).  The  implication  of  this  variable 
vector  is  shortness  and  the  mean  percentage  contributed  to  the  length  of  the  digit 
III  (Table  A14)  tends  to  support  this.  The  high  extremes  in  the  range  of  variation 
of  this  percentage  are  noteworthy.  The  second  phalanx  constitutes  33.77  per 
cent  of  the  total  length  of  digit  III  in  Miniopterus.  Similarly,  this  phalanx  is 
proportionately  long  in  Lasionycteris,  Chalinolobus,  and  Kerivoula  (27.31, 
26.77 ,  and  26.42  per  cent,  respectively).  The  vector  for  the  length  of  the  third 
phalanx  of  digit  III  (F)  is  moderately  important  in  the  discrimination  of  the  family. 
However,  its  precise  effect  on  the  dispersion  of  the  group  centroid  is  difficult  to 
assess  because  this  phalanx  is  indistinguishable  or  absent  in  some  species  and 
markedly  elongate  in  others.  In  most  vespertilionid  species,  this  phalanx  com¬ 
prises  10  per  cent  or  less  of  the  length  of  digit  III  (Table  A15);  20.13  per  cent  is 
contributed  by  this  element  in  the  wing  of  Eudiscopus. 

The  interaction  among  the  elements  that  compose  the  fourth  digit  is  complex 
and,  as  will  be  noted  in  Tables  A16-A18,  the  range  of  variation  is  wide.  The 


282 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


vector  for  the  metacarpal  (G)  implies  large  size.  The  effect  of  this  vector  on  the 
ordination  of  the  vespertilionids  appears  to  be  similar  to  that  exerted  in  the 
Molossidae  (Figs.  4D-E,  5,  6),  although,  in  the  latter,  all  vectors  associated  with 
elements  of  the  fourth  digit  appear  to  apply  a  more  direct  force  on  the  ordination. 
The  first  phalangeal  element  of  the  fourth  digit  is  about  average  in  its  pro¬ 
portional  length  as  compared  to  that  of  other  bats  (Table  A17).  The  vector  for  the 
length  of  the  second  phalanx  of  digit  IV  (I)  emphasizes  shortness  and  this  is 
generally  supported  in  Table  A18,  although,  again,  the  range  of  variation  is  wide. 

The  length  of  the  fifth  digit  of  vespertilionids  appears  to  be  controlled  mainly 
by  the  length  of  the  metacarpal  element.  The  vector  for  this  variable  (J)  is  important 
in  the  overall  ordination  of  the  vespertilionids  as  indicated  by  the  relatively  high 
percentage  (16.53)  contributed  to  the  discrimination  vector  of  the  group  (Table 
6).  The  vespertilionids  rank  second  highest  with  regard  to  the  mean  per  cent 
contributed  by  the  fifth  metacarpal  to  the  total  length  of  digit  V  (Table  A 19). 
Whereas  noctilionids  average  larger  than  vespertilionids  with  regard  to  the 
proportional  length  of  the  fifth  metacarpal,  the  high  extremes  in  the  range  of 
variation  among  vespertilionids  far  exceed  that  of  any  other  bats.  Notable  among 
these  extremes  are  Mimetillus  (82.35  per  cent),  Philetor  (75.87), 
Scotophilus  (73.07),  Tylonycteris  (72.84),  and  Nyctalus  (72.56).  The  vector  for 
length  of  the  first  phalanx  of  digit  V  (K)  implies  shortness,  but  this  variable  is 
of  minor  importance  in  the  discrimination  of  the  group  (Table  6).  The  vector  for 
the  length  of  the  distal  phalanx  of  this  digit  (L)  is  somewhat  stronger  in  its  influence 
on  the  group  discrimination  vector  (Table  6)  and  it  suggests  shortness. 

In  a  general  descriptive  sense,  the  wings  of  vespertilionids  are  not  particularly 
striking;  they  are  about  average  or  slightly  below  average  in  most  respects  as 
compared  to  those  of  other  members  of  the  order.  However,  in  terms  of  internal 
composition,  wing  variation  in  vespertilionids  is  the  most  complex  of  any  family 
we  have  examined.  This  is  particularly  true  of  species  that  depart  from  the  family 
norm,  that  is,  those  vespertilionids  with  wings  of  higher  than  average  aspect  ratio. 

To  illustrate  some  of  this  variation,  we  can  examine  the  construction  of  the  tip 
region  in  three  species — Eudiscopus  denticulus,  Lasiurus  cinereus,  and  Mimetillus 
moloneyi.  The  aspect  ratio  for  the  tip  followed  by  the  tip  index  (in  parentheses)  for 
each  of  these  species  is  5.98  (2.16),  5.04  (1.99),  and  4.41  (1.59),  respectively.  In 
Eudiscopus ,  the  third  metacarpal  is  proportionately  short  (41.82  per  cent  of 
digital  length),  the  bulk  of  the  length  being  contributed  by  the  phalangeal  elements, 
especially  the  third  phalanx.  The  fourth  and  fifth  metacarpals  are  proportionately 
longer  (61.55  and  64.55  per  cent,  respectively)  than  the  third,  but  nearly  half 
the  length  of  each  of  these  digits  is  accounted  for  by  the  phalanges.  In  Lasiurus, 
the  metacarpals  are  proportionately  longer  (54.07,  65.01,  and  70.63  per  cent, 
respectively)  than  in  Eudiscopus,  and  the  first  and  second  phalanx  of  digit  III 
account  for  most  of  the  remaining  length  of  this  digit;  the  third  phalanx  is  markedly 
shortened.  The  phalanges  of  digits  IV  and  V  are  nearly  equal  in  length.  The  third 
metacarpal  of  Mimetillus  is  proportionately  longer  than  that  of  either  of  the  two 
aforementioned  species  (63.40  per  cent  of  the  length).  The  third  phalanx  of  digit 
III  is  indistinguishable,  and  the  remaining  two  are  about  equal  in  length.  The 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


283 


majority  of  the  length  of  the  fourth  and  fifth  digits  is  contributed  by  the  metacarpal 
elements  (73.59  and  82.35  per  cent,  respectively).  The  second  phalanx  of  digit 
IV  is  much  reduced,  (comprising  less  than  five  per  cent  of  the  length  of  the  digit). 
Both  phalanges  of  digit  V  are  extremely  short  and  equal  or  subequal  in  length. 
Together,  they  comprise  17.65  per  cent  of  the  length  of  this  digit. 

These  three  species  are  only  exemplary  of  the  kinds  of  variation  that  exist 
within  the  family  Vespertilionidae.  This  would  seem  to  confirm  the  wide  variety 
of  flight  behaviors  reported  for  the  family,  which  range  from  the  swift, 
sustained  flight  of  migratory  species  to  the  erratic,  highly  maneuverable  flight  of 
some  of  the  smaller  nonmigratory  species.  Norberg  (1972,  1976a,  1976/?)  has 
clearly  demonstrated  the  hovering  ability  of  Plecotus  auritus,  and  certainly  other 
species  will  be  shown  to  possess  this  flight  behavior. 

Mystacinidae 

The  group  centroid  for  this  rather  unusual,  monotypic  family  ordinates  into  the 
upper  right-hand  quadrant  of  discriminant  space  (Figs.  4D-E,  5,  6).  As  we  have 
noted  above,  this  portion  of  discriminant  space  is  defined  generally  by  a  relatively 
long  and  broad  chiropatagium  and  relatively  short  and  broad  plagiopatagium. 
The  centroid  of  Mystacina  tuberculata  is  most  closely  associated  with  that  of  the 
Mormoopidae  in  the  first  two  canonical  axes  (Fig.  6).  However,  interactions  among 
variable  vectors  in  the  third  canonical  axis  (Figs.  4E,  5)  cause  a  rather  marked 
dissociation  of  these  two  centroids,  suggesting  basic  differences  in  the  composition 
of  the  wings  of  these  two  families. 

The  effect  of  the  vector  for  length  of  forearm  (B),  albeit  weak  as  compared  to 
that  of  other  groups  of  bats,  is  somewhat  stronger  in  discrimination  of  Mystacina 
than  in  mormoopids  (Table  6).  In  both  groups  and  in  the  first  two  canonical  axes 
(Figs.  4D,  6),  this  variable  vector  generally  suggests  longness.  In  the  third  canonical 
axis  (Figs.  4E,  5),  the  mystacinid  centroid  appears  to  be  more  strongly  influenced 
by  the  tail  (smallness)  end  of  this  vector,  whereas  the  mormoopid  centroid  is 
aligned  closer  to  the  positive  (longness)  end.  The  relative  length  of  the  forearm 
of  Mystacina  ranks  slightly  below  the  mean  for  all  bats:  mormoopids  rank  above 
this  mean  (Table  A8).  This  indicates  a  somewhat  greater  length  of  head  and  body 
for  Mystacina  as  compared  with  that  of  mormoopids. 

Interactions  among  variable  vectors  associated  with  length  of  digit  III  (C,  D,  E, 
F)  of  Mystacina  are  similar  to  those  discussed  for  mormoopids.  The  vector  for 
the  length  of  the  third  metacarpal  (C)  of  both  these  families  implies  large  size 
(Figs.  4D-E,  5,  6).  The  proportional  length  of  this  wing  element  is  slightly  greater 
in  Mystacina  than  in  mormoopids  and,  in  both,  contributes  more  than  50  per 
cent  to  the  length  of  digit  III  (Table  A12).  As  appears  to  be  typical  of  bats  arrayed 
in  this  portion  of  discriminant  space,  the  vector  for  the  length  of  the  first  phalanx 
of  digit  III  (D)  suggests  shortness.  This  wing  element  comprises  only  14.33  per 
cent  of  the  length  of  the  third  digit  in  Mystacina,  which  is  only  slightly  higher  than 
that  contributed  in  mormoopids  (Table  A 13).  These  two  variable  vectors  appear 
to  be  important  in  the  group  discrimination  vectors  of  both  Mystacina  and  mor¬ 
moopids  (Table  6).  Although  the  variable  vectors  for  the  two  distal  phalanges 


284 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


(E,  F)  of  both  families  suggest  long  length,  the  contribution  of  each  of  these  ele¬ 
ments  to  the  wing  of  these  two  groups  is  somewhat  different.  The  proportional 
lengths  of  all  three  phalangeal  elements  are  maintained  nearly  equal  or  subequal 
in  the  wing  of  Mystacina  (Fig.  9;  Tables  A13-15).  On  the  other  hand,  there  ap¬ 
pears  to  be  a  definite  allometric  relationship  among  these  phalangeal  elements 
in  the  wing  of  mormoopids.  The  relative  length  of  the  third  digit  of  the  mystacinid 
wing  lies  below  the  average  for  all  bats  (Table  A9).  Likewise,  the  tip  index  of 
Mystacina  is  below  the  average  computed  for  all  bats  (Table  A2).  However,  as 
noted  above,  the  long  forearm  tends  to  mask  the  length  of  digit  III  in  these  bats 
as  well  as  in  emballonurids  and  mormoopids. 

The  effects  of  the  vectors  for  elements  of  the  fourth  digit  (G,  H,  I)  of  Mystacina 
are  similar  to  those  discussed  for  mormoopids.  The  vector  for  the  length  of  the 
fourth  metacarpal  (G)  indicates  shortness  in  the  first  two  canonical  axes  (Figs. 
4D,  6),  but  a  slightly  positive  (longness)  influence  is  suggested  in  the  third  axis 
(Figs.  4E,  5).  The  proportional  length  of  this  element  is  well  above  the  average 
for  all  bats  (Table  A16).  The  variable  vectors  for  the  phalangeal  elements  of 
digit  IV  (H,  I)  both  imply  longness.  In  terms  of  the  group  discrimination  vector, 
the  variable  vector  for  the  first  phalanx  of  this  digit  (H)  appears  to  be  important 
(Table  6).  The  proportional  length  of  the  second  phalanx  of  digit  IV  ranks  slightly 
above  the  mean  for  all  bats  and  this  element  contributes  21.04  per  cent  of  the 
length  of  the  digit  (Table  A 18). 

The  greatest  differences  in  composition  of  the  wing  of  Mystacina  and  that  of 
mormoopids  concern  features  of  the  fifth  digit.  In  Mystacina ,  variable  vector  for 
the  length  of  the  fifth  metacarpal  (J)  suggests  longness  in  the  first  two  canonical 
axes  (Figs.  4D,  6).  However,  the  implication  shifts  toward  smallness  in  the  third 
axis  (Figs.  4E,  5).  Paradoxically,  the  proportional  length  of  this  wing  element 
(67.42)  ranks  well  above  the  average  for  all  bats  (61.02),  whereas  that  for  mor¬ 
moopids  (59.29)  falls  below  the  average  (Table  A19).  This  variable  vector  ap¬ 
pears  to  be  relatively  unimportant  in  the  discrimination  of  the  Mystacinidae 
(Table  6).  The  strongest  vectors  in  this  regard  are  those  for  lengths  of  the  first  and 
second  phalanx  of  digit  V  (K  L).  The  vector  for  the  first  phalanx  (K)  strongly 
suggests  shortness  in  all  three  canonical  axes  (Figs.  4D-E,  5,  6).  The  proportional 
length  of  this  element  averages  the  shortest  among  all  bats  (Table  A20);  mor¬ 
moopids  rank  above  the  overall  average  with  regard  to  this  feature.  On  the  other 
hand,  the  vector  for  the  length  of  the  second  phalanx  of  digit  V  (L)  strongly 
implies  longness  and  this  element  contributes  21.00  per  cent  to  the  length  of  this 
digit  (Table  A21). 

The  overall  aspect  ratio  of  the  wing,  as  well  as  that  of  the  tip,  of  Mystacina  falls 
slightly  below  the  average  of  all  bats  (Tables  A3,  A4).  However,  the  relatively 
long  forearm  and  comparatively  short  fifth  digit  contribute  to  the  higher  than 
average  aspect  ratio  of  the  plagiopatagium  (Table  A5). 

Little  is  known  concerning  the  flight  behavior  of  Mystacina  tuberculata.  The 
family  is  endemic  to  New  Zealand  where  it  and  Chalinolobus  tuberculatus 
(Vespertilionidae)  comprise  the  total  chiropteran  fauna.  The  phylogenetic  re¬ 
lationships  of  the  family  are  poorly  understood  although  relationship  to  the 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


285 


Molossidae  has  been  suggested  by  various  authors  (Dobson,  1875,  Miller,  1907). 
In  terms  of  wing  shape,  Mystacina  most  closely  resembles  mormoopids  and 
phyllostomatids.  This  is  particularly  interesting  in  view  of  Daniel’s  (1976)  recent 
report  on  the  food  habits  of  Mystacina  in  which  he  included  fruit  and  possibly 
nectar  along  with  aerial  and  terrestrial  insects  in  the  feeding  regime.  If  the  mor¬ 
phometric  resemblance  between  Mystacina,  mormoopids,  and  phyllostomatids 
is  conveyed  in  functional  similarity,  the  wing  of  Mystacina  should  be  found  to  be 
relatively  versatile. 


Molossidae 

The  shape  of  the  wing  in  this  family  is  perhaps  the  most  distinctive  among  all 
bats.  The  molossid  wing  is  extremely  narrow  and  has  an  unusually  long  tip  region 
(Fig.  9).  As  a  consequence,  the  wing  is  highest  in  overall  aspect  ratio  among 
bats.  We  have  already  discussed  some  features  of  molossid  wings  in  the  accounts 
of  emballonurids  and  noctilionids.  Of  particular  interest  is  the  fact  that,  even 
though  the  bats  of  these  three  families  possess  wings  of  high  aspect,  the  mode  by 
which  their  wings  are  constructed  is  markedly  different. 

Whereas  the  forearm  is  usually  long  in  most  other  groups  of  bats,  especially  those 
that  possess  high  aspect  wings,  the  relative  length  of  the  forearm  of  molossids 
averages  the  shortest  among  all  bats  (Table  A8).  The  vector  for  this  variable  (B) 
is  oriented  almost  directly  away  from  the  group  centroid  in  the  first  three  canonical 
axes  and  thereby  suggests  shortness  (Figs.  4D-E,  5,  6).  The  forearm  contributes 
only  30  to  35  per  cent  to  the  total  span  of  the  wing.  Among  molossids,  Cheiromeles, 
Otomops,  and  Eumops  possess  the  largest  forearms,  whereas  Sauromys  and 
Molossus  have  the  shortest.  As  the  orientation  and  length  of  the  variable  vector 
indicate,  the  length  of  the  forearm  is  an  important  factor  in  the  group  dis¬ 
crimination  vector  (Table  6). 

The  great  length  of  the  wing  is  reflected  in  the  generally  positive  orientation  of 
all  vectors  associated  with  elements  of  the  third  digit  toward  the  molossid  centroid 
(Figs.  4D-E,  5,  and  6).  The  vectors  for  the  metacarpal,  and  second  and  third 
phalanges  (C,  E,  F)  are  not  as  positively  associated  with  the  molossid  centroid  as 
was  noted  for  the  noctilionids,  mormoopids,  phyllostomatids,  and  vespertilionids. 
Nonetheless,  these  vectors  do  imply  longness  of  these  elements  in  the  Molossidae. 
The  vector  for  the  first  phalanx  of  digit  III  (D)  strongly  suggests  longness  in  the  first 
two  canonical  axes  and  to  a  certain  extent  in  the  third  axis.  Proportionately,  the 
length  of  this  phalanx  (19  to  26  per  cent  of  the  length  of  digit  III)  averages  among 
the  largest  for  all  bats  (Table  A1 3).  Although  the  proportional  length  of  the  second 
phalanx  averages  below  the  mean  for  all  bats  (Table  A 14),  these  two  phalangeal 
elements  in  consort  with  the  metacarpal  produce  the  major  portion  of  the  span 
of  digit  III.  It  is  difficult  to  interpret  the  vector  for  the  third  phalanx  of  this  digit 
because,  by  comparison,  it  is  rather  short.  However,  this  vector  appears  to  be 
rather  important  in  the  group  discrimination  vector  (Table  6).  In  this  case,  the 
vector  seems  to  imply  simple  presence  of  the  phalanx  rather  than  length.  Shortness 
or  absence  of  the  distal  phalanx  of  digit  III  seems  to  be  the  case  in  other  families 


286 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


that  have  proportionately  long  first  and  second  phalanges  (for  example,  embal- 
lonurids,  noctilionids,  and  vespertilionids). 

In  most  of  the  other  groups  of  bats  considered  in  this  study,  vectors  associated 
with  elements  of  the  fourth  digit  (G,  H,  I)  are  not  easily  interpreted,  mostly 
because  of  their  tangential  orientation  to  centroids  and  their  synergistic  inter¬ 
action  with  other  variables.  All  three  of  these  vectors  are  directed  toward  the 
molossid  centroid  and  all  imply  longness.  The  most  powerful  among  these  are  the 
vectors  for  length  of  metacarpal  (G)  and  second  phalanx  (I).  In  addition,  these 
two  vectors  are  important  in  the  group  discrimination  vector  (Table  6).  The 
relative  length  of  the  fourth  digit  is  not  particularly  impressive  and  it  averages 
below  the  mean  for  all  bats  (Table  A 10).  However,  this  value  is  greatly  masked  by 
the  generally  long  length  of  head  and  body  of  these  bats.  The  fourth  digit  of 
molossids  constitutes  nearly  60  per  cent  of  the  span  of  the  wing,  and  in  these  terms, 
is  the  largest  among  all  bats.  The  metacarpal  alone  contributes  55  to  69  per  cent 
of  the  length  of  this  digit  (Table  A16).  The  first  phalanx  constitutes  the  bulk  of 
the  remaining  length  (18  to  28  per  cent,  Table  A17).  The  length  of  the  second 
phalanx  of  digit  IV  is  variable  and  can  contribute  as  much  as  18.11  per  cent 
( Sauromys )  or  as  little  as  2.88  and  3.94  ( Tadarida  and  Promops,  respectively) 
to  the  length  of  this  digit.  Eumops  and  Molossus,  on  the  average,  possess  a  rather 
short  second  phalanx  on  digit  IV. 

Unusually  long  third  and  fourth  digits  have  been  discussed  in  the  accounts 
of  several  groups,  especially  the  Phyllostomatidae,  which  have  generally  long- 
tipped,  low  aspect  wings.  Perhaps  the  most  striking  feature  of  the  molossid  wing 
is  the  markedly  short  fifth  digit,  which  converts  the  long  tip  region  into  a  high 
aspect  surface.  The  vector  for  the  length  of  the  fifth  metacarpal  (J)  strongly  implies 
shortness  in  the  first  three  canonical  axes  (Figs.  4D-E,  5,  6).  Similarly,  the  vectors 
for  the  lengths  of  the  two  phalangeal  elements  of  this  digit  (K,  L)  orient  away 
from  the  molossid  centroid  and  thereby  imply  shortness.  All  three  of  these 
variable  vectors  are  important  factors  in  the  discrimination  of  the  group  (Table 
6). 

In  other  high  aspect  wings  such  as  those  of  emballonurids  and  noctilionids, 
the  shortening  of  the  fifth  digit  is  accomplished  by  shortening  the  phalangeal 
elements  while  maintaining  the  metacarpal  more  or  less  isometric  with  the  third 
and  fourth  metacarpals.  If  the  apparent  versatility  in  flight  behavior  of  these 
bats  is  any  indication,  we  could  assume  the  formation  of  a  high  aspect  wing  in  this 
fashion  to  be  a  less  than  total  commitment  to  swift  flight.  On  the  other  hand,  by 
shortening  the  fifth  metacarpal,  molossids  gain  dexterous  control  of  a  smaller 
portion  of  the  camberable  surface  but  at  the  same  time  might  lose  a  sizable  degree 
of  flight  versatility.  In  this  light,  it  is  interesting  to  note  that  the  genus  Tadarida 
(the  most  diverse,  yet  least  specialized,  of  the  family  with  some  45  or  so  species) 
exhibits  a  wide  range  of  variation  in  the  composition  of  the  fifth  digit  and  other 
digital  elements. 

To  illustrate  the  degree  of  variation  in  wing  composition  within  the  Molossidae, 
we  have  used  Cheiromeles ,  Otomops,  Sauromys,  and  Tadarida.  Cheiromeles 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


287 


torquatus  is  the  largest  molossid,  with  a  head  and  body  length  of  115  to  135 
millimeters  and  a  weight  of  150  to  170  grams.  The  proportional  lengths  of  the 
metacarpal  elements  of  digits  III  to  V  are  shortest  among  the  family  (43.21, 
54.62,  and  49.37  per  cent  contribution  to  digital  length,  respectively).  On  the  other 
hand,  the  phalanges  of  the  third  and  fourth  digits  are  proportionately  longer  than 
those  of  any  other  molossid.  The  first  phalanx  of  the  fifth  digit  is  proportionately 
longer  than  that  of  any  other  molossid,  comprising  nearly  40  per  cent  of  the  length 
of  the  digit;  the  second  phalanx  is  about  average  for  the  family  (1 1.50  per  cent  of 
digital  length). 

On  the  average,  Otomops  possesses  the  proportionately  longest  third  and 
fourth  metacarpals  of  any  molossid  (54.42  and  68.78  per  cent  of  the  digital  length, 
respectively),  although  individual  species  of  Tadarida  and  Eumops  possess  longer 
fourth  metacarpal  elements  (72.52  and  70.18  per  cent,  respectively).  The  pro¬ 
portional  lengths  of  phalangeal  elements  vary  in  Otomops.  Generally  the  major 
portion  of  the  length  of  digit  III  is  contributed  by  the  second,  first,  and  third 
phalanx  (21.18,  20.59,  and  4.65  per  cent,  respectively).  Otomops  possesses  the 
shortest  first  phalangeal  element  of  digit  IV  of  the  family  (17.82  per  cent  of  the 
digital  length),  and  the  proportional  length  of  the  second  phalanx  (7.21  per  cent) 
is  well  below  the  average  for  the  family.  Whereas  the  metacarpal  of  digit  V  is  ex¬ 
tremely  short,  the  proportional  length  of  the  phalanges  of  this  digit  are  nearly  the 
largest  for  the  family  (35.27  and  12.57  per  cent,  respectively). 

Whereas  the  two  genera  discussed  above  might  be  considered  as  among  the  more 
specialized  molossids,  Sauromys  appears  to  be  among  the  least  specialized.  The 
metacarpal  elements  of  digits  III  and  IV  are  proportionately  average  for  the  fam¬ 
ily  (50.81  and  59.05  per  cent,  respectively);  the  fifth  metacarpal  is  unusually  long 
for  the  family  (63.56  per  cent  of  the  digital  length).  The  proportional  lengths  of 
the  first  and  second  phalangeal  elements  of  digits  III  to  V  vary  although  they  are 
generally  isometric  and  range  between  22  and  1 5  per  cent  of  the  digital  length.  The 
third  phalanx  of  digit  III  is  proportionately  long  for  the  family  (7.93  per  cent  of 
the  digital  length). 

Finally,  Tadarida  is  perhaps  the  most  variable  among  the  molossids  in  terms 
of  wing  composition.  The  proportional  lengths  of  the  metacarpals  of  digits  III  to 
V  rank  near  the  family  average,  but  the  range  is  broad  (53.97  to  46.02,  72.52  to 
57.16,  and  67.51  to  53.55  per  cent  of  the  digital  length,  respectively).  There  is  a 
general  trend  of  isometry  among  the  proportional  lengths  of  the  first  and  second 
phalanx  of  digit  III  (20.59  to  19.81  and  23.34  to  18.02  per  cent,  respectively). 
The  proportional  length  of  the  third  phalanx  of  this  digit  varies  (10.80  to  4.92 
per  cent).  With  regard  to  the  phalanges  of  the  fourth  digit,  patterns  of  allometry 
and  isometry  vary  markedly,  especially  with  respect  to  proportional  length  of 
the  second  phalanx  (27.10  to  19.94  and  20.49  to  2.88  per  cent  of  digital  length, 
respectively).  Freeman  (1977)  noted  this  allometric  variation  in  the  composition 
of  digit  IV  and  interpreted  it  in  terms  of  zoogeographic  distribution.  Allometry 
is  even  more  pronounced  in  the  proportional  lengths  of  the  phalangeal  complement 
of  the  fifth  digit  (36.43  to  21.81  and  15.05  to  7.97  per  cent,  respectively). 


288 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Generally  speaking,  the  wings  of  molossids  are  highly  specialized,  although 
we  remind  the  reader  that  within  this  family  a  degree  of  variability  exists.  Wing 
loading  is  normally  high  (Table  A6):  Eumops  averages  highest  in  the  family  (28.47 
Nt/m2;  Eumops  auripendulus  is  highest  among  all  bats  with  58.00  Nt/m2);  and 
Tadarida,  although  nearly  average  in  this  feature  (19.54  Nt/m2),  exhibits  loadings 
down  to  8.43  Nt/m2.  The  general  composition  of  the  molossid  wing  suggests  a 
reduction  in  the  control  of  camberable  surface.  No  doubt  the  “automatic”  flexing 
and  extending  devices  in  the  elbow  and  shoulder  regions  discussed  by  Vaughan 
( 1 959,  1 970a)  relate  to  this  alar  composition.  We  suspect  that  the  more  generalized 
species  of  the  family  will  be  shown  to  have  a  greater  degree  of  “manual”  control 
of  their  flight  surfaces. 


Classification 

As  has  been  discussed  above,  in  the  discriminant  analysis  a  discrimination 
vector  is  computed  for  each  group  (in  this  case  families  or  subfamilies)  based  on 
the  synergistic  interaction  among  variables.  In  the  classification  phase  of  the 
analysis,  each  case  (species  in  this  analysis)  is  scrutinized  and  assigned  to  that 
group  to  which  it  is  most  closely  aligned  in  discriminant  space  (Fig.  17).  Inasmuch 
as  the  discrimination  vector  for  each  group  is  an  expression  of  the  complex 
qualitative  and  quantitative  aspects  of  wing  shape,  species  are  grouped  together 
based  on  similarity  of  wing  shape. 

In  the  overall  classificatory  analysis,  only  14  of  466  species  (three  per  cent) 
were  incorrectly  assigned.  The  high  degree  of  correct  associations  appears  to 
indicate  a  rather  large  phylogenetic  component  in  the  overall  shape  of  bat  wings. 
“Misclassifications”  may  be  attributed  to  several  possible  sources  of  error. 

The  first  of  these  is  insufficient  sample  size,  which  could  have  greatly  effected 
the  formulation  of  an  accurate  discrimination  vector  for  various  groups.  We 
suspect  this  might  be  the  case  with  regard  to  the  Thyropteridae  and  Myzapodidae, 
in  which  the  sample  sizes  were  extremely  small.  We  would  not  be  particularly 
surprised  if  the  association  of  these  two  families  with  the  Vespertilionidae  was 
found  not  to  be  related  to  the  sample  size,  because  the  shape  of  the  wing  in  these 
two  families  is  in  fact  similar  to  that  of  the  vespertilionids.  Another  source  of 
error  involves  “leakage”  of  taxa  that  ordinate  close  to  the  grand  centroid  for  all 
bats.  This  we  suspect  is  the  explanation  for  most  of  the  “misclassifications” 
encountered  in  the  Phyllostomatidae. 

Yet  another  source  of  error  might  be  that  of  functional  similarity.  With  regard 
to  the  two  species  assigned  to  the  Megadermatidae,  as  well  as  Hypsignathus 
monstrosus  (Pteropodidae)  and  Rhinolophus  Indus  (Rhinolophidae),  it  is  note¬ 
worthy  to  point  out  that  each  has  a  relatively  long  second  phalanx  in  digit  III, 
which  is  a  major  feature  of  megadermatids.  Similarly,  the  association  of  Pteronotus 
parnellii  with  the  Phyllonycterinae  appears  to  relate  to  the  overall  similarity  of 
alar  shape,  especially  with  respect  to  length  of  forearm.  As  noted  above,  this 
association  also  may  reflect  some  phylogenetic  similarity  between  mormoopids 
and  phyllostomatids.  One  molossid  species,  Tadarida  loriae ,  is  classified  as  a 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


PTEROP 
RHINOP 
CRASEO 
EMBALL 
RHINOL 
NYCTER 
MEGAD 
NOCTIL 
MORMO 
PHYLOS 
GLOSSO 
CARO  LL 
STENOD 
PHYNYC 
DESMOD 
NATAL  L 
THYROP 
MYZAPO 
VESPER 
MYSTAC 
MO  LOS 

Fig.  17. — Classification  graph  from  the  discriminant  analysis.  Numbers  on  the  diagonal 
represent  number  of  species  correctly  associated  by  the  group  discrimination  vector  for  each 
group,  with  their  respective  taxonomic  category.  Numbers  in  rows,  and  off  the  diagonal, 
represent  number  of  “misclassifications"  to  other  taxonomic  groups.  This  analysis  resulted  in 
97  per  cent  correct  associations.  See  text  for  discussion. 

vespertilionid.  This  is  not  surprising  because  other  generalized  species  of  Tadarida 
are  ordinated  toward  the  vespertilionid  dispersion. 

The  Phyllostomatidae,  as  a  whole,  illustrates  a  rather  high  affinity  to  its  various 
group  discrimination  vectors;  only  4.32  per  cent  of  its  species  are  assigned  outside 
the  limits  of  the  family.  However,  within  the  family  there  is  a  relatively  high 
percentage  of  “misclassification”  (22.30  per  cent);  this  could  reflect  phylogenetic 
infidelity  or,  again,  it  simply  might  be  attributable  to  functional  similarities 
in  wing  shape. 

The  Desmodontinae  is  the  only  phyllostomatid  subfamily  that  does  not  exhibit 
a  “misclassification.”  However,  three  species  of  Phyllostomus  are  confused  as 


289 


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290 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


desmodontines.  Of  these,  two  different  samples  of  P.  discolor  follow  this  trend 
with  91  to  50  per  cent  affinity,  respectively,  to  the  discrimination  vector  of 
vampires.  Phyllostomus  latifolius  exhibits  44  per  cent  affinity  and  P.  hastatus, 
in  two  separate  analyses,  showed  100  per  cent  affinity  with  this  subfamily.  A 
tentative  explanation  of  this  might  be  that  these  large-sized  phyllostomatines  have 
flight  requirements  similar  to  those  of  vampires  (high  weight-bearing  capacity) 
and  hence  wings  of  similar  shape. 

The  glossophagines  illustrate  the  tightest  packing  of  taxa  among  the  phyl- 
lostomatids.  Only  two  species,  Hylonycteris  underwoodi  and  Platalina  gen- 
ovensium,  are  associated  outside  of  the  group.  It  is  difficult  to  assess  their  re¬ 
lationship  with  the  stenodermines  other  than  to  say  that  these  two  species  appear 
to  be  similar  to  Vampyrops  and  Sturnira. 

The  stenodermines,  although  not  so  tightly  packed,  occupy  a  fairly  discrete 
portion  of  discriminant  space.  The  seven  “misclassified”  species  are  located  in  the 
congested  region  near  the  grand  centroid  for  all  bats.  The  Carolliinae  is  practically 
engulfed  in  this  congestion,  and  they  show  little  fidelity  to  their  group  discrimi¬ 
nation  vector.  The  fact  that  this  congested  area  exists  and  that  it  is  composed 
primarily  of  phyllostomatines  would  suggest  the  generalized  nature  of  the  wings 
of  this  subfamily.  Also  this  seems  generally  to  support  the  basal  assignment  of 
this  group  in  terms  of  phylogenetic  relationships  within  the  family. 

Conclusions 

As  stated  in  our  introductory  comments,  the  wide  range  of  variation  and 
complicated  nature  of  the  interactions  among  the  intrinsic  wing  elements  of 
chiropteran  species  makes  impossible  a  precise  and  definitive  explanation  of 
wing  shape.  However,  the  essence  of  wing  shape  and  the  variables  that  affect 
it  can  be  perceived  in  multidimensional  space  using  such  multivariate  procedures 
as  discriminant  analysis.  This  study  has  been  as  much  an  analysis  of  chiropteran 
wings  as  it  has  been  an  example  of  this  morphometric  procedure. 

The  interactions  among  the  variables  utilized  in  this  study  are  summarized 
below. 

1.  Length  of  head  and  body  appears  to  have  little  effect  on  the  shape  of 
chiropteran  wings.  Generally  speaking,  bats  tend  to  possess  wings  that  range 
between  one  and  one  half  and  two  and  one-half  times  the  length  of  the  head  and 
body.  Extremes  in  excess  of  three  times  the  length  of  head  and  body  were  noted 
among  the  Emballonuridae.  Whereas  small-sized  bats  tend  to  have  longer  wings 
with  lighter  loading  than  do  larger  bats,  there  is  a  great  deal  of  variation  and  the 
picture  appears  to  be  more  complex  than  simple  bivariate  analysis  indicates. 
We  do  not  believe  that  the  questions  of  mass,  area,  and  wing  shape  have  been 
adequately  dealt  with,  and  certainly  these  considerations  were  beyond  the  focus 
of  our  analysis.  We  suggest  that  these  questions  will  require  further  analysis  under 
free-flight  conditions. 

2.  Lengths  of  forearm  and  digit  III  certainly  constitute  the  majority  of  the 
wing  span.  However,  derived  variables  that  describe  their  relative  proportions 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


291 


(such  as  tip  index)  do  not  adequately  represent  their  influence  on  wing  shape.  We 
have  shown  that  the  forearm  can  be  relatively  short  or  relatively  long  and  in 
conjunction  with  the  span  of  digit  III  produce  a  wing  of  similar  or  different  shape. 
Pteropodids,  emballonuroids,  and  rhinolophoids  tend  to  emphasize  the  long 
length  of  forearm  in  wing  construction.  The  remaining  chiropteran  families 
generally  possess  shorter  forearms. 

3.  The  composite  length  of  digit  III  can  be  relatively  short  (Rhinopomatidae) 
or  long  (Phyllostomatidae  and  Molossidae).  The  interactions  among  the  bony 
elements  that  comprise  the  length  of  the  third  digit  are  extremely  complex. 
Chiropteran  families  are  ordinated,  rather  markedly,  into  two  general  groups  in 
discriminant  space  by  the  presence  or  absence  of  the  third  phalanx  of  this  digit. 
However,  wing  tips  of  nearly  equal  proportional  length  are  achieved  by  members 
of  both  groups.  Those  that  possess  long  wing  tips,  the  phyllostomatoids  and 
vespertilionoids  (except  molossids)  tend  to  have  a  lengthened  second  phalangeal 
element  of  digit  III.  The  phyllostomatoids  generally  have  a  lengthened  third 
phalanx  as  well  and  a  shortened  first  phalanx.  Vespertilionoids  (except  molossids) 
tend  to  possess  a  lengthened  first  and  second  phalanx,  in  an  isometric  fashion, 
and  have  a  shortened  terminal  phalanx  of  digit  III.  Molossids  follow  the  general 
pattern  of  vespertilionoids,  but  also  have  a  lengthened  metacarpal  of  this  digit. 
Those  bats  with  generally  short  wing  tips  illustrate  an  allometric  mixture  in  the 
composition  of  digit  III.  Most,  with  the  notable  exceptions  of  the  Pteropodidae  and 
Craseonycteridae,  possess  a  moderately  long  metacarpal.  However,  most  of 
the  span  of  digit  III  is  contributed  by  a  relatively  long  second  phalanx  or,  in  some 
cases,  moderately  long  first  and  second  phalanges  of  nearly  equal  length. 

4.  The  effect  of  the  fourth  digit  on  the  shape  of  the  wing  is  complex  and,  in 
most  cases,  the  influence  of  its  elements  are  involved  in  an  overall  synergism  among 
variables.  In  those  bats  with  low  aspect  tip  regions,  the  length  of  this  digit  is 
intermediate  between  digits  III  and  V.  The  fourth  digit  is  relatively  long  in  the  high 
aspect  wing  tips  of  noctilionids  and  molossids;  in  those  of  emballonurids  this 
digit  is  shortened.  The  composition  of  the  fourth  digit  also  varies.  In  the  phyl¬ 
lostomatoids,  the  metacarpal  is  moderately  long  and  has  proportionately  length¬ 
ened  first  and  second  phalanges.  The  terminal  phalanx  is  especially  long  in 
noctilionids.  The  metacarpal  element  is  lengthened  in  emballonuroids  and 
rhinolophoids,  and  the  first  phalanx  also  tends  to  be  proportionately  long.  In 
the  Pteropodidae,  the  metacarpal  is  markedly  shortened,  and  the  length  of  the 
digit  is  produced  by  proportionately  long  first  and  second  phalangeal  elements. 
The  long  fourth  digit  of  molossids  is  comprised  of  the  long  metacarpal  and  first 
phalanx. 

5.  Whereas  digit  III  is  important  in  determining  the  span  of  the  wing,  digit  V 
determines  the  chord.  The  interactions  between  this  digit  and  other  wing  com¬ 
ponents  in  determining  shape  are  somewhat  dualistic  in  nature.  The  aspect 
ratio  of  the  plagiopatagium  can  be  affected  either  by  a  lengthening  or  shortening 
of  digit  V  or  the  forearm.  Thus,  in  the  Rhinopomatidae  and  Emballonuridae, 
a  relatively  long  forearm  in  combination  with  a  moderately  long  fifth  digit  produces 


292 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


a  high  aspect  plagiopatagium.  On  the  other  hand,  in  the  Molossidae,  shortening 
of  both  elements  produces  an  aspect  ratio  of  similar  or  higher  magnitude.  The 
interaction  of  digit  III  with  the  fifth  digit  yields  a  tip  region  of  high  aspect  in  the 
Emballonuridae  and  Molossidae;  by  comparison,  that  of  rhinopomatids  is  low 
in  aspect.  In  the  Phyllostomatidae,  a  long  fifth  digit  tends  to  offset  the  effects 
of  the  long  span  of  digit  III  and,  in  combination  with  a  relatively  short  forearm, 
produces  an  overall  low  aspect  wing.  The  composition  of  the  fifth  digit,  like  that 
of  digits  III  and  IV,  varies  from  group  to  group.  Most  bats  lengthen  or  shorten 
the  fifth  digit  by  differentially  lengthening  or  shortening  phalangeal  elements; 
most  taxa,  especially  molossids,  retain  a  moderately  long  metacarpal.  The 
pteropodids,  as  we  have  noted,  have  markedly  short  metacarpal  elements  in  all 
three  digits.  Of  the  Microchiroptera,  the  molossids  illustrate  the  most  drastic 
proportional  shortening  of  the  fifth  metacarpal. 

6.  Finally,  we  reemphasize  that  although  the  overall  shape  of  the  wing 
(silhouette)  may  be  important  from  the  standpoint  of  such  aerodynamic  features 
as  wetted  surface  area  and  wing  loading,  it  is  the  internal  composition  of  the 
wing  that  determines  the  camberability  and  ultimately  the  dynamics  of  lifting 
potential.  Far  too  little  is  known  concerning  the  comparative  aspects  of  actual 
free-flight  behavior  of  bats  to  permit  meaningful  functional  interpretation  of 
wing  shape.  It  is  to  this  end  that  we  suggest  future  morphometric  analyses  be 
directed,  for  without  this,  functional  speculations  can  only  be  misleading  and  may 
further  confound  an  understanding  of  mammalian  flight. 

Literature  Cited 

Atchley,  W.  R.  1978.  Ratios,  regression  intercepts,  and  the  scaling  of  data.  Syst. 
Zool.,  27:78-83. 

Atchley,  W.  R„  and  D.  Anderson.  1978.  Ratios  and  the  statistical  analysis  of  biological 
data.  Syst.  Zool.,  27:71-78. 

Atchley,  W.  R.,  C.  T.  Gaskins,  and  D.  Anderson.  1976.  Statistical  properties  of  ratios. 
I.  Empirical  results.  Syst.  Zool.,  25:137-148. 

Bader,  R.  S.,  and  J.  S.  Hall.  1960.  Osteometric  variation  and  function  in  bats.  Evolu¬ 
tion,  14:8-17. 

Cooley,  W.  W.,  and  P.  R.  Lohnes.  1971.  Multivariate  data  analysis.  John  Wiley  and 
Son,  New  York,  xii-f  364  pp. 

Daniel,  M.  J.  1976.  Feeding  by  the  short-tailed  bat  ( Mystacina  tuberculata )  on  fruit 
and  possibly  nectar.  New  Zealand  J.  Zool.,  3:391-398. 

Davis,  R.  1969.  Wing  loading  in  pallid  bats.  J.  Mamm.,  50:140-144. 

Dixon,  W.  J.  (ed.).  1973.  Biomedical  computer  programs.  Univ.  California  press, 

Berkeley,  3rd  ed.,  viii  +  773  pp. 

Dobson,  G.  E.  1875.  Conspectus  of  the  suborders,  families,  and  genera  of  Chiroptera 
arranged  according  to  their  natural  affinities.  Ann.  Mag.  Nat.  Hist.,  ser.  4, 
16:345-357. 

Dwyer,  P.  D.  1965.  Flight  patterns  of  some  eastern  Australian  bats.  Victoria  Nat., 
82:36-41. 

Eisentraut,  M.  1936.  Beitrag  zur  Mechanik  des  Fledermausfluges.  Zeit.  wiss.  Zool., 
148:159-188. 

Farney,  J.,  and  E.  Fleharty.  1969.  Aspect  ratio,  loading,  wing  span,  and  membrane 
area  of  bats.  J.  Mamm.,  50:362-367. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


293 


Findley,  J.  S.,  E.  H.  Studier,  and  D.  E.  Wilson.  1972.  Morphologic  properties  of  bat 
wings.  J.  Mamm.,  53:429-444. 

Freeman,  P.  W.  1977.  Multivariate  study  of  the  family  Molossidae:  functional  mor¬ 
phology  and  ecology.  Unpublished  Ph.D.  dissertation,  Univ.  New  Mexico,  xi  + 
270  pp. 

Gaisler,  J.  1964.  Comment  volent  les  chauves-souris?  Sci.  Nat.,  66: 1 1-16. 

Harrison,  D.  1964.  The  mammals  of  Arabia.  Ernest  Benn,  London,  1  :xx  +  1-192. 

Hartman,  F.  A.  1963.  Some  flight  mechanisms  of  bats.  Ohio  J.  Sci.,  63:59-65. 

Hill,  J.  E.  1974.  A  new  family,  genus  and  species  of  bat  (Mammalia:  Chiroptera)  from 
Thailand.  Bull.  British  Mus.  (Nat.  Hist.),  27:301-336. 

Jones,  C.  1967.  Growth,  development,  and  wing  loading  in  the  evening  bat,  Nycticeius 
humeral  is  (Rafinesque).  J.  Mamm.,  48: 1-19. 

Jones,  C.,  and  R.  D.  Suttkus.  1971.  Wing  loading  in  Plecotus  rafinesquii.  J.  Mamm., 
52:458-460. 

Koons,  P.  B.  1962.  Canonical  analysis.  Pp.  266-279,  in  Computer  applications  in  the 
behavioral  sciences  (H.  Borko,  ed.),  Prentice  Hall,  New  York,  xx  +  633  pp. 

Kulzer,  E.  1968.  Der  Flug  des  afrikanischen  Flughundes  Eidolon  helvum.  Natur  Mus., 
98:181-194. 

Miller,  G.  S.,  Jr.  1907.  The  families  and  genera  of  bats.  Bull.  U.S.  Nat.  Mus., 
57:xvii  + 1-282. 

Nie,  N.  H.,  C.  H.  Hull,  J.  G.  Jenkins,  K.  Steinbrenner,  and  D.  H.  Bent.  1975.  Statistical 
package  for  the  social  sciences.  McGraw  Hill,  New  York,  xxiv  +  675  pp. 

Norberg,  U.  M.  1969.  An  arrangement  giving  a  stiff  leading  edge  to  the  hand  wing  in 
bats.  J.  Mamm.,  50:766-770. 

- .  1970.  Hovering  flight  of  Plecotus  aurittis  Linnaeus.  Bijdr.  Dierk.,  40:62-66. 

- .  1972.  Bat  wing  structures  important  for  aerodynamics  and  rigidity  (Mammalia, 

Chiroptera).  Z.  Morph.  Tiere,  73:45-61. 

- .  1976a.  Some  advanced  flight  manoeuvres  of  bats.  J.  Exp.  Biol.,  64:489-495. 

- .  19766.  Aerodynamics  of  hovering  flight  in  the  long-eared  bat  Plecotus  auritus. 

J.  Exp.  Biol.,  65:459-470. 

O’Farrell,  M.  J.,  and  E.  H.  Studier.  1976.  Seasonal  changes  in  wing  loading,  body 
composition  and  organ  weights  in  Myotis  thysanodes  and  M.  lucifugus  (Chiroptera: 
Vespertilionidae).  Bull.  S.  California  Acad.  Sci.,  75:258-266. 

Orr,  R.  1954.  Natural  history  of  the  pallid  bat,  Antrozous  pallidus  (Le  Conte).  Proc. 
California  Acad.  Sci.,  28:165-246. 

Pearson,  O.,  M.  Koford,  and  A.  Pearson.  1952.  Reproduction  of  the  lump-nosed  bat 
( Corynorhinus  rafinesquei )  in  California.  J.  Mamm.,  33:273-320. 

Pennycuick,  C.  J.  1971.  Gliding  flight  of  the  dog-faced  bat,  Rousettus  aegyptiacus, 
observed  in  a  wind  tunnel.  J.  Exp.  Biol.,  55:833-845. 

Pimentel,  R.  A.  1978.  Morphometries:  multivariate  analysis  of  biological  data.  Ken¬ 
dall/Hunt  Press,  Dubuque,  Iowa,  in  press. 

Poole,  E.  1936.  Relative  wing  ratios  of  bats  and  birds.  J.  Mamm,  17:412-413. 

Revilliod,  P.  1916.  A  propos  de  1’adaptation  au  vol  chez  les  microchiropteres.  Verh. 
naturf.  Ges.  Basel,  27:156-183. 

Schnitzler,  H.-U.  1971.  Fledermause  in  Windkanal.  Z.  Vergl.  Physiol.,  73:209-221. 

Short,  H.  L.  1961.  Growth  and  development  of  Mexican  free-tailed  bats.  Southwestern 
Nat.,  6:156-163. 

Smith,  J.  D.  1972.  Systematics  of  the  chiropteran  family  Mormoopidae.  Misc.  Publ. 
Mus.  Nat.  Hist.,  Univ.  Kansas,  56:1-132. 

Struhsaker,  T.  1961.  Morphological  factors  regulating  flight  in  bats.  J.  Mamm., 
42:152-159. 

Vaughan,  T.  1959.  Functional  morphology  of  three  bats:  Eumops,  Myotis ,  Macrotus. 
Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  12:1-153. 


294 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


- .  1970a.  The  skeletal  system.  Pp.  97-138,  in  Biology  of  bats  (W.  A.  Wimsatt,  ed.), 

Academic  Press,  New  York,  l:xxii  +  1-406. 

- .  19706.  The  muscular  system.  Pp.  139-194,  in  Biology  of  bats  (W.  A.  Wimsatt, 

ed.),  Academic  Press,  New  York,  l:xii  + 1-406. 

- .  1970c.  Flight  patterns  and  aerodynamics.  Pp.  195-216,  in  Biology  of  bats 

(W.  A.  Wimsatt,  ed.),  Academic  Press,  New  York,  l:xii  +  1-406. 

Vaughan,  T.  A.,  and  G.  C.  Bateman.  1970.  Functional  morphology  of  the  forelimb  of 
mormoopid  bats.  J.  Mamm.,  51:217-235. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


295 


Appendix 

Tables  A1-A21  follow  and  consist  of  ranked  means  and  statistics  for  selected 
derived  variables.  Statistics  include  mean  for  taxa  (range  in  parentheses),  plus 
or  minus  one  standard  error  of  the  mean,  and  the  coefficient  of  variation.  Vari¬ 
able  means  are  based  on  genera  within  families  or  subfamilies,  or  species  within 
genera.  Familial  means  are  ranked  from  largest  to  smallest.  Within  the  Phyllosto- 
matidae,  subfamilial  means  are  similarly  ranked  as  are  genera  within  subfamilies. 
The  grand  mean  for  all  bats  is  ranked  with  the  familial  ranking.  These  tables  were 
generated  aside  from  the  primary  principal  component  and  discriminant  analyses, 
and  are  discussed  in  the  text  to  illuminate  the  interpretation  of  these  multivariate 
procedures. 


296 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  Al. — Ranked  means  and  statistics  for  the  alpha  angle. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Rhinopomatidae 

1 

40.25 

Rhinolophidae 

7 

39.10 

(42.55-37.09) 

0.663 

4.488 

Nycteridae 

1 

37.75 

Furipteridae 

1 

37.74 

Megadermatidae 

5 

37.52 

(38.63-37.10) 

0.281 

1.672 

Natalidae 

1 

36.74 

Pteropodidae 

30 

36.45 

(39.23-33.61) 

0.206 

3.094 

Craseonycteridae 

1 

35.83 

Thyropteridae 

1 

35.73 

Myzapodidae 

1 

35.37 

Phyllostomatidae 

49 

35.32 

(39.29-31.73) 

0.239 

4.740 

Phyllonycterinae 

3 

37.71 

(38.08-37.24) 

0.249 

1.145 

Phyllonycteris 

2 

38.08 

(38.49-37.68) 

0.403 

1.498 

Erophylla 

2 

37.80 

(38.16-37.45) 

0.352 

1.316 

Brachyphylla 

2 

37.24 

(37.37-37.11) 

0.131 

0.497 

Phyllostomatinae 

11 

36.70 

(39.29-34.69) 

0.470 

4.247 

Tonal  ia 

7 

39.29 

(40.74-36.98) 

0.517 

3.481 

Vampyrum 

1 

38.49 

Chrotopterus 

1 

38.44 

Micronycteris 

12 

37.31 

(39.21-35.40) 

0.395 

3.670 

Macrotus 

2 

36.83 

(37.74-35.93) 

0.904 

3.469 

Trachops 

1 

36.72 

Macrophylluin 

2 

36.37 

(36.68-36.07) 

0.308 

1.199 

Phylloderma 

1 

35.32 

Minion 

5 

35.15 

(37.91-33.04) 

1.066 

6.782 

Lonchorhina 

3 

35.12 

(36.29-34.37) 

0.592 

2.918 

Phyllostomus 

5 

34.69 

(35.85-33.31) 

0.435 

2.802 

Desmodontinae 

3 

36.27 

(39.05-33.86) 

1.509 

7.204 

Desniodus 

2 

39.05 

(39.37-38.72) 

0.329 

1.190 

Diphylla 

1 

35.91 

Diaemus 

1 

33.86 

Carolliinae 

2 

35.41 

(35.91-34.90) 

0.505 

2.018 

Caro  Ilia 

4 

35.91 

(36.31-35.66) 

0.143 

0.796 

Rhinophylla 

3 

34.90 

(35.95-33.95) 

0.578 

2.870 

Stenoderminae 

17 

35.04 

(36.58-34.07) 

0.165 

1.943 

Ainetrida 

2 

36.58 

(37.16-36.01) 

0.578 

2.234 

Phyllops 

2 

35.94 

(36.18-35.70) 

0.241 

0.950 

Ar  ileus 

1 

35.75 

Cenlurio 

1 

35.42 

Ardops 

1 

35.41 

Pvgoderma 

1 

35.37 

Artibeas 

13 

35.12 

(35.71-33.99) 

0.141 

1.444 

Sphaeronvcteris 

1 

35.08 

Vampyrodes 

1 

35.03 

Eclophylla 

2 

35.02 

(35.66-34.38) 

0.641 

2.589 

Enchisthenes 

1 

34.89 

Stnrnira 

10 

34.77 

(35.22-33.97) 

0.129 

1.177 

Slenoderma 

1 

34.42 

Uroderina 

2 

34.29 

(34.53-34.06) 

0.231 

0.954 

Vampyressa 

5 

34.29 

(35.39-33.81) 

0.279 

1.822 

Vampyrops 

10 

34.16 

(34.70-33.46) 

0.145 

1.346 

Chiroderma 

5 

34.07 

(34.66-33.49) 

0.234 

1.538 

Glossophaginae 

13 

33.73 

(35.41-31.73) 

0.290 

3.101 

Glossophaga 

4 

35.41 

(35.63-35.20) 

0.089 

0.504 

Platalina 

1 

35.34 

Leptonyderis 

3 

34.93 

(35.43-34.22) 

0.365 

1.811 

Lonchophvlla 

5 

34.47 

(35.18-33.74) 

0.246 

1.598 

Choeronycteris 

1 

33.62 

Lionycteris 

1 

33.48 

Choeroniscus 

5 

33.46 

(34.25-32.43) 

0.331 

2.209 

Lichonycteris 

3 

33.45 

(33.85-33.00) 

0.246 

1.273 

Hvlonycteris 

1 

33.39 

Musonycteris 

1 

33.22 

Scleronycteris 

1 

33.00 

Monophylltis 

2 

32.97 

(33.38-32.57) 

0.407 

1.747 

Anoura 

5 

31.73 

(32.57-30.79) 

0.320 

2.255 

Mormoopidae 

2 

35.17 

(37.13-33.21) 

1.963 

7.894 

All  bats 

153 

35.08 

(42.55-24.09) 

0.237 

8.373 

Vespertilionidae 

31 

35.02 

(38.84-31 .41) 

0.391 

6.220 

Mystacinidae 

1 

34.78 

Emballonuridae 

12 

32.95 

(35.66-29.07) 

0.507 

5.330 

Noctilionidae 

1 

30.38 

Molossidae 

9 

26.93 

(29.82-24.09) 

0.516 

5.754 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


297 


Table  A2. — Ranked  means  and  statistics  for  the  tip  index. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Phyllostomatidae 

49 

2.04 

(2.35-1.59) 

0.025 

8.546 

Carolliinae 

2 

2.24 

(2.30-2.18) 

0.061 

3.831 

Rhinophylla 

3 

2.30 

(2.50-2.20) 

0.102 

7.674 

Carol  lia 

4 

2.18 

(2.22-2.15) 

0.016 

1.457 

Stenoderminae 

17 

2.15 

(2.35-2.04) 

0.020 

3.902 

Pygoderma 

1 

2.35 

Anieirida 

2 

2.27 

(2.28-2.25) 

0.018 

1.1  14 

Vampyrops 

10 

2.22 

(2.34-2.07) 

0.024 

3.464 

Sphaeronycteris 

1 

2.22 

Ariteus 

1 

2.21 

Vampyressa 

5 

2.20 

(2.33-2.08) 

0.046 

4.699 

Chirodernta 

5 

2.19 

(2.24-2.10) 

0.026 

2.657 

Sturniru 

10 

2.17 

(2.29-2.10) 

0.020 

2.898 

Stenoderma 

1 

2.15 

Vampyrodes 

1 

2.14 

Ardops 

1 

2.11 

Ectophylla 

Uroaerma 

2 

2.10 

(2.14-2.07) 

0.036 

2.446 

2 

2.09 

(2.09-2.08) 

0.004 

0.286 

Phyllops 

2 

2.08 

(2.11-2.05) 

0.030 

2.015 

Centurio 

1 

2.05 

Enchisthenes 

1 

2.05 

Artibeus 

13 

2.04 

(2.13-1.91) 

0.020 

3.550 

Glossophaginae 

13 

2.06 

(2.20-1.81) 

0.033 

5.776 

Scleronycteris 

1 

2.20 

A  no  lira 

5 

2.20 

(2.29-2.12) 

0.035 

3.574 

Lichonycteris 

3 

2.19 

(2.28-2.09) 

0.055 

4.329 

Hylonycteris 

1 

2.19 

Choeroniscus 

5 

2.11 

(2.21-2.03) 

0.035 

3.740 

Lonchophylla 

5 

2.07 

(2.19-1.92) 

0.044 

4.799 

Choeronycteris 

1 

2.05 

Lionycteris 

1 

2.02 

Glossophagu 

4 

2.00 

(2.03-1.95) 

0.020 

2.021 

Monophyllus 

2 

1.99 

(2.00-1.99) 

0.004 

0.252 

Musonycteris 

1 

1.96 

Platalina 

1 

1.96 

Leptonycteris 

3 

1.81 

(1.85-1.76) 

0.024 

2.328 

Phyllostomatinae 

11 

1.92 

(2.11-1.68) 

0.033 

5.696 

Mucrophyllum 

2 

2.11 

(2.13-2.10) 

0.017 

1.158 

Phvlloderma 

1 

2.03 

Million 

5 

1.98 

(2.10-1.84) 

0.045 

5.076 

Trachops 

1 

1.96 

Lonchorhina 

3 

1.95 

(2.02-1.89) 

0.036 

3.190 

Vampyrum 

1 

1.94 

Chrotopterus 

1 

1.90 

(2.16-1.68) 

0.038 

7.029 

Micronycteris 

12 

1.89 

Tonal  ia 

7 

1.87 

(1.95-1.80) 

0.018 

2.495 

Phyllostomus 

5 

1.85 

(1.97-1.75) 

0.038 

4.548 

Macrotus 

2 

1.68 

(1.70-1.66) 

0.020 

1.643 

Desmodontinae 

3 

1.86 

(2.03-1.59) 

0.141 

13.083 

Diphylla 

1 

2.03 

Diaemus 

1 

1.98 

Desmodus 

2 

1.59 

(1.60-1.57) 

0.015 

1.348 

Phyllonycterinae 

3 

1.69 

(1.74-1.66) 

0.025 

2.613 

Brachyphylla 

2 

1.74 

(1.74-1.74) 

0.000 

0.025 

Erophylla 

2 

1.66 

(1.69-1.64) 

0.023 

1.955 

Phyllonycteris 

2 

1.66 

(1.68-1.65) 

0.016 

1.321 

Molossidae 

9 

1.98 

(2.30-1.81) 

0.044 

6.669 

Noctilionidae 

1 

1.95 

Craseonycteridae 

1 

1.86 

Natalidae 

1 

1.85 

All  bats 

153 

1.85 

(2.35-1.09) 

0.018 

12.280 

Vespertilionidae 

31 

1.81 

(2.17-1.60) 

0.027 

8.396 

Thyropteridae 

1 

1.81 

Myzapodidae 

1 

1.79 

6.596 

Pteropodidae 

30 

1.78 

(2.06-1.57) 

0.021 

Nycteridae 

1 

1.78 

0.017 

2.101 

Megadermatidae 

5 

1.76 

(1.83-1.74) 

Mystacinidae 

1 

1.75 

0.123 

10.221 

Mormoopidae 

2 

1.70 

(1.82-1.58) 

Emballonuridae 

12 

1.62 

(1.92-1.48) 

0.032 

6.950 

Furipteridae 

1 

1.58 

0.036 

6.764 

Rhinolophidae 

7 

1.40 

(1.52-1.29) 

Rhinopomatidae 

1 

1.09 

298 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A3. — Ranked  means  and  statistics  for  the  overall  aspect  ratio. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Molossidae 

9 

7.54 

(8.05-6.46) 

0.174 

6.935 

Emballonuridae 

12 

6.71 

(7.93-6.05) 

0.147 

7.620 

Noctilionidae 

1 

6.69 

Mormoopidae 

2 

6.04 

(6.39-5.68) 

0.356 

8.346 

All  bats 

153 

5.86 

(8.05-4.71) 

0.051 

10.866 

Phyllostomatidae 

49 

5.74 

(6.50-5.05) 

0.046 

5.615 

Glossophaginae 

13 

6.09 

(6.50-5.71) 

0.061 

3.623 

Anoura 

5 

6.50 

(6.74-6.23) 

0.087 

2.991 

Musonycteris 

1 

6.30 

Scleronycteris 

1 

6.23 

Lionycteris 

1 

6.19 

Monophyllus 

2 

6.18 

(6.29-6.07) 

0.109 

2.482 

Choeroniscus 

5 

6.17 

(6.40-6.01) 

0.073 

2.653 

Lichonycteris 

3 

6.13 

(6.30-5.98) 

0.091 

2.563 

Hvlonycteris 

1 

6.09 

Choeronycteris 

1 

6.09 

Lonchophylla 

5 

5.94 

(6.10-5.81) 

0.056 

2.102 

Leplonycteris 

3 

5.92 

(6.09-5.82) 

0.083 

2.425 

Platalina 

1 

5.72 

Glossophaga 

4 

5.71 

(5.80-5.64) 

0.033 

1.149 

Stenoderminae 

17 

5.71 

(5.96-5.26) 

0.050 

3.632 

Chiroderma 

5 

5.96 

(6.14-5.78) 

0.060 

2.240 

Uroderma 

2 

5.94 

(5.97-5.91) 

0.027 

0.648 

Vampyrops 

10 

5.93 

(6.11-5.78) 

0.030 

1.606 

Stenoderma 

1 

5.88 

Vampyressa 

5 

5.88 

(5.96-5.69) 

0.049 

1.869 

Centurio 

1 

5.88 

Ectophvllu 

2 

5.76 

(5.92-5.61) 

0.151 

3.713 

Sturnira 

10 

5.76 

(5.88-5.62) 

0.026 

1.404 

Artibeus 

13 

5.75 

(6.04-5.57) 

0.036 

2.231 

Vampyrodes 

1 

5.71 

Ardops 

1 

5.66 

Pvgoaenna 

1 

5.63 

Enchisthenes 

1 

5.62 

Phyllops 

2 

5.58 

(5.64-5.53) 

0.054 

1.365 

Ariteus 

1 

5.49 

Sphaeronycteris 

Ametrida 

1 

2 

5.32 

5.26 

(5.33-5.19) 

0.070 

1.874 

Carolliinae 

2 

5.69 

(5.74-5.64) 

0.049 

1.205 

Rhinophylla 

3 

5.74 

(5.95-5.52) 

0.124 

3.748 

Carollia 

4 

5.64 

(5.74-5.54) 

0.041 

1.442 

Phyllostomatinae 

11 

5.55 

(5.92-5.05) 

0.094 

5.620 

Minion 

5 

5.92 

(6.36-5.41) 

0.209 

7.892 

Phyllostomus 

5 

5.91 

(6.18-5.54) 

0.109 

4.107 

Lonchorhina 

3 

5.89 

(6.04-5.60) 

0.140 

4.135 

Phvlloderina 

1 

5.78 

Macrotus 

2 

5.77 

(6.06-5.49) 

0.284 

6.945 

Macrophyllum 

2 

5.45 

(5.48-5.43) 

0.029 

0.747 

Trachops 

1 

5.42 

Micronycteris 

12 

5.41 

(5.84-5.02) 

0.076 

4.867 

Chrotopierus 

1 

5.25 

Vampyrum 

1 

5.23 

Tonalia 

7 

5.05 

(5.57-4.74) 

0.113 

5.913 

Desmodontinae 

3 

5.50 

(5.81-5.17) 

0.186 

5.874 

D  iaem  us 

1 

5.81 

D  ip  h  vita 

1 

5.51 

Desnwdus 

2 

5.17 

(5.23-5.11) 

0.057 

1.572 

Phyllonycterinae 

3 

5.40 

(5.44-5.35) 

0.027 

0.872 

Brachvphvlla 

2 

5.44 

(5.46-5.42) 

0.016 

0.428 

Erophvlla 

2 

5.42 

(5.52-5.32) 

0.098 

2.559 

Phyllonvcteris 

2 

5.35 

(5.43-5.27) 

0.082 

2.169 

Vespertilionidae 

31 

5.73 

(6.90-4.92) 

0.085 

8.229 

Mystacinidae 

1 

5.71 

Thyropteridae 

1 

5.70 

Myzapodidae 

1 

5.65 

Craseonycteridae 

1 

5.64 

Natalidae 

1 

5.60 

Rhinopomatidae 

Megadermatidae 

1 

5.58 

5 

5.55 

(5.74-5.29) 

0.073 

2.938 

Furipteridae 

1 

5.52 

Pteropodidae 

30 

5.49 

(5.97-5.01) 

0.035 

3.533 

Nycteridae 

1 

5.48 

Rhinolophidae 

7 

5.42 

(5.99-4.71) 

0.157 

7.645 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


299 


Table  A4. — Ranked  means  and  statistics  for  the  aspect  ratio  of  the  wing  tip. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Molossidae 

9 

5.79 

(6.31-4.79) 

0.163 

8.465 

Embalonuridae 

12 

5.22 

(6.35-4.77) 

0.133 

8.790 

Noctilionidae 

1 

5.21 

Mormoopidae 

2 

4.73 

(5.12-4.34) 

0.390 

11.655 

Phyllostomatidae 

49 

4.67 

(5.41-3.86) 

0.048 

7.155 

Glossophaginae 

13 

4.98 

(5.41-4.60) 

0.066 

4.804 

Anoura 

5 

5.41 

(5.68-5.10) 

0.105 

4.328 

Scleronycteris 

1 

5.21 

Musonyeteris 

1 

5.13 

Lichanycteris 

3 

5.13 

(5.32-5.03) 

0.096 

3.252 

Choeroniscus 

5 

5.11 

(5.29-4.91) 

0.076 

3.320 

Lionycteris 

1 

5.06 

Hvlonvcteris 

1 

5.06 

Choeronycteris 

1 

4.97 

Monophvllus 

2 

4.95 

(5.04-4.85) 

0.097 

2.779 

Lonchophylla 

5 

4.89 

(5.11-4.67) 

0.088 

4.006 

Leptonvcteris 

3 

4.67 

(4.81-4.59) 

0.072 

2.673 

Glossophaga 

4 

4.62 

(4.67-4.58) 

0.025 

1.094 

Platalina 

1 

4.60 

Carolliinae 

2 

4.78 

(4.83-4.74) 

0.043 

1.279 

Rhinophylla 

3 

4.83 

(5.13-4.58) 

0.163 

5.865 

Carollia 

4 

4.74 

(4.85-4.62) 

0.052 

2.215 

Stenoderminae 

17 

4.70 

(4.96-4.22) 

0.053 

4.628 

Vampyrops 

10 

4.96 

(5.15-4.84) 

0.034 

2.146 

Chiroderma 

5 

4.96 

(5.16-4.71) 

0.073 

3.313 

Centurio 

1 

4.93 

Uroderma 

2 

4.88 

(4.89-4.87) 

0.009 

0.269 

Vampyressa 

5 

4.88 

(4.99-4.70) 

0.057 

2.615 

Stenoderma 

1 

4.87 

Pvyoderma 

1 

4.76 

Sturnira 

10 

4.74 

(4.88-4.66) 

0.020 

1.301 

Ectophylla 

2 

4.73 

(4.84-4.63) 

0.104 

3.107 

Vampyrodes 

1 

4.70 

Artibeus 

13 

4.67 

(4.87-4.48) 

0.037 

2.883 

Ardops 

1 

4.65 

Phyllops 

2 

4.57 

(4.65-4.50) 

0.071 

2.194 

Ariteus 

1 

4.52 

Enchisthenes 

1 

4.47 

Ametrida 

2 

4.33 

(4.37-4.28) 

0.045 

1.484 

Sphaeronvcteris 

1 

4.22 

Phyllostomatinae 

11 

4.48 

(4.86-4.04) 

0.079 

5.878 

M  ini  on 

5 

4.86 

(5.26-4.38) 

0.193 

8.892 

Lonchorhina 

3 

4.79 

(4.90-4.60) 

0.094 

3.413 

Phylloderma 

1 

4.74 

Phyllostomus 

5 

4.66 

(4.83-4.32) 

0.098 

4.710 

Macrotus 

2 

4.59 

(4.87-4.32) 

0.275 

8.475 

Macrophyllum 

2 

4.47 

(4.49-4.45) 

0.017 

0.535 

Trachops 

1 

4.33 

Micronycteris 

12 

4.32 

(4.73-3.90) 

0.076 

6.058 

Vampyrum 

1 

4.27 

Chrotopterus 

1 

4.25 

Tonatia 

7 

4.04 

(4.60-3.69) 

0.120 

7.839 

Desmodontinae 

3 

4.28 

(4.54-3.86) 

0.209 

8.457 

Diaemus 

1 

4.54 

D  ip  h  vlla 

1 

4.42 

Desmodus 

2 

3.86 

(3.88-3.84) 

0.019 

0.692 

Phyllonycterinae 

3 

4.16 

(4.21-4.10) 

0.032 

1.320 

Brachyphylla 

2 

4.21 

(4.22-4.20) 

0.008 

0.254 

Erophylla 

2 

4.17 

(4.24-4.10) 

0.069 

2.356 

Phyllonycteris 

2 

4.10 

(4.19-4.01) 

0.087 

2.999 

All  bats 

153 

4.58 

(6.35-3.44) 

0.044 

12.004 

Natalidae 

1 

4.51 

Thyropteridae 

1 

4.46 

Craseonycteridae 

1 

4.45 

Megadermatidae 

5 

4.45 

(4.72-4.19) 

0.089 

4.463 

Nycteridae 

1 

4.39 

Vespertilionidae 

31 

4.38 

(5.99-3.72) 

0.083 

10.512 

Myzapodidae 

1 

4.33 

Mystacinidae 

1 

4.27 

Pteropodidae 

30 

4.22 

(4.48-3.65) 

0.029 

3.782 

Furipteridae 

1 

4.19 

Rhinolophidae 

7 

3.93 

(4.27-3.44) 

0.099 

6.667 

Rhinopomatidae 

1 

3.69 

300 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A5. — Ranked  means  and  statistics  for  the  aspect  ratio  of  the  plagiopatagium. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Rhinopomatidae 

1 

2.17 

Molossidae 

9 

2.00 

(2.21-1.82) 

0.038 

5.762 

Emballonuridae 

12 

1.93 

(2.20-1.66) 

0.054 

9.630 

Rhinolophidae 

7 

1.78 

(2.06-1.51) 

0.077 

1 1 .440 

Noctilionidae 

1 

1.75 

Mormoopidae 

2 

1.68 

(1.68-1.68) 

0.002 

0.158 

Mystacinidae 

1 

1.65 

Furipteridae 

1 

1.64 

Vespertilionidae 

31 

1.59 

(1.99-1.32) 

0.028 

9.742 

All  bats 

153 

1.58 

(2.21-1.20) 

0.018 

14.364 

Myzapodidae 

1 

1.57 

Thyropteridae 

1 

1.54 

Pteropodidae 

30 

1.52 

(1.74-1.34) 

0.018 

6.619 

Craseonycteridae 

1 

1.49 

Megadermatidae 

5 

1.48 

(1.52-1.43) 

0.018 

2.688 

Nycteridae 

1 

1.46 

Natalidae 

1 

1.45 

Phyllostomatidae 

49 

1.40 

(1.59-1.19) 

0.015 

7.265 

Phyllonycterinae 

3 

1.53 

(1.55-1.51) 

0.011 

1.215 

Erophylla 

2 

1.55 

(1.59-1.51) 

0.040 

3.670 

Phyllonycteris 

2 

1.54 

(1.54-1.53) 

0.007 

0.673 

Brachyphylla 

2 

1.51 

(1.52-1.51) 

0.006 

0.583 

Desmodontinae 

3 

1.47 

(1.56-1.36) 

0.060 

7.004 

Desmudus 

2 

1.56 

(1.59-1.52) 

0.034 

3.082 

Diaem  us 

1 

1.51 

Diphylla 

1 

1.36 

Glossophaginae 

13 

1.46 

(1.58-1.38) 

0.019 

4.670 

Leptonycteris 

3 

1.58 

(1.62-1.54) 

0.023 

2.530 

Musonycteris 

1 

1.55 

Monophyllus 

2 

1.55 

(1.57-1.53) 

0.022 

2.013 

Lionycteris 

1 

1.50 

Ano  lira 

5 

1.47 

(1.51-1.44) 

0.010 

1.538 

Choeronycteris 

1 

1.46 

Platalina 

1 

1.44 

Choeroniscus 

5 

1.44 

(1.50-1.37) 

0.021 

3.253 

Lonchophylla 

5 

1.41 

(1.52-1.36) 

0.028 

4.507 

Glossophcigu 

4 

1.41 

(1.46-1.37) 

0.019 

2.700 

Scleronycteris 

1 

1.40 

Hvlonycteris 

1 

1.38 

Lichonycteris 

3 

1.38 

(1.45-1.31) 

0.040 

4.986 

Phyllostomatinae 

11 

1.40 

(1.59-1.29) 

0.031 

7.392 

Macrotus 

2 

1.59 

(1.66-1.52) 

0.071 

6.292 

Phyllostomus 

5 

1.57 

(1.67-1.44) 

0.048 

6.789 

Lonchorhina 

3 

1.46 

(1.53-1.35) 

0.055 

6.526 

Mimon 

5 

1.44 

(1.51-1.35) 

0.032 

5.007 

Micronycteris 

12 

1.39 

(1.51-1.24) 

0.025 

6.144 

Phylloderma 

1 

1.39 

Trachops 

1 

1.37 

Chrotopterus 

1 

1.33 

Tonatia 

7 

1.31 

(1.36-1.26) 

0.015 

2.985 

Vanipyrum 

1 

1.30 

Macrophyllum 

2 

1.29 

(1.29-1.28) 

0.003 

0.380 

Stenoderminae 

17 

1.33 

(1.41-1.19) 

0.015 

4.689 

Uroderma 

2 

1.41 

(1.42-1.40) 

0.009 

0.949 

Enchisthenes 

1 

1.40 

Artibeus 

13 

1.40 

(1.50-1.32) 

0.016 

4.023 

Centurio 

1 

1.37 

Ectophylla 

2 

1.36 

(1.41-1.30) 

0.056 

5.803 

Stenoderina 

1 

1.36 

Chiroderma 

5 

1.35 

(1.38-1.33) 

0.009 

1.414 

Vumpvressa 

5 

1.34 

(1.42-1.27) 

0.026 

4.300 

Vampyrodes 

1 

1.33 

Stnrnira 

10 

1.33 

(1.38-1.25) 

0.013 

2.999 

Ardops 

1 

1.33 

Phyllops 

2 

1.33 

(1.34-1.32) 

0.007 

0.762 

Vampyrops 

10 

1.33 

(1.40-1.26) 

0.012 

2.823 

Sphaeronvcteris 

1 

1.28 

Ariteus 

1 

1.26 

Pygoderma 

1 

1.20 

Ameirida 

2 

1.19 

(1.21-1.17) 

0.015 

1.818 

Carolliinae 

2 

1.26 

(1.27-1.25) 

0.009 

1.006 

Carollia 

4 

1.27 

(1.28-1.25) 

0.006 

0.870 

Rhinophylla 

3 

1.25 

(1.31-1.19) 

0.036 

4.936 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


301 


Table  A6. — Ranked  means  and  statistics  for  wing  loading  in  newtons  per  square  meter. 


Taxon 

N 

Mean 

Max-min 

±1  SE 

CV 

Molossidae 

9 

21.41 

(28.47-15.56) 

1.239 

17.358 

Pteropodidae 

30 

19.18 

(36.24-11.48) 

1.084 

30.959 

Noctilionidae 

1 

17.65 

Craseonycteridae 

1 

16.70 

All  bats 

153 

14.62 

(36.08-  3.69) 

0.507 

42.905 

Phyllostomatidae 

Desmodontinae 

49 

14.50 

(28.89-  3.92) 

0.686 

33.1  19 

3 

20.87 

(29.23-14.99) 

4.293 

35.639 

Diaemus 

1 

32.71 

Diphylla 

1 

20.34 

Desmodus 

2 

15.72 

(17.26-14.17) 

1.548 

13.931 

Phyllonycterinae 

3 

18.40 

(21.04-13.75) 

2.331 

21.940 

Phyllonycteris 

2 

21.04 

(24.56-17.52) 

3.522 

23.675 

Brcichvphvllu 

2 

20.41 

(22.74-18.08) 

2.329 

16.134 

Erophylla 

Stenoaerminae 

2 

13.75 

(15.46-12.05) 

1.708 

17.557 

17 

15.01 

(22.66-10.96) 

0.776 

21.319 

Enchisthenes 

1 

22.66 

Sturniru 

10 

17.85 

(28.58-10.68) 

2.071 

36.690 

Ariteus 

1 

17.42 

Vampyrodes 

1 

17.10 

Chiroaerma 

5 

17.09 

(21.86-13.15) 

1.424 

18.632 

Sphaeronycteris 

Centurio 

1 

16.96 

1 

16.76 

Vampyrops 

10 

16.09 

(21.13-10.38) 

0.895 

17.576 

Artibeus 

13 

15.94 

(23.23-10.56) 

1.084 

24.515 

Uroderma 

2 

14.04 

(16.59-1 1 .49) 

2.549 

25.675 

Stenodenna 

1 

13.39 

Ametridu 

2 

12.77 

(13.85-11.69) 

1.079 

11.954 

Ardops 

1 

11.98 

Vampyressa 

5 

11.50 

(13.01-  8.90) 

0.840 

16.327 

Ectophylla 

2 

11.47 

(12.73-10.22) 

1.256 

15.485 

Phyllops 

2 

11.16 

(13.29-  9.03) 

2.128 

26.968 

Pygodernia 

1 

10.96 

Phyilostomatinae 

11 

14.04 

(19.94-  7.88) 

1.205 

28.473 

Phyllostomus 

5 

19.94 

(24.11-16.47) 

1.242 

13.927 

Chrotopterus 

1 

18.81 

Trachons 

Phylloaernia 

1 

17.36 

1 

16.30 

Vampyruni 

1 

15.48 

Tonal  ia 

7 

14.64 

(19.35-10.86) 

1.206 

21.797 

Macrotus 

2 

12.96 

(14.68-11.25) 

1.715 

18.708 

Mimon 

5 

11.23 

(13.94-  6.81) 

1.295 

25.786 

Lonchorhina 

3 

11.05 

(13.22-  9.56) 

1.110 

17.392 

Micronycteris 

12 

8.77 

(15.43-  5.47) 

0.809 

31.949 

Mucrophyllum 

2 

7.88 

(10.24-  5.52) 

2.361 

42.356 

Glossophaginae 

13 

12.51 

(15.58-10.01) 

0.421 

12.128 

Musonycteris 

1 

15.58 

Leptonycteris 

3 

14.17 

(15.97-11.77) 

1.249 

15.270 

Choeronycteris 

1 

13.75 

Hylonycteris 

1 

13.50 

Lonchophyllu 

5 

13.21 

(17.03-11.46) 

1.005 

17.015 

Glossophaga 

4 

12.54 

(14.81-11.32) 

0.774 

12.343 

Anoura 

5 

12.31 

(17.35-  9.36) 

1.415 

25.706 

Choeroniscus 

5 

12.16 

(14.01-11.25) 

0.510 

9.390 

Menophyllus 

2 

11.93 

(12.45-11.41) 

0.519 

6.151 

Lichonycteris 

3 

11.63 

(12.03-11.27) 

0.220 

3.276 

Lionycteris 

1 

10.94 

Platalinu 

1 

10.94 

Scleronycteris 

1 

10.01 

Carolliinae 

2 

10.98 

(11.14-10.81) 

0.168 

2.168 

Rhinophylla 

3 

11.14 

(12.46-  9.70) 

0.800 

12.428 

Caro  Ilia 

4 

10.81 

(12.87-  9.10) 

0.857 

15.863 

Rhinopomatidae 

Megadermatidae 

1 

5 

11.82 

11.33 

(15.16-  8.32) 

1.428 

25.220 

Mystacinidae 

1 

11.15 

1.825 

24.406 

Mormoopidae 

2 

10.57 

(12.40-  8.75) 

Vespertilionidae 

31 

10.54 

(19.71-  6.92) 

0.544 

28.713 

Nycteridae 

1 

9.82 

1.364 

48.847 

Emballonuridae 

12 

9.67 

(21.16-  4.73) 

Rhinolophidae 

7 

8.05 

(14.48-  1.84) 

1.769 

58.147 

Myzapodidae 

1 

7.41 

Thyropteridae 

1 

5.91 

Natalidae 

1 

5.43 

Furipteridae 

1 

4.20 

302 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A7. — Ranked  means  and  statistics  for  the  relative  length  of  the  wing. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Furipteridae 

1 

2.62 

Natalidae 

1 

2.61 

Noctilionidae 

1 

2.53 

Emballonuridae 

12 

2.44 

(3.34-2.06) 

0.110 

15.553 

Myzapodidae 

1 

2.43 

Thyropteridae 

1 

2.42 

Craseonycteridae 

1 

2.35 

Nycteridae 

1 

2.31 

Mormoopidae 

2 

2.21 

(2.33-2.09) 

0.118 

7.531 

Rhinolophidae 

7 

2.20 

(2.51-2.03) 

0.064 

7.644 

Megadermatidae 

5 

2.18 

(2.43-1.95) 

0.084 

8.643 

Vespertilionidae 

31 

2.07 

(2.49-1.44) 

0.048 

12.907 

Phyllostomatidae 

49 

2.07 

(2.40-1.69) 

0.024 

8.174 

Carolliinae 

2 

2.22 

(2.24-2.20) 

0.018 

1.162 

Caro  Ilia 

4 

2.24 

(2.46-2.04) 

0.090 

8.029 

Rhinophylla 

3 

2.20 

(2.39-2.03) 

0.105 

8.251 

Phyllostomatinae 

11 

2.18 

(2.40-2.00) 

0.034 

5.103 

Minion 

5 

2.40 

(2.75-2.22) 

0.096 

8.969 

Lonchorhina 

3 

2.31 

(2.49-2.18) 

0.094 

7.087 

Macrophvlhwi 

2 

2.25 

(2.36-2.14) 

0.106 

6.656 

Micronycteris 

12 

2.22 

(2.56-1.82) 

0.058 

9.078 

Phylloderma 

1 

2.19 

Macrotus 

2 

2.18 

(2.29-2.07) 

0.111 

7.204 

Trachops 

1 

2.13 

Vampyrum 

1 

2.12 

Chrotopterus 

1 

2.11 

Phyllostomus 

5 

2.08 

(2.18-1.99) 

0.036 

3.845 

Tonal  ia 

7 

2.00 

(2.22-1.60) 

0.081 

10.764 

Stenoderminae 

17 

2.14 

(2.36-1.89) 

0.035 

6.840 

Ardops 

1 

2.36 

Stenodenna 

1 

2.34 

Phyllops 

2 

2.27 

(2.49-2.06) 

0.212 

13.188 

Centurio 

1 

2.24 

Vampyrodes 

1 

2.22 

Pygoderma 

1 

2.21 

Vampyrops 

10 

2.20 

(2.46-1.99) 

0.042 

5.999 

Artibeus 

13 

2.19 

(2.44-2.00) 

0.039 

6.343 

Ectophylla 

2 

2.19 

(2.34-2.03) 

0.150 

9.731 

Chiroderma 

5 

2.16 

(2.23-2.10) 

0.024 

2.528 

Vampyressa 

5 

2.15 

(2.41-2.03) 

0.074 

7.713 

Sturnira 

10 

2.05 

(2.22-1.83) 

0.043 

6.606 

Uroderina 

2 

2.03 

(2.06-2.00) 

0.029 

2.034 

Ariteus 

1 

1.96 

Ametrida 

2 

1.96 

(2.04-1.87) 

0.083 

5.968 

Enchisthenes 

1 

1.90 

Sphaeronycteris 

1 

1.89 

Glossophaginae 

13 

1.93 

(2.16-1.75) 

0.033 

6.112 

Scleronycteris 

1 

2.16 

Lionycteris 

1 

2.10 

Choeronycteris 

1 

2.01 

Anoura 

5 

1.98 

(2.26-1.75) 

0.102 

11.532 

Lichonycteris 

3 

1.98 

(2.03-1.93) 

0.029 

2.571 

Monophyllus 

2 

1.96 

(2.04-1.87) 

0.085 

6.174 

Choeroniscus 

5 

1.92 

(2.00-1.83) 

0.029 

3.312 

Platalina 

1 

1.91 

Glossophaga 

4 

1.87 

(1.91-1.84) 

0.016 

1.683 

Leptonycteris 

3 

1.84 

(1.97-1.75) 

0.068 

6.354 

Lonchophylla 

5 

1.84 

(1.90-1.79) 

0.021 

2.516 

Hylonycteris 

1 

1.79 

Musonycteris 

1 

1.75 

Desmodontinae 

3 

1.93 

(2.05-1.69) 

0.118 

10.617 

Desmodus 

2 

2.05 

(2.06-2.05) 

0.005 

0.314 

D  ip  h  vlla 

1 

2.05 

Diaemus 

1 

1.69 

Phyllonycterinae 

3 

1.92 

(2.05-1.78) 

0.077 

6.951 

Erophylla 

2 

2.05 

(2.14-1.96) 

0.093 

6.384 

Brachyphylla 

2 

1.94 

(2.00-1.87) 

0.068 

4.933 

Phy  lionycteris 

2 

1.78 

(1.89-1.68) 

0.106 

8.400 

All  bats 

153 

2.06 

(3.34-1.44) 

0.023 

13.586 

Mystacinidae 

1 

1.97 

Rhinopomatidae 

1 

1.97 

Molossidae 

9 

1.86 

(2.10-1.63) 

0.053 

8.476 

Pteropodidae 

30 

1.80 

(2.13-1.59) 

0.027 

8.108 

t 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


303 


Table  A8. — Ranked  means  and  statistics  for  the  relative  length  of  the  forearm. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Furipteridae 

1 

1.02 

Rhinopomatidae 

1 

0.94 

Emballonuridae 

12 

0.93 

(1.14-0.76) 

0.034 

12.771 

Rhinolophidae 

7 

0.92 

(1.01-0.86) 

0.022 

6.479 

Natalidae 

1 

0.92 

Myzapodidae 

1 

0.87 

Thyropteridae 

1 

0.86 

Noctilionidae 

1 

0.86 

Nycteridae 

1 

0.83 

Craseonyctridae 

1 

0.82 

Mormoopidae 

2 

0.82 

(0.82-0.81) 

0.007 

1.130 

Megadermatidae 

5 

0.79 

(0.88-0.71) 

0.028 

7.967 

Vespertilionidae 

31 

0.74 

(0.92-0.56) 

0.014 

10.850 

All  bats 

153 

0.73 

(1.14-0.52) 

0.010 

16.238 

Mystacinidae 

1 

0.72 

Phyllostomatidae 

49 

0.68 

(0.81-0.56) 

0.009 

9.145 

Phyllostomatinae 

11 

0.75 

(0.81-0.69) 

0.012 

5.232 

Macrotus 

2 

0.81 

(0.85-0.78) 

0.035 

6.162 

Mimon 

5 

0.80 

(0.91-0.71) 

0.034 

9.502 

Lonchorhinu 

3 

0.78 

(0.85-0.74) 

0.032 

7.171 

Micronycteris 

12 

0.77 

(0.90-0.64) 

0.021 

9.585 

Phyllostomus 

5 

0.73 

(0.77-0.70) 

0.013 

3.857 

Chrotopterus 

1 

0.73 

Macrophyllum 

2 

0.72 

(0.75-0.69) 

0.030 

5.940 

Phylloaerma 

1 

0.72 

Vampyrum 

Trachops 

1 

1 

0.72 

0.72 

Tonatia 

7 

0.69 

(0.77-0.57) 

0.026 

9.746 

Phyllonycterinae 

3 

0.72 

(0.77-0.67) 

0.029 

6.996 

Erophylla 

2 

0.77 

(0.81-0.73) 

0.042 

7.640 

Brachyphytla 

2 

0.71 

(0.73-0.68) 

0.025 

4.913 

Phyllonycteris 

2 

0.67 

(0.71-0.63) 

0.044 

9.255 

Caroiliinae 

2 

0.69 

(0.70-0.67) 

0.018 

3.807 

Carollia 

4 

0.70 

(0.77-0.65) 

0.026 

7.338 

Rhinophylla 

3 

0.67 

(0.69-0.63) 

0.018 

4.555 

Desmodontinae 

3 

0.68 

(0.79-0.57) 

0.065 

16.603 

Desmodus 

2 

0.79 

(0.80-0.79) 

0.003 

0.523 

Diphylla 

1 

0.67 

Diaemus 

1 

0.57 

Stenoderminae 

17 

0.68 

(0.76-0.59) 

0.013 

7.756 

Ardops 

1 

0.76 

Sienoderma 

1 

0.74 

Phyllops 

2 

0.74 

(0.80-0.68) 

0.062 

11.844 

Centurio 

1 

0.73 

Artibeus 

13 

0.72 

(0.80-0.66) 

0.011 

5.679 

Vampyrodes 

1 

0.71 

Ectophylla 

2 

0.70 

(0.76-0.65) 

0.057 

11.414 

Vumpyrops 

10 

0.68 

(0.76-0.61) 

0.013 

5.890 

Chiroderma 

5 

0.68 

(0.69-0.66) 

0.006 

2.041 

Vumpyressa 

5 

0.67 

(0.74-0.64) 

0.017 

5.645 

Pygoderma 

1 

0.66 

Uruderma 

2 

0.66 

(0.67-0.65) 

0.010 

2.229 

Sturnira 

10 

0.65 

(0.70-0.56) 

0.014 

6.773 

Enchisihenes 

1 

0.62 

Ariteus 

1 

0.61 

Ainetrida 

2 

0.60 

(0.63-0.57) 

0.028 

6.712 

Sphaeronycteris 

1 

0.59 

Glossophaginae 

13 

0.63 

(0.70-0.56) 

0.010 

5.819 

Lionycteris 

1 

0.70 

Scleronycteris 

1 

0.68 

Choerortycteris 

1 

0.66 

Leptonycteris 

3 

0.66 

(0.71-0.62) 

0.030 

7.848 

Monophyllus 

2 

0.65 

(0.68-0.63) 

0.028 

6.059 

Platalinci 

1 

0.65 

Glossophaga 

4 

0.62 

(0.65-0.61) 

0.008 

2.675 

Lichortycteris 

3 

0.62 

(0.64-0.59) 

0.016 

4.520 

Choeroniscus 

5 

0.62 

(0.65-0.59) 

0.010 

3.751 

Anuura 

5 

0.62 

(0.69-0.55) 

0.027 

9.645 

Lonchophylla 

5 

0.60 

(0.63-0.56) 

0.013 

4.816 

Musonycteris 

1 

0.59 

Hvlonycteris 

1 

0.56 

Pteropodidae 

30 

0.65 

(0.80-0.52) 

0.012 

9.730 

Molossidae 

9 

0.63 

(0.72-0.56) 

0.018 

8.610 

304 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A9. — Ranked  means  and  statistics  for  the  relative  length  of  digit  III. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Natalidae 

1 

1.69 

Noctilionidae 

1 

1.67 

Furipteridae 

1 

1.61 

Myzapodidae 

1 

1.56 

Thyropteridae 

1 

1.56 

Craseonycteridae 

1 

1.53 

Emballonuridae 

12 

1.51 

(2.19-1.28) 

0.077 

17.721 

Nycteridae 

1 

1.48 

Mormoopidae 

2 

1.39 

(1.50-1.28) 

0.111 

1 1.291 

Megadermatidae 

5 

1.39 

(1.55-1.24) 

0.057 

9.091 

Phyllostomatidae 

49 

1.39 

(1.60-1.11) 

0.018 

9.206 

Carolliinae 

2 

1.54 

(1.54-1.54) 

0.000 

0.018 

Rhinophvlla 

3 

1.54 

(1.71-1.39) 

0.092 

10.325 

Caro  Ilia 

4 

1.54 

(1.69-1.40) 

0.064 

8.370 

Stenoderminae 

17 

1.46 

(1.60-1.28) 

0.024 

6.728 

Ardops 

1 

1.60 

Stenoderma 

1 

1.59 

Pvgodenna 

1 

1.55 

Phyllops 

2 

1.54 

(1.69-1.39) 

0.150 

13.834 

Vampyrops 

10 

1.51 

(1.70-1.36) 

0.030 

6.345 

Vainpyrodes 

1 

1.51 

Cenlurio 

1 

1.51 

Chirodenna 

5 

1.48 

(1.54-1.42) 

0.021 

3.098 

Ectophvlla 

2 

1.48 

(1.57-1.39) 

0.094 

8.931 

Vumpyressa 

5 

1.48 

(1.68-1.38) 

0.059 

8.884 

Anioeus 

13 

1.47 

(1.66-1.31) 

0.028 

6.936 

Sturnira 

10 

1.40 

(1.54-1.27) 

0.030 

6.720 

Uroderma 

2 

1.37 

(1.39-1.35) 

0.019 

1.941 

Ametrida 

2 

1.36 

(1.41-1.30) 

0.054 

5.639 

Arireus 

1 

1.35 

Sphaeronvcteris 

1 

1.30 

Enchisthenes 

1 

1.28 

Phyllostomatinae 

11 

1.43 

(1.59-1.30) 

0.026 

6.095 

Mimon 

5 

1.59 

(1.84-1.48) 

0.065 

9.137 

Macrophvlluni 

2 

1.53 

(1.60-1.45) 

0.076 

6.995 

Lonchorhina 

3 

1.52 

(1.65-1.43) 

0.064 

7.265 

Phvlloderma 

1 

1.47 

Micronycteris 

12 

1.45 

(1.66-1.18) 

0.040 

9.658 

Trachops 

1 

1.41 

Vampvrum 

1 

1.40 

Chrotopterus 

1 

1.38 

Macrotus 

2 

1.37 

(1.44-1.29) 

0.076 

7.823 

Phyllostomus 

5 

1.35 

(1.41-1.27) 

0.028 

4.658 

Tonal  ia 

7 

1.30 

(1.46-1.03) 

0.056 

11.356 

Glossophaginae 

13 

1.30 

(1.49-1.16) 

0.026 

7.090 

Scleronycteris 

1 

1.49 

Lionycteris 

1 

1.41 

Anoura 

5 

1.36 

(1.57-1.20) 

0.076 

12.464 

Lichonycteris 

3 

1.36 

(1.40-1.34) 

0.020 

2.487 

Choeronycteris 

1 

1.35 

Choeroniscus 

5 

1.30 

(1.38-1.24) 

0.022 

3.759 

Monophyllus 

2 

1.30 

(1.36-1.25) 

0.057 

6.231 

Platalina 

1 

1.27 

Glossophaga 

4 

1.25 

(1.27-1.23) 

0.009 

1.423 

Lonchophylla 

5 

1.24 

(1.28-1.21) 

0.013 

2.432 

Hvlonycteris 

1 

1.23 

Leptonycteris 

3 

1.19 

(1.26-1.14) 

0.038 

5.529 

Musonycteris 

1 

1.16 

Desmodontinae 

3 

1.25 

(1.37-1.13) 

0.071 

9.809 

Diphylla 

1 

1.37 

Desmodus 

2 

1.26 

(1.27-1.25) 

0.007 

0.841 

Diaemus 

1 

1.13 

Phyllonycterinae 

3 

1.21 

(1.28-1.11) 

0.050 

7.107 

Erophylla 

2 

1.28 

(1.33-1.23 

0.051 

5.629 

Brachyphvlla 

2 

1.23 

(1.27-1.19) 

0.043 

4.945 

Phyllonycteris 

2 

1.11 

(1.17-1.05) 

0.062 

7.885 

Vespertilionidae 

31 

1.34 

(1.69-0.89) 

0.035 

14.739 

All  bats 

153 

1.33 

(2.19-0.89) 

0.015 

14.209 

Rhinolophidae 

7 

1.28 

(1.52-1.15) 

0.046 

9.478 

Mystacinidae 

1 

1.26 

Molossidae 

9 

1.24 

(1.41-1.07) 

0.038 

9.294 

Pteropodidae 

30 

1.15 

(1.33-0.99) 

0.017 

8.043 

Rhinopomatidae 

1 

1.03 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


305 


Table  A10. — Ranked  means  and  statistics  for  the  relative  length  of  digit  IV. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Noctilionidae 

1 

1.27 

Natalidae 

1 

1.25 

Furipteridae 

1 

1.25 

Myzapodidae 

1 

1.25 

Thyropteridae 

1 

1.20 

Craseonycteridae 

1 

1.18 

Nycteridae 

1 

1.10 

Emballonuridae 

12 

1.07 

(1.43-0.85) 

0.049 

15.900 

Vespertilionidae 

31 

1.07 

(1.27-0.74) 

0.022 

11.381 

Rhinolophidae 

7 

1.04 

(1.20-0.89) 

0.038 

9.718 

Mystacinidae 

1 

1.03 

Mormoopidae 

2 

1.03 

(1.07-0.99) 

0.044 

5.998 

Phyllostomatidae 

49 

1.03 

(1.19-0.82) 

0.012 

8.208 

Carolliinae 

2 

1.11 

(1.11-1.10) 

0.004 

0.516 

Rhinophvlla 

3 

1.11 

(1.19-1.04) 

0.045 

6.930 

Carollia 

4 

1.10 

(1.19-1.01) 

0.038 

6.933 

Stenoderminae 

17 

1.08 

(1.19-1.00) 

0.013 

4.954 

Ardops 

1 

1.19 

Stenoderma 

1 

1.16 

Phvl  lops 

2 

1.14 

(1.24-1.04) 

0.101 

12.438 

Pvgoderma 

1 

1.13 

Vampyrodes 

1 

1.12 

Artibeus 

13 

1.10 

(1.22-1.00) 

0.016 

5.277 

Ectophvlla 

2 

1.09 

(1.15-1.03) 

0.062 

8.082 

Vampyrops 

10 

1.09 

(1.19-0.98) 

0.019 

5.659 

Vampvressa 

5 

1.08 

(1.21-1.02) 

0.037 

7.644 

Sphaeronvcteris 

1 

1.07 

Chiroderma 

5 

1.07 

(1.09-1.05) 

0.008 

1.605 

Centurio 

1 

1.06 

Ametrida 

2 

1.05 

(1.09-1.01) 

0.039 

5.234 

Sturnira 

10 

1.04 

(1.16-0.93) 

0.023 

6.938 

Ariteus 

1 

1.02 

Uroderma 

2 

1.00 

(1.01-0.99) 

0.010 

1.403 

Enchisthenes 

1 

1.00 

Phyllostomatinae 

1 1 

1.07 

(1.15-1.00) 

0.015 

4.726 

Macrophtlhnn 

2 

1.15 

(1.21-1.09) 

0.061 

7.502 

Minton 

5 

1.14 

(1.30-1.05) 

0.046 

9.065 

Lonchorhina 

3 

1.11 

(1.20-1.02) 

0.050 

7.846 

Micronycteris 

12 

1.11 

(1.23-0.92) 

0.026 

8.110 

Trachops 

Phvlloaerma 

1 

1.09 

1 

1.07 

Vampyrunt 

1 

1.05 

Chrotopterus 

1 

1.05 

0.035 

9.170 

Tonatia 

7 

1.02 

(1.16-0.86) 

Phyllostomus 

5 

1.02 

( 1 .10-0.97) 

0.028 

6.068 

Macrotus 

2 

1.00 

(1.09-0.90) 

0.094 

13.335 

Desmodontinae 

3 

0.99 

(1.06-0.89) 

0.052 

9.075 

Diphylla 

1 

1.06 

Desntodus 

2 

1.04 

(1.04-1.03) 

0.004 

0.520 

Diaemus 

1 

0.89 

Phyllonycterinae 

3 

0.95 

(1.00-0.88) 

0.036 

6.533 

Erophylla 

2 

1.00 

(1.03-0.97) 

0.031 

4.418 

Brachyphylla 

2 

0.97 

(1.00-0.93) 

0.035 

5.093 

Phvllonycteris 

2 

0.88 

(0.94-0.82) 

0.060 

9.631 

Glossophaginae 

13 

0.94 

(1.05-0.82) 

0.016 

6.247 

Scleronveteris 

1 

1.05 

Lionycteris 

1 

1.01 

Choeronvcreris 

1 

0.98 

Monophvlhts 

2 

0.97 

(1.00-0.93) 

0.034 

5.003 

Lichonycteris 

3 

0.96 

(0.97-0.96) 

0.003 

0.515 

A  no  lira 

5 

0.96 

(1.08-0.83) 

0.047 

10.987 

Plcilalina 

1 

0.95 

Glossophaga 

4 

0.93 

(0.94-0.92) 

0.007 

1.420 

Choeroniscus 

5 

0.93 

(0.98-0.88) 

0.017 

4.156 

Lonchophvlla 

5 

0.90 

(0.93-0.86) 

0.014 

3.618 

Leptonycteris 

3 

0.89 

(0.94-0.86) 

0.028 

5.496 

Hvlonycieris 

1 

0.88 

Mtisonvcteris 

1 

0.82 

Megadermatidae 

5 

1.03 

(1.09-0.95) 

0.029 

6.294 

All  bats 

153 

1.02 

(1.43-0.74) 

0.010 

11.544 

Molossidae 

9 

0.95 

(1.10-0.85) 

0.026 

8.256 

Pteropodidae 

30 

0.92 

(1.04U80) 

0.013 

7.458 

Rhinopomatidae 

1 

0.86 

306 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A 1 1. — Ranked  means  and  statistics  for  the  relative  length  of  digit  V. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Natalidae 

1 

1.26 

Furipteridae 

1 

1.24 

Nycteridae 

1 

1.14 

Thyropteridae 

1 

1.12 

Myzapodidae 

1 

1.11 

Craseonycteridae 

1 

1.10 

Megadermatidae 

5 

1.07 

(1.18-0.99) 

0.038 

7.920 

Rhinolophidae 

7 

1.04 

(1.21-0.86) 

0.046 

11.774 

Phyllostomatidae 

49 

0.98 

(1.14-0.76) 

0.015 

10.554 

Carolliinae 

2 

1.09 

(1.11-1.07) 

0.021 

2.713 

Caro  Ilia 

4 

1.11 

(1.21-1.01) 

0.043 

7.690 

Rhinoph  ylla 

3 

1.07 

(1.15-1.01) 

0.041 

6.683 

Phyllostomatinae 

11 

1.07 

(1.12-0.93) 

0.017 

5.211 

Maerophyllum 

2 

1.12 

(1.17-1.08) 

0.044 

5.498 

Minion 

5 

1.12 

(1.24-1.00) 

0.052 

10.289 

Vampyrum 

1 

1.11 

Micronycteris 

12 

1.10 

(1.19-0.96) 

0.021 

6.469 

Chrotopterus 

1 

1.10 

Lonchorhinu 

3 

1.07 

(1.13-0.99) 

0.042 

6.774 

Tonatia 

7 

1.06 

(1.21-0.89) 

0.040 

10.035 

Trachons 

PhyUoaermu 

1 

1.05 

1 

1.04 

Macrotus 

2 

1.03 

(1.12-0.94) 

0.090 

12.441 

Phyllostonius 

5 

0.93 

(1.01-0.88) 

0.025 

5.971 

Stenoderminae 

17 

1.02 

(1.14-0.89) 

0.017 

6.888 

Ardops 

1 

1.14 

Phyllops 

2 

1.11 

(1.21-1.01) 

0.099 

12.597 

Pvgoderma 

1 

1.10 

Stenodermu 

1 

1.09 

Cenlurio 

1 

1.07 

Vamp  yrodes 

1 

1.06 

Ectophylla 

Art  in e  us 

2 

1.04 

(1.08-0.99) 

0.042 

5.669 

13 

1.04 

(1.17-0.93) 

0.019 

6.750 

Vampyrops 

10 

1.03 

(1.13-0.94) 

0.016 

4.962 

Ametrida 

2 

1.01 

(1.07-0.95) 

0.061 

8.608 

Vumpyressu 

5 

1.01 

(1.12-0.93) 

0.035 

7.779 

Chiroderma 

5 

1.00 

(1.02-0.98) 

0.007 

1.554 

Sturnira 

10 

0.98 

(1.08-0.85) 

0.024 

7.917 

Ariteus 

1 

0.97 

Uroderma 

2 

0.94 

(0.96-0.92) 

0.021 

3.186 

Sphaeronycieris 

Enchisthenes 

1 

0.91 

1 

0.89 

Phyllonycterinae 

3 

0.93 

(0.99-0.87) 

0.034 

6.389 

Erophylla 

2 

0.99 

(1.02-0.96) 

0.027 

3.915 

Brachyphylla 

2 

0.93 

(0.96-0.91) 

0.028 

4.249 

Phyllonycteris 

2 

0.87 

(0.93-0.81) 

0.061 

9.876 

Desmodontinae 

3 

0.92 

(1.02-0.76) 

0.085 

15.868 

Desmodus 

2 

1.02 

(1.04-1.00) 

0.018 

2.472 

Diphvlla 

1 

0.99 

Diaemus 

1 

0.76 

Glossophaginae 

13 

0.87 

(0.97-0.76) 

0.015 

6.188 

Scleronvcteris 

1 

0.97 

Lionycteris 

1 

0.93 

Lichonycteris 

3 

0.90 

(0.91-0.89) 

0.006 

1.213 

Plalalina 

1 

0.90 

Choeronycteris 

1 

0.90 

Gtossophaga 

4 

0.89 

(0.90-0.87) 

0.007 

1.554 

Choeroniscus 

5 

0.86 

(0.91-0.79) 

0.021 

5.493 

Lonchophylla 

5 

0.85 

(0.90-0.82) 

0.016 

4.147 

Monophyllus 

2 

0.84 

(0.87-0.82) 

0.024 

4.045 

Anoura 

5 

0.84 

(0.93-0.75) 

0.037 

9.846 

Leptonycteris 

3 

0.83 

(0.90-0.79) 

0.034 

7.116 

Hvlonycteris 

1 

0.81 

Musonycteris 

1 

0.76 

Noctilionidae 

1 

0.98 

Emballonuridae 

12 

0.98 

(1.38-0.72) 

0.052 

18.283 

Mormoopidae 

2 

0.98 

(0.98-0.97) 

0.009 

1.250 

All  bats 

153 

0.94 

(1.38-0.57) 

0.012 

16.083 

Vespertilionidae 

31 

0.94 

(1.22-0.57) 

0.025 

15.026 

Mystacinidae 

1 

0.87 

Rhinopomatidae 

1 

0.87 

Pteropodidae 

30 

0.85 

(1.01-0.72) 

0.014 

8.678 

Molossidae 

9 

0.63 

(0.72-0.57) 

0.015 

7.133 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


307 


Table  A12. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  III 

by  the  metacarpal. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

CV 

Rhinopomatidae 

1 

61.70 

Emballonuridae 

12 

57.67 

(63.21-53.06) 

0.920 

5.524 

Furipteridae 

1 

54.80 

Thyropteridae 

1 

53.85 

Mystacinidae 

1 

53.37 

Mormoopidae 

2 

52.87 

(56.37-49.36) 

3.505 

9.375 

Vespertilionidae 

31 

51.74 

(63.40-41.82) 

0.708 

7.621 

Rhinolophidae 

7 

51.57 

(56.03-46.57) 

1.246 

6.391 

Molossidae 

9 

50.70 

(54.42-43.21) 

1.015 

6.004 

Natalidae 

1 

49.92 

All  bats 

153 

47.04 

(63.40-35.39) 

0.533 

14.017 

Phyllostomatidae 

Desmodontinae 

49 

3 

45.12 

50.53 

(54.67-36.15) 

(54.67-48.44) 

0.456 

2.068 

7.081 

7.088 

Desmodus 

2 

54.67 

(55.89-53.44) 

1.227 

3.175 

Diaemus 

1 

48.48 

Diphvlla 

1 

48.44 

Phyllonycterinae 

3 

48.71 

(49.70-47.69) 

0.581 

2.065 

Phyllonycteris 

2 

49.70 

(50.43-48.98) 

0.727 

2.069 

Brachyphvlta 

2 

48.73 

(49.53-47.92) 

0.805 

2.336 

Erophylla 

2 

47.69 

(47.85-47.54) 

0.156 

0.462 

Glossophaginae 

13 

46.99 

(49.43-44.66) 

0.413 

3.167 

Leptonycteris 

3 

49.43 

(50.21-48.95) 

0.393 

1.377 

Lionycteris 

1 

48.71 

Plaialina 

1 

48.37 

Glossophaga 

4 

48.02 

(48.33-47.18) 

0.283 

1.177 

Lonchophylla 

5 

47.72 

(49.55-46.75) 

0.534 

2.503 

Monophyllus 

2 

47.13 

(47.47-46.79) 

0.342 

1.026 

Musonycteris 

1 

47.12 

Hvlonycteris 

1 

46.98 

Choeroniscus 

5 

46.80 

(47.75-45.32) 

0.435 

2.077 

Choeronycteris 

1 

45.84 

Lichonycteris 

3 

45.13 

(45.65-44.22) 

0.460 

1.765 

Scleronycteris 

1 

45.02 

Anoura 

5 

44.66 

(46.46-43.17) 

0.542 

2.714 

Phyliostomatinae 

11 

43.61 

(48.03-36.15) 

1.178 

8.960 

Phyllostomus 

5 

48.03 

(49.21-46.62) 

0.500 

2.330 

Lonchorhinu 

3 

47.34 

(48.55-45.80) 

0.810 

2.963 

Macrophylluni 

2 

46.66 

(46.82-46.50) 

0.158 

0.480 

Phylloderma 

1 

46.23 

Micronvcteris 

12 

46.17 

(48.80-43.72) 

0.588 

4.415 

Macrotus 

2 

44.35 

(44.39-44.32) 

0.035 

0.111 

Minion 

5 

43.70 

(45.18-42.78) 

0.473 

2.419 

Tonal  ia 

7 

42.21 

(43.08-41.27) 

0.276 

1.729 

Trachops 

1 

40.82 

Chrotopterus 

1 

38.08 

Vampyrum 

1 

36. 1 5 

Stenoderminae 

17 

43.41 

(45.49-40.92) 

0.273 

2.594 

Uroderma 

2 

45.49 

(45.52-45.46) 

0.028 

0.088 

Artibeus 

13 

44.76 

(46.66-43.21) 

0.279 

2.251 

Enchisthenes 

1 

44.75 

Ariteus 

1 

44.34 

Vampyrodes 

1 

43.97 

Ectophylla 

2 

43.68 

(43.69-43.68) 

0.005 

0.015 

Sphaeronycteris 

Ardops 

1 

1 

43.67 

43.58 

Chiroderma 

5 

43.57 

(45.49-42.29) 

0.676 

3.467 

Stenodenna 

1 

43.42 

Sturnira 

10 

43.11 

(44.51-41.67) 

0.323 

2.370 

Vampyressa 

5 

43.11 

(44.34-40.57) 

0.659 

3.416 

Phyllops 

2 

42.97 

(44.08-41.86) 

1.115 

3.669 

Centurio 

1 

42.37 

Vampyrops 

10 

42.25 

(44.06-40.46) 

0.426 

3.187 

Amelrida 

2 

42.00 

(42.35-41.66) 

0.345 

1.161 

Pygoderma 

1 

40.92 

1.634 

Carolliinae 

2 

42.34 

(42.83-41.85) 

0.489 

Carollia 

4 

42.83 

(43.39-42.37) 

0.241 

1.126 

Rhinophyllu 

3 

41.85 

(42.81-40.06) 

0.896 

3.708 

Myzapodiaae 

1 

44.82 

Noctilionidae 

1 

44.77 

Craseonycteridae 

1 

43.44 

Nycteridae 

1 

41.44 

Megadermatidae 

5 

39.43 

(40.53-37.48) 

0.566 

3.210 

Pteropodidae 

30 

38.71 

(42.49-35.39) 

0.311 

4.396 

308 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A13. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  III 

by  the  first  phalanx. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Nycteridae 

1 

28.50 

Pteropodidae 

30 

27.16 

(29.52-25.32) 

0.186 

3.743 

Thyropteridae 

Megaaermatidae 

1 

5 

23.32 

22.65 

(25.68-21.22) 

0.788 

7.776 

Molossidae 

9 

22.30 

(26.36-19.15) 

0.701 

9.425 

Natalidae 

1 

21.38 

Rhinolophidae 

7 

20.39 

(23.54-12.46) 

1.390 

18.039 

Myzapodidae 

1 

20.26 

All  bats 

153 

20.14 

(29.52-  9.96) 

0.379 

23.292 

Vespertilionidae 

31 

19.94 

(24.70-11.77) 

0.373 

10.417 

Emballonuridae 

12 

17.33 

(21.77-10.76) 

1.086 

21.711 

Phyllostomatidae 

49 

16.89 

(21.00-10.21) 

0.329 

13.638 

Carolliinae 

2 

18.93 

(19.00-18. 87) 

0.065 

0.488 

Caroltia 

4 

19.00 

(19.23-18.55) 

0. 1 54 

1.619 

Rhinophylla 

3 

18.87 

(19.34-18.37) 

0.279 

2.560 

Phyllonycterinae 

3 

17.77 

(20.14-15.60) 

1.312 

12.791 

Erophvlla 

2 

20.14 

(20.45-19.83) 

0.309 

2.170 

Phyllonvcteris 

2 

17.57 

(18.13-17.01) 

0.562 

4.525 

Bracliyphvlla 

2 

15.60 

(16.06-15.15) 

0.453 

4.108 

Phyllostomatinae 

11 

17.73 

(21.00-13.03) 

0.667 

12.472 

Vampyruni 

1 

21.00 

Chrotopterus 

1 

19.99 

Macrotus 

2 

18.84 

(19.15-18.54) 

0.305 

2.292 

Micronycteris 

12 

18.80 

(21.17-16.87) 

0.437 

8.047 

Tonal  in 

7 

18.77 

(20.16-17.72) 

0.320 

4.511 

Macrophvllum 

2 

18.46 

(18.90-18.02) 

0.439 

3.366 

Trachops 

1 

17.55 

Million 

5 

16.29 

(19.37-14.51) 

1.053 

14.445 

Lonchorhina 

3 

16.28 

(17.61-15.50) 

0.668 

7.105 

Phvlloderma 

1 

16.04 

Phyllostomus 

5 

13.03 

(14.25-12.07) 

0.426 

7.304 

Glossophaginae 

13 

17.08 

(18.76-14.98) 

0.365 

7.706 

Platalina 

1 

18.76 

Choeronycieris 

1 

18.53 

Scleronycteris 

1 

18.11 

Lichonycteris 

3 

18.09 

(18.61-17.48) 

0.329 

3.150 

Hvlonycteris 

1 

17.95 

Glossophaga 

4 

17.67 

(18.09-16.89) 

0.267 

3.021 

Musonycteris 

1 

17.47 

Choeroniscus 

5 

17.15 

(17.81-16.41) 

0.225 

2.937 

Lonchophyllu 

5 

16.90 

(19.41-16.05) 

0.632 

8.354 

Monophyllus 

2 

15.99 

(16.30-15.68) 

0.308 

2.721 

Anoura 

5 

15.23 

(16.09-14.65) 

0.265 

3.886 

Lionvcteris 

1 

15.22 

Leptonvcteris 

3 

14.98 

(15.48-14.40) 

0.312 

3.609 

Stenoderminae 

17 

16.94 

(19.58-14.97) 

0.350 

8.513 

Centurio 

1 

19.58 

Pvgoderma 

1 

19.44 

Phyllops 

2 

19.13 

(19.48-18.77) 

0.356 

2.632 

Vampyressa 

5 

17.75 

(19.57-16.58) 

0.664 

8.365 

Chiroderina 

5 

17.25 

(18.45-15.81) 

0.488 

6.329 

Sphaeronvcteris 

1 

17.25 

Ectophvlla 

2 

17.20 

(19.08-15.33) 

1.876 

15.423 

Sturnira 

10 

17.19 

(18.55-16.55) 

0.182 

3.344 

Enchisthenes 

1 

17.01 

Vampyrops 

Uroaerma 

10 

16.78 

(17.69-15.31) 

0.237 

4.465 

2 

16.52 

(16.72-16.32) 

0.200 

1.716 

Vampvrodes 

1 

16.37 

Arlibeus 

13 

15.80 

(17.66-14.11) 

0.316 

7.220 

Ainetrida 

2 

15.44 

(15.57-15.31) 

0.131 

1.204 

Slenoderina 

1 

15.25 

Ardops 

1 

15.08 

Ariteus 

1 

14.97 

Desmodontinae 

3 

10.51 

(10.99-10.21) 

0.244 

4.015 

Desmodus 

2 

10.99 

(11.03-10.95) 

0.041 

0.525 

Diphvlla 

1 

10.33 

Diaemus 

1 

10.21 

Mystacinidae 

1 

14.33 

Craseonycteridae 

1 

14.32 

Rhinopomatidae 

1 

13.42 

Noctilionidae 

1 

12.30 

Mormoopidae 

2 

11.18 

(11.36-11.00) 

0.182 

2.302 

Furipteridae 

1 

9.96 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


309 


Table  A14. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  III 

by  the  second  phalanx. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Craseonycteridae 

1 

42.23 

Megadermatidae 

5 

37.92 

(40.32-35.44) 

0.782 

4.612 

Furipteridae 

1 

35.23 

Pteropodidae 

30 

34.13 

(37.82-29.33) 

0.347 

5.566 

Noctilionidae 

1 

30.95 

Nycteridae 

1 

30.05 

Natalidae 

1 

28.70 

Rhinolophidae 

7 

28.03 

(39.31-23.55) 

2.245 

21.185 

All  bats 

153 

25.47 

(42.23-14.21) 

0.528 

25.647 

Emballonuridae 

12 

24.99 

(28.71-21.06) 

0.691 

9.574 

Rhinopomatidae 

1 

24.88 

Phyllostomatidae 

49 

23.62 

(28.68-18.04) 

0.297 

8.810 

Carolliinae 

2 

24.68 

(25.03-24.33) 

0.354 

2.030 

Rhinophylla 

3 

25.03 

(25.93-24.17) 

0.507 

3.508 

Carollia 

4 

24.33 

(25.30-23.44) 

0.396 

3.257 

Stenoderminae 

17 

24.39 

(28.68-21.11) 

0.405 

6.840 

Pvgoderma 

1 

28.68 

Vampyrodes 

1 

25.84 

Ariteus 

1 

25.37 

Stenoderma 

1 

25.18 

Centurio 

1 

25.00 

Ardops 

1 

24.89 

Sphaeronycteris 

1 

24.57 

Vampyrops 

10 

24.51 

(26.17-23.57) 

0.280 

3.610 

Chiroderma 

5 

24.49 

(25.47-23.68) 

0.340 

3.105 

Vampyressa 

5 

24.43 

(25.66-22.83) 

0.469 

4.294 

Artibeus 

13 

24.28 

(25.43-23.52) 

0.164 

2.440 

Ametrida 

2 

24.17 

(24.50-23.84) 

0.330 

1.930 

Uroderma 

2 

23.77 

(24.14-23.41) 

0.362 

2.153 

Ectophylla 

2 

23.67 

(24.51-22.84) 

0.835 

4.989 

Phyllops 

2 

23.23 

(23.27-23.19) 

0.039 

0.236 

Sturnira 

10 

21.44 

(22.84-20.41) 

0.237 

3.494 

Enchisthends 

1 

21.11 

Glossophaginae 

13 

23.73 

(26.05-21.92) 

0.296 

4.504 

Lionyeteris 

1 

26.05 

Anoura 

5 

24.79 

(25.53-24.30) 

0.248 

2.234 

Lichonycteris 

3 

24.60 

(25.07-23.91) 

0.350 

2.467 

Scleronycieris 

1 

24.06 

Hvlonycteris 

1 

23.98 

Choeroniscus 

5 

23.93 

(24.54-23.18) 

0.233 

2.178 

Leptonycteris 

3 

23.87 

(24.54-23.14) 

0.404 

2.934 

Lonchophylla 

5 

23.44 

(24.38-22.69) 

0.272 

2.593 

Monophyllus 

2 

23.27 

(24.09-22.45) 

0.820 

4.982 

Musonycteris 

1 

23.11 

Choeronycteris 

1 

23.03 

Glossophaga 

4 

22.41 

(23.25-21.54) 

0.358 

3.198 

Platalina 

1 

21.92 

Desmodontinae 

3 

23.54 

(26.32-18.80) 

2.385 

17.547 

Diaemus 

1 

26.32 

Diphvlla 

1 

25.52 

1.445 

Desmodus 

2 

18.80 

(18.99-18.60) 

0.192 

Phyllostomatinae 

11 

23.45 

(26.41-20.27) 

0.529 

7.484 

True  hops 

1 

26.41 

Lonchorhina 

3 

25.77 

(26.26-24.97) 

0.402 

2.704 

Mimon 

5 

24.42 

(25.29-23.17) 

0.346 

3.168 

Vampyrum 

1 

23.84 

Phyllostomus 

5 

23.75 

(24.31-22.50) 

0.323 

3.041 

Macrophyllum 

2 

23.42 

(23.43-23.42) 

0.003 

0.019 

Chrotopterus 

1 

23.34 

0.630 

9.612 

Micronvcteris 

12 

22.69 

(27.07-20.16) 

Phylloderma 

1 

22.55 

(22.41-20.36) 

0.302 

3.724 

Tonal  ia 

7 

21.45 

Macrotus 

2 

20.27 

(20.28-20.27) 

0.005 

0.036 

Phyllonycterinae 

3 

18.87 

(20.47-18.04) 

0.803 

7.371 

Brachyphylla 

2 

20.47 

(20.50-20.44) 

0.031 

0.213 

Erophylla 

2 

18.08 

(18.72-17.45) 

0.636 

4.976 

Phyllonycteris 

2 

18.04 

(18.57-17.52) 

0.523 

4.103 

Myzapodidae 

1 

21.59 

0.272 

4.023 

Molossidae 

9 

20.31 

(21.51-18.76) 

Mormoopidae 

2 

19.31 

(23.16-15.46) 

3.850 

28.203 

Vespertilionidae 

31 

19.30 

(33.77-14.21) 

0.790 

22.797 

Mystacinidae 

1 

17.60 

Thyropteridae 

1 

14.86 

310 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A15. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  III 

by  the  third  phalanx. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Mormoopidae 

2 

16.64 

(16.81-16.48) 

0.163 

1.386 

Mystacinidae 

1 

14.70 

Phyllostomatidae 

Desmodontinae 

49 

14.36 

(19.01-10.02) 

0.318 

15.510 

3 

15.42 

(15.71-14.99) 

0.218 

2.449 

D  ip  h  Vila 

1 

15.71 

Desmodus 

2 

15.55 

(16.62-14.47) 

1.076 

9.789 

Diaemus 

1 

14.99 

Stenoderminae 

17 

15.26 

(18.39-10.96) 

0.446 

12.044 

Anietrida 

2 

18.39 

(19.19-17.58) 

0.806 

6.200 

Sturnira 

10 

18.26 

(20.31-16.48) 

0.359 

6.222 

Enchisthenes 

1 

17.13 

Vampyrops 

10 

16.46 

(18.17-14.15) 

0.384 

7.374 

Ardops 

1 

16.45 

Stenodenna 

1 

16.15 

Ectaphylla 

2 

15.44 

(16.49-14.40) 

1.045 

9.574 

Ariteus 

1 

15.32 

Artiheus 

13 

15.16 

(16.79-13.92) 

0.240 

5.708 

Vampyressa 

5 

14.71 

(15.72-14.08) 

0.326 

4.953 

Chiroderma 

5 

14.69 

(15.40-13.98) 

0.241 

3.674 

Phvllops 

2 

14.67 

(16.18-13.16) 

1.510 

14.551 

Sphaeronycteris 

1 

14.52 

Urodermu 

2 

14.22 

(14.41-14.03) 

0.190 

1.887 

Vampyrodes 

1 

13.82 

Centurio 

1 

13.05 

Pvgoderma 

1 

10.96 

Phyilostomatinae 

11 

15.21 

(19.01-10.62) 

0.835 

18.209 

Vamp  yrum 

1 

19.01 

Chrotopterus 

1 

18.59 

Tonal  ia 

7 

17.57 

(19.17-15.18) 

0.467 

7.037 

Macrotus 

2 

16.53 

(16.80-16.27) 

0.265 

2.270 

Mimon 

5 

15.59 

(17.89-12.56) 

0.988 

14.167 

Trachops 

1 

15.23 

Phyllostonius 

5 

15.20 

(16.03-14.18) 

0.335 

4.925 

Phylloderma 

1 

15.18 

Micronycteris 

12 

12.35 

(15.57-  7.56) 

0.671 

18.828 

Macrophyllum 

2 

11.46 

<11.74-1 1.18) 

0.278 

3.429 

Lonchorhina 

3 

10.62 

(11.62-  9.69) 

0.560 

9. 1 30 

Phyllonycterinae 

3 

14.66 

(15.20-14.09) 

0.321 

3.791 

Brachyphvlla 

2 

15.20 

(15.52-14.88) 

0.321 

2.984 

Phyllonycteris 

2 

14.68 

(16.50-12.87) 

1.813 

17.458 

Erophylta 

2 

14.09 

(14.26-13.92) 

0.172 

1.722 

Carolliinae 

2 

14.05 

(14.25-13.85) 

0.200 

2.013 

Rhino  phyllu 

3 

14.25 

(17.40-12.37) 

1.584 

19.259 

Carollia 

4 

13.85 

(14.98-13.10) 

0.430 

6.217 

Glossophaginae 

13 

12.20 

(15.32-10.02) 

0.360 

10.640 

Anoura 

5 

15.32 

(17.46-11.92) 

1.011 

14.753 

Monophyllus 

2 

13.61 

(14.47-12.76) 

0.854 

8.868 

Scleronycteris 

1 

12.81 

Choeronycteris 

1 

12.60 

Musonycteris 

1 

12.31 

Lichonyeteris 

3 

12.18 

(12.36-11.92) 

0.133 

1.885 

Choeroniscus 

5 

12.13 

(13.02-1  1.55) 

0.250 

4.601 

Lonchophylla 

5 

11.94 

(14.28-  9.91) 

0.808 

15.128 

Glossophaya 

4 

11. 90 

(12.71-10.99) 

0.382 

6.415 

Leptonvcteri s 

3 

1 1.73 

(12.72-10.21) 

0.773 

11.415 

Hvlonycteris 

1 

1 1.08 

Piatalina 

1 

10.96 

Lionyceris 

1 

10.02 

Myzapodidae 

1 

13.33 

All  bats 

93 

12.14 

(20.13-  4.64) 

0.378 

30.027 

Noctilionidae 

1 

11.98 

Vespertilionidae 

31 

9.02 

(20.13-  0.00) 

0.672 

41.474 

Thyropteridae 

1 

7.97 

Molossidae 

9 

6.68 

(  8.92-  4.64) 

0.506 

22.711 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


311 


Table  A16. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  IV 

by  the  metacarpal. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

cv 

Thyropteridae 

1 

68.56 

Emballonuridae 

12 

66.37 

(70.34-63.19) 

0.690 

3.600 

Mormoopidae 

2 

65.88 

(66.33-65.44) 

0.442 

0.949 

Nataiidae 

1 

64.78 

Furipteridae 

1 

64.38 

Molossidae 

9 

63.63 

(68.78-54.62) 

1.557 

7.342 

Mystacinidae 

1 

62.95 

Rhinolophidae 

7 

62.92 

(65.78-59.65) 

0.864 

3.633 

Vespertilionidae 

31 

62.67 

(73.59-58.48) 

0.577 

5.128 

Craseonycteridae 

1 

61.46 

Noctilionidae 

1 

60.13 

Nycteridae 

1 

59.88 

Rhinopomatidae 

1 

59.77 

Phyllostomatidae 

Desmodontinae 

49 

58.95 

(67.11-47.84) 

0.425 

5.045 

3 

63.63 

(67.11-61.36) 

1.765 

4.805 

Desmodus 

2 

67.11 

(68.09-66.13) 

0.980 

2.066 

Diphvtla 

1 

62.43 

Diaem  us 

1 

61.36 

Glossophaginae 

13 

60.72 

(63.09-59.66) 

0.263 

1.562 

Hvlonycteris 

1 

63.09 

Lionycleris 

1 

61.70 

Scleronycteris 

1 

61.28 

Anoura 

5 

61.17 

(62.12-58.96) 

0.574 

2.100 

Lichonycteris 

3 

60.85 

(62.43-59.77) 

0.808 

2.299 

Monophyllus 

2 

60.83 

(61.49-60.18) 

0.656 

1.525 

Choeroniscus 

5 

60.46 

(61.05-59.61) 

0.277 

1.024 

Musonycteris 

1 

60.43 

0.483 

Glossophaga 

4 

60.12 

(61.07-58.97) 

1.606 

Lonchophylla 

5 

60.06 

(61.59-58.75) 

0.499 

1.857 

Choeronycteris 

1 

59.84 

Leptonycteris 

3 

59.79 

(60.33-58.78) 

0.507 

1.470 

Platalina 

1 

59.66 

Phyllonycterinae 

3 

59.16 

(59.54-58.72) 

0.237 

0.695 

Phyllonycieris 

2 

59.54 

(60.32-58.77) 

0.775 

1.841 

Brachyphyllu 

2 

59.21 

(59.77-58.64) 

0.568 

1.357 

Eraphylla 

2 

58.72 

(58.88-58.57) 

0.152 

0.366 

Phyllostomatinae 

11 

58.48 

(63.24-52.65) 

1.060 

6.013 

Phyllodermu 

1 

63.24 

Phyllosromus 

5 

63.01 

(64.30-61.83) 

0.485 

1.723 

Lonchorhina 

3 

61.41 

(63.66-59.56) 

1.201 

3.388 

Minion 

5 

60.75 

(64.94-56.94) 

1.495 

5.502 

Micronycteris 

12 

59.79 

(63.15-56.99) 

0.578 

3.347 

Macrophvlluni 

2 

59.14 

(60.04-58.24) 

0.898 

2.147 

Macrotus 

2 

56.79 

(57.44-56-13) 

0.659 

1.641 

Trachops 

1 

55.96 

4.957 

Tonal  ia 

7 

55.93 

(60.05-53.09) 

1.048 

Chrotopterus 

1 

54.63 

Vampyrum 

1 

52.65 

0.628 

4.519 

Stenoderminae 

17 

57.29 

(59.08-47.84) 

Uroderma 

2 

59.08 

(59.11-59.05) 

0.034 

0.081 

Srenoderma 

1 

58.75 

Chiroderma 

5 

58.67 

(60.47-57.08) 

0.735 

2.802 

Artibeus 

13 

58.56 

(59.93-56.96) 

0.229 

1.410 

Phyllops 

2 

58.53 

(59.46-57.60) 

0.929 

2.244 

Ardops 

1 

58.38 

0.173 

Ectophylla 

2 

58.16 

(58.33-57.99) 

0.422 

Vampyressu 

5 

58.12 

(58.83-56.63) 

0.395 

1.521 

Arireus 

1 

58.04 

Vampyrodes 

1 

57.98 

Siurnira 

10 

57.65 

(59.91-55.77) 

0.368 

2.020 

Enchisthenes 

1 

57.63 

Vampvrops 

10 

57.60 

(60.27-56.03) 

0.426 

2.339 

Sphaeronvcteris 

1 

56.84 

Centurio 

1 

56.08 

Pvgoderma 

1 

55.92 

1.241 

Ametrida 

2 

47.84 

(48.26-47.42) 

0.420 

Carolliinae 

2 

56.97 

(57.16-56.77) 

0.195 

0.484 

Caro  lli  a 

4 

57.16 

(58.11-56.20) 

0.459 

1.607 

Rhinophylla 

3 

56.77 

(58.04-55.50) 

0.734 

2.239 

Megadermatidae 

5 

58.66 

(60.84-56.58) 

0.712 

2.715 

All  bats 

153 

58.59 

(73.59-41.96) 

0.567 

11.978 

Myzapodidae 

1 

57.18 

0.451 

5.306 

Pteropodidae 

30 

46.52 

(53.21-41.96) 

312 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A17. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  IV 

by  the  first  phalanx. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

CV 

Pteropodidae 

30 

26.23 

(28.69-22.90) 

0.291 

6.069 

Molossidae 

9 

23.62 

(27.95-17.82) 

1.128 

14.327 

Rhinopomatidae 

1 

22.57 

Nycteridae 

1 

21.62 

Myzapodidae 

1 

20.88 

All  bats 

153 

20.49 

(28.69-10.23) 

0.327 

19.719 

Vespertilionidae 

31 

20.25 

(25.71-13.30) 

0.448 

12.318 

Rhinolophidae 

7 

19.41 

(21.07-15.88) 

0.687 

9.358 

Thyropteridae 

1 

18.93 

Emballonuridae 

12 

18.88 

(22.35-14.01) 

0.684 

12.559 

Phyllostomatidae 

49 

18.10 

(22.93-11.92) 

0.362 

14.008 

Carolliinae 

2 

20.60 

(20.99-20.20) 

0.394 

2.708 

Caro  Ilia 

4 

20.99 

(21.21-20.63) 

0. 1 36 

1.295 

Rhino  phylla 

3 

20.20 

(21.15-19.38) 

0.514 

4.407 

Phyllostomatinae 

11 

19.23 

(22.93-14.33) 

0.818 

14.113 

Mucrotus 

2 

22.93 

(22.98-22.88) 

0.048 

0.298 

Vampyrum 

1 

22.00 

Tonal  ia 

7 

21.84 

(23.10-20.50) 

0.429 

5.197 

Chrotopterus 

1 

21.04 

Micronycteris 

12 

19.48 

(22.83-17.35) 

0.556 

9.888 

Trachops 

1 

19.15 

Macrophyllum 

2 

18.95 

(19.44-18.46) 

0.488 

3.640 

Lonehorhina 

3 

18.49 

(19.91-17.42) 

0.738 

6.914 

Mimon 

5 

18.21 

(20.75-15.92) 

0.886 

10.877 

Phylloderma 

1 

15.17 

Phvllostomus 

5 

14.33 

(16.93-12.10) 

0.910 

14.207 

Stenoderminae 

17 

19.08 

(22.12-16.84) 

0.353 

7.619 

Centurio 

1 

22.12 

Ametrida 

2 

21.31 

(21.35-21.27) 

0.039 

0.260 

Phyllops 

2 

20.42 

(20.47-20.38) 

0.044 

0.306 

Ectophylla 

2 

20.07 

(21.53-18.60) 

1.465 

10.328 

Chiroderma 

5 

19.91 

(21.13-18.80) 

0.416 

4.673 

Vampyressa 

5 

19.55 

(21.48-17.70) 

0.751 

8.592 

Vampyrops 

10 

19.45 

(20.51-18.04) 

0.256 

4.161 

Enchisihenes 

1 

19.26 

Sturnira 

10 

19.03 

(19.89-18.37) 

0.156 

2.590 

Uroderma 

2 

18.90 

(18.93-18.87) 

0.029 

0.217 

Vampyrodes 

1 

18.76 

Pvgodernta 

1 

18.34 

Stenoderma 

1 

18.34 

Artiheus 

13 

17.99 

(19.85-16.12) 

0.349 

6.992 

Ardops 

1 

17.09 

Ariteus 

1 

16.99 

Sphaeronvcteris 

1 

16.84 

Phyllonycterinae 

3 

18.64 

(20.44-17.25) 

0.944 

8.770 

Erophylla 

2 

20.44 

(20.56-20.32) 

0.121 

0.834 

Phyllonycteris 

2 

18.22 

(18.67-17.77) 

0.453 

3.517 

Brachyphvlla 

2 

17.25 

(17.47-17.03) 

0.222 

1.824 

Glossophaginae 

13 

16.75 

(18.58-14.31) 

0.340 

7.314 

Glossophaga 

4 

18.58 

(19.15-18.03) 

0.260 

2.794 

Platalina 

1 

18.35 

Choeronycteris 

1 

17.62 

Scleronycteris 

1 

17.25 

Lonchophylla 

5 

17.22 

(17.99-16.08) 

0.316 

4.102 

Lichonycteris 

3 

17.1 1 

(17.89-16.51) 

0.408 

4.136 

Choeroniscus 

5 

16.99 

(17.71-15.70) 

0.365 

4.807 

Musonyeteris 

1 

16.88 

Leptonycteris 

3 

16.45 

(16.75-16.06) 

0.202 

2.123 

Anoura 

5 

16.17 

(17.15-15.34) 

0.320 

4.420 

Hylonycteris 

1 

16.02 

Monophyllus 

2 

14.81 

(14.92-14.70) 

0.112 

1.074 

Lionycteris 

1 

14.31 

Desmodontinae 

3 

12.05 

(12.14-11.92) 

0.065 

0.931 

Diphylla 

1 

12.14 

Desmodus 

2 

12.09 

(12.63-11.54) 

0.548 

6.412 

Diaemus 

1 

11.92 

Megadermatidae 

5 

17.86 

(19.85-16.18) 

0.746 

9.338 

Natalidae 

1 

17.28 

Mormoopidae 

2 

16.64 

(18.28-15.01) 

1.634 

13.885 

Mystacinidae 

1 

16.01 

Furipteridae 

1 

15.98 

Noctilionidae 

1 

10.61 

Craseonycteridae 

1 

10.23 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


313 


Table  A18. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  IV 

by  the  second  phalanx. 


Taxon 

N 

Mean 

Max-min 

±  1  SE 

CV 

Noctilionidae 

1 

29.27 

Craseonycteridae 

1 

28.31 

Pteropodidae 

30 

27.25 

(29.77-22.67) 

0.335 

6.742 

Megadermatidae 

5 

23.48 

(24.93-19.30) 

1.054 

10.039 

Phyllostomatidae 

49 

22.95 

(30.84-20.10) 

0.280 

8.528 

Desmodontinae 

3 

24.32 

(26.71-20.80) 

0.795 

12.788 

Diaemus 

1 

26.71 

Diphylla 

1 

25.44 

Desmodus 

2 

20.80 

(21.24-20.37) 

0.432 

2.940 

Stenoderminae 

17 

23.63 

(30.84-21.05) 

0.577 

10.061 

Ametrida 

2 

30.84 

(31.22-30.46) 

0.380 

1.745 

Sphaeronycteris 

1 

26.32 

Pygoderma 

1 

25.74 

Ariteus 

1 

24.97 

Ardops 

1 

24.53 

Artibeus 

13 

23.45 

(25.33-21.41) 

0.257 

3.949 

Sturnira 

10 

23.32 

(25.59-21.69) 

0.349 

4.730 

Vampyrodes 

1 

23.26 

Enchisthenes 

1 

23.11 

Vampyrops 

10 

22.95 

(23.94-20.97) 

0.285 

3.923 

Sienoderma 

1 

22.91 

Vampyressa 

Uroaerma 

5 

22.33 

(23.71-20.43) 

0.557 

5.579 

2 

22.02 

(22.08-21.95) 

0.063 

0.405 

Centurio 

1 

21.80 

Ectophvlla 

2 

21.77 

(23.06-20.48) 

1.292 

8.393 

Chiroderma 

5 

21.42 

(23.08-19.05) 

0.712 

7.430 

Phyllops 

2 

21.05 

(21.93-20.16) 

0.885 

5.944 

Glossophaginae 

13 

22.53 

(24.35-20.90) 

0.287 

4.599 

Monophyllus 

2 

24.35 

(24.90-23.81) 

0.544 

3.157 

Lionycteris 

1 

23.99 

Leptonvcteris 

3 

23.76 

(24.47-23.14) 

0.387 

2.822 

Lonchophylla 

5 

22.72 

(25.17-21.29) 

0.658 

6.474 

Musonycteris 

1 

22.70 

Anoura 

5 

22.66 

(23.89-22.09) 

0.321 

3.169 

Choeroniscus 

5 

22.55 

(23.25-22.13) 

0.212 

2.106 

Choeronycteris 

1 

22.54 

Lichonycteris 

3 

22.04 

(23.30-21.06) 

0.662 

5.199 

Platalina 

1 

21.99 

Scleronycteris 

1 

21.47 

4.239 

Glossophaga 

4 

21.29 

(22.03-20.05) 

0.451 

Hvlonycteris 

1 

20.90 

Carolliiriae 

2 

22.43 

(23.02-21.85) 

0.589 

3.716 

Rhinophylla 

3 

23.02 

(25.11-21.88) 

1.046 

7.872 

Carollia 

4 

21.85 

(22.60-21.25) 

0.334 

3.061 

Phyllostomatinae 

11 

22.28 

(25.35-20.10) 

0.555 

8.257 

Vampyrum 

1 

25.35 

Trachops 

1 

24.89 

Chrotopterus 

1 

24.33 

0.692 

6.824 

Phyllostomus 

5 

22.66 

(24.06-20.15) 

Tonal  ia 

7 

22.23 

(23.96-18.74) 

0.773 

9.204 

Macro  phy  llum 

2 

21.91 

(23.30-20.52) 

1.386 

8.945 

Phylloaerma 

1 

21.58 

0.632 

6.713 

Mimon 

5 

21.05 

(22.44-19.13) 

Micronycteris 

12 

20.73 

(22.77-18.64) 

0.359 

6.005 

Macrotus 

2 

20.29 

(20.99-19.58) 

0.707 

4.931 

Lonchorhina 

3 

20.10 

(20.84-18.92) 

0.596 

5.135 

Phyllonycterinae 

3 

22.21 

(23.54-20.84) 

0.781 

6.091 

Brachyphylla 

2 

23.54 

(23.89-23.20) 

0.346 

2.077 

Phyllonycteris 

2 

22.24 

(22.56-21.92) 

0.322 

2.048 

Erophvlla 

2 

20.84 

(21.11-20.57) 

0.272 

1.849 

Myzapodicfae 

1 

21.94 

Mystacinidae 

1 

21.04 

26.054 

All  bats 

153 

20.92 

(31.22-  5.27) 

0.441 

Furipteridae 

1 

19.63 

Nycteridae 

1 

18.51 

Natalidae 

1 

17.94 

Rhinolophidae 

7 

17.67 

(24.48-14.13) 

1.458 

21.842 

Rhinopomatidae 

1 

17.66 

1.192 

9.646 

Mormoopidae 

2 

17.47 

(18.67-16.28) 

Vespertilionidae 

31 

17.08 

(26.56-  5.31) 

0.771 

25.135 

Emballonuridae 

12 

14.75 

(19.47-10.33) 

0.851 

19.977 

Molossidae 

9 

12.75 

(18.11-  5.27) 

1.683 

39.616 

Thyropteridae 

1 

12.51 

314 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A19. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  V 

by  the  metacarpal. 


Taxon 

N 

Mean 

Max-Min 

±  1  SE 

CV 

Noctilionidae 

1 

74.69 

Vespertilionidae 

31 

68.57 

(82.35-59.25) 

0.808 

6.557 

Rhinopomatidae 

1 

68.18 

Mystacinidae 

1 

67.42 

Thyropteridae 

1 

65.80 

Craseonycteridae 

1 

65.57 

Furipteridae 

1 

63.91 

Natalidae 

1 

63.86 

Myzapodidae 

1 

63.85 

Emballonuridae 

12 

63.57 

(67.72-57.70) 

0.811 

4.417 

Phyllostomatidae 

Desmodontinae 

49 

62.12 

(67.99-52.13) 

0.429 

4.838 

3 

66.34 

(67.76-64.29) 

1.049 

2.740 

Diaemus 

1 

67.76 

Desmodus 

2 

66.96 

(67.54-66.38) 

0.578 

1.221 

Diphrlla 

1 

64.29 

Phyllonycterinae 

3 

62.93 

(65.09-61.36) 

1.113 

3.063 

Brachyphylla 

2 

65.09 

(65.97-64.20) 

0.889 

1.932 

Phyllonycteris 

2 

62.36 

(62.76-61.96) 

0.401 

0.910 

Eronhyllu 

Stenoaerminae 

2 

61.36 

(62.06-60.66) 

0.697 

1.606 

17 

62.53 

(64.80-52.13) 

0.785 

5.178 

XJroderma 

2 

64.80 

(65.16-64.44) 

0.362 

0.789 

Stenoderma 

1 

64.67 

Arliheus 

13 

64.58 

(67.64-62.21) 

0.522 

2.916 

Sturnira 

10 

64.52 

(66.05-62.75) 

0.439 

2.153 

Vampvrodes 

1 

64.48 

Chiroderma 

5 

63.89 

(65.51-62.95) 

0.432 

1.514 

Vampyressa 

5 

63.69 

(66.00-61.89) 

0.707 

2.482 

Enchisthenes 

1 

63.67 

Sphaeronvcteris 

1 

63.60 

Vampyrops 

10 

63.50 

(65.47-61.98) 

0.372 

1.854 

Ariteus 

1 

63.48 

Ectophvlla 

2 

63.42 

(64.68-62.16) 

1.258 

2.806 

Ardops 

1 

63.00 

Phyllops 

2 

62.14 

(62.72-61.56) 

0.581 

1.323 

Pvgoderma 

1 

59.15 

Centurio 

1 

58.32 

Ametrida 

2 

52.13 

(52.49-51.76) 

0.366 

0.993 

Phyllostomatinae 

1 1 

61.51 

(67.99-55.66) 

1.213 

6.542 

Phvllostonuts 

5 

67.99 

(70.39-66.08) 

0.880 

2.894 

Phvllodenmi 

1 

67.07 

Minion 

5 

64.20 

(69.16-59.32) 

1.886 

6.570 

Lonchorhina 

3 

63.38 

(65.11-60.19) 

1.600 

4.372 

Macrophyllum 

2 

62.22 

(62.30-62.15) 

0.074 

0.169 

Micronvcteris 

12 

61.67 

(65.46-58.16) 

0.718 

4.030 

Macrotus 

2 

60.48 

(61.61-59.35) 

1.130 

2.643 

True  hops 

1 

59.89 

Chrotopterus 

1 

57.18 

Tonal  in 

7 

56.91 

(60.78-54.38) 

0.928 

4.316 

Vunipvruni 

1 

55.66 

Glossophaginae 

13 

61.22 

(62.70-59.21) 

0.254 

1.495 

Choeronycteris 

1 

62.70 

Leptonycteris 

3 

62.40 

(62.96-61.91) 

0.306 

0.848 

Lonchophylla 

5 

62.01 

(63.12-61.52) 

0.290 

1.046 

Musonycteris 

1 

61.87 

Glossophaya 

4 

61.35 

(61.96-60.62) 

0.291 

0.950 

Choeroniscus 

5 

61.32 

(62.51-59.84) 

0.488 

1.780 

Hvlonycteris 

1 

61.28 

Lionycteris 

1 

60.99 

Lichonycteris 

3 

60.83 

(61.27-60.22) 

0.315 

0.896 

Platalina 

1 

60.77 

Scleronvcteris 

1 

60.76 

Anoura 

5 

60.36 

(61.06-59.07) 

0.379 

1.404 

Monophvllns 

2 

59.21 

(59.65-58.78) 

0.435 

1.040 

Carolliinae 

2 

60.26 

(60.48-60.04) 

0.222 

0.522 

Rhinophylla 

3 

60.48 

(61.79-59.66) 

0.660 

1.890 

Carol  1  in 

4 

60.04 

(60.28-59.59) 

0.154 

0.515 

All  bats 

153 

61.02 

(82.35-46.71) 

0.551 

11.164 

Megadermatidae 

5 

59.85 

(61.13-58.53) 

0.542 

2.027 

Nycteridae 

1 

59.39 

Mormoopidae 

Molossidae 

2 

59.29 

(64.15-54.43) 

4.861 

11.593 

9 

58.88 

(63.56-49.37) 

1.679 

8.557 

Rhinolophidae 

7 

57. 1 3 

(60.70-52.96) 

1.091 

5.052 

Pteropodidae 

30 

50.78 

(53.41-46.71) 

0.295 

3.184 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


315 


Table  A20. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  V 

by  the  first  phalanx. 


Taxon 

N 

Mean 

Max-min 

±1  SE 

cv 

Molossidae 

9 

29.46 

(39.13-20.66) 

1.837 

18.703 

Pteropodidae 

30 

24.16 

(26.06-22.91) 

0.160 

3.636 

Emballonuridae 

12 

23.42 

(29.75-18.26) 

1.038 

15.348 

Furipteridae 

1 

23.22 

Rhinolophidae 

7 

23.12 

(27.43-17.74) 

1.464 

16.759 

Mormoopidae 

2 

22.22 

(26.70-17.74) 

4.480 

28.512 

Nycteridae 

1 

20.58 

All  bats 

153 

20.13 

(39.13-  8.83) 

0.374 

22.977 

Megadermatidae 

5 

20.05 

(25.43-17.91) 

1.395 

15.555 

Myzapodidae 

1 

18.92 

Thyropteridae 

1 

17.75 

Natalidae 

1 

17.61 

Vespertilionidae 

31 

17.44 

(20.61-  8.83) 

0.395 

12.610 

Rhinopomatidae 

1 

17.20 

Noctilionidae 

1 

17.15 

Phyllostomatidae 

49 

16.88 

(22.86-12.74) 

0.303 

12.565 

Carolliinae 

2 

18.61 

(19.00-18.23) 

0.389 

2.952 

Caro  Ilia 

4 

19.00 

(19.62-18.63) 

0.224 

2.359 

Rhinophylla 

3 

18.23 

(19.52-17.06) 

0.715 

6.799 

Phyllostomatinae 

11 

18.25 

(21.50-13.93) 

0.708 

12.858 

Tonal  ia 

7 

21.50 

(23.1  1-20.52) 

0.337 

4.143 

Macrotus 

2 

20.67 

(21.28-20.06) 

0.613 

4.196 

Vampyrum 

1 

19.99 

Chrotopterus 

1 

19.66 

Micronycteris 

12 

19.60 

(21.83-17.53) 

0.478 

8.448 

Trachops 

1 

18.66 

Lonchorhina 

3 

17.47 

(18.81-16.80) 

0.668 

6.617 

Macrophyllum 

2 

17.39 

(17.60-17.18) 

0.211 

1.715 

Mimon 

5 

16.74 

(20.05-14.74) 

1.106 

14.780 

Phylloderma 

1 

15.19 

Phyllostomus 

5 

13.93 

(15.96-12.05) 

0.701 

11.264 

Phyllonycterinae 

3 

16.89 

(19.00-15.58) 

1.067 

10.940 

Erophylla 

2 

19.00 

(19.19-18.81) 

0.190 

1.413 

Brachyphylla 

2 

16.09 

(16.33-15.84) 

0.247 

2.170 

Phyllonycteris 

2 

15.58 

(16.35-14.81) 

0.769 

6.986 

Stenoderminae 

17 

16.71 

(22.86-14.88) 

0.547 

13.491 

Centurio 

1 

22.86 

Ameirida 

2 

21.07 

(21.35-20.79) 

0.279 

1.872 

Sphaeronycteris 

1 

18.90 

Phyllops 

2 

17.26 

(17.42-17.10) 

0.161 

1.317 

Ectophylla 

2 

16.98 

(17.89-16.08) 

0.904 

7.524 

Pyyoderma 

1 

16.64 

Vampyre  ssa 

5 

16.32 

(17.70-14.64) 

0.515 

7.063 

Chiroderma 

5 

16.14 

(16.84-15.08) 

0.320 

4.426 

Vumpyrops 

10 

16.09 

(17.30-15.05) 

0.241 

4.744 

Stenoderina 

1 

15.71 

Vainnyrodes 

1 

15.36 

Uroaerma 

2 

15.27 

(16.09-14.45) 

0.818 

7.576 

Sturnira 

10 

15.24 

(16.57-14.08) 

0.259 

5.366 

Arileu.s 

1 

15.21 

Ardops 

1 

15.10 

Artibeus 

13 

14.94 

(16.89-13.48) 

0.317 

7.652 

Enchisthenes 

1 

14.88 

Glossophaginae 

13 

16.38 

(17.83-14.89) 

0.246 

5.422 

Platalina 

1 

17.83 

Glossophaga 

4 

17.69 

(18.31-16.83) 

0.327 

3.697 

Monophyllus 

2 

17.15 

(17.31-16.99) 

0.162 

1.338 

Scleronycteris 

1 

16.93 

Lonchophylla 

5 

16.66 

(18.16-15.52) 

0.437 

5.862 

Choeroniscus 

5 

16.38 

(17.21-15.19) 

0.332 

4.537 

Lionycteris 

1 

16.34 

Leptonycieris 

3 

16.24 

(16.25-16.23) 

0.006 

0.059 

Lichonycteris 

3 

16.21 

(16.57-15.73) 

0.250 

2.670 

Musonycteris 

1 

15.77 

Choeronycieris 

1 

15.76 

Hylonycteris 

1 

15.11 

Ano  ura 

5 

14.89 

(15.58-14.18) 

0.279 

4.196 

Desmodontinae 

3 

13.75 

(15.13-12.74) 

0.712 

8.970 

Diphylla 

1 

15.13 

Diaeinus 

1 

13.39 

Desmodus 

2 

12.74 

(12.85-12.64) 

0.102 

1.129 

Craseonycteridae 

1 

15.79 

Mystacinidae 

1 

11.59 

316 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  A21. — Ranked  means  and  statistics  for  the  percentage  contributed  to  length  of  digit  V 

by  the  second  phalanx. 


Taxon 

N 

Mean 

Max-min 

±1  SE 

cv 

Pteropodidae 

30 

25.06 

(28.54-22.08) 

0.291 

6.357 

Phyllostomatidae 

49 

21.00 

(26.80-17.49) 

0.275 

9.161 

Glossophaginae 

13 

22.40 

(24.75-20.95) 

0.310 

4.990 

Anoura 

5 

24.75 

(25.35-24.47) 

0.163 

1.473 

Monophyllus 

2 

23.64 

(23.91-23.36) 

0.273 

1.634 

Hvlonycteris 

1 

23.62 

Lichonycteris 

3 

22.96 

(23.44-22.42) 

0.295 

2.229 

Lionycteris 

1 

22.67 

Musonycteris 

1 

22.36 

Scleronycteris 

1 

22.31 

Choeroniscus 

5 

22.30 

(24.97-20.80) 

0.711 

7.129 

Choeronycleris 

1 

21.54 

Platalina 

1 

21.40 

Leptonycteris 

3 

21.36 

(21.84-20.80) 

0.302 

2.448 

Lonchophylla 

5 

21.33 

(22.51-19.98) 

0.464 

4.860 

Glossophaga 

4 

20.95 

(21.82-19.73) 

0.465 

4.442 

Carolliinae 

2 

21.13 

(21.29-20.96) 

0.166 

1.112 

Rhinophylla 

3 

21.29 

(21.90-20.81) 

0.322 

2.620 

Caro  Ilia 

4 

20.96 

(21.17-20.78) 

0.102 

0.970 

Stenoderminae 

17 

20.76 

(26.80-17.49) 

0.509 

10.101 

Ametrida 

2 

26.80 

(26.89-26.72) 

0.087 

0.459 

Pvgodernia 

1 

24.21 

Ardops 

1 

21.89 

Enchisthenes 

1 

21.45 

Art  ileus 

1 

21.31 

Phvt  lops 

2 

20.60 

(21.02-20.18) 

0.421 

2.889 

Ariibeus 

13 

20.47 

(22.13-18.87) 

0.282 

4.964 

Vampyrops 

10 

20.40 

(21.39-19.16) 

0.277 

4.288 

Sturnira 

10 

20.24 

(22.43-18.28) 

0.367 

5.730 

Vampyrodes 

1 

20.16 

Vampyressa 

5 

19.99 

(22.58-17.42) 

1.073 

11.997 

Chirodenna 

5 

19.97 

(21.06-17.72) 

0.578 

6.476 

Uroderma 

2 

19.93 

(20.38-19.47) 

0.456 

3.238 

Stenoderma 

1 

19.61 

Ectophylla 

2 

19.60 

(19.95-19.24) 

0.355 

2.559 

Centurio 

1 

18.82 

Sphaeronycteris 

1 

17.49 

Phyllostomatinae 

11 

20.23 

(24.35-17.75) 

0.649 

10.646 

Vampyrum 

1 

24.35 

Chrotopterus 

1 

23.16 

Tonal  ia 

7 

21.60 

(23.92-18.62) 

0.680 

8.334 

Trachops 

1 

21.45 

Macro  ph  yll  uni 

2 

20.39 

(20.52-20.25) 

0.137 

0.948 

Lonchorhina 

3 

19.14 

(21.00-18.07 

0.934 

8.451 

Minion 

5 

19.06 

(20.93-16.02) 

0.880 

10.327 

Macrotus 

2 

18.85 

(19.37-18.33) 

0.517 

3.878 

Micronycteris 

12 

18.73 

(21.43-16.26) 

0.441 

8.151 

Phyllostomus 

5 

18.08 

(18.90-17.51) 

0.269 

3.323 

Phylloderma 

1 

17.75 

Phyllonycterinae 

3 

20.18 

(22.07-18.83) 

0.973 

8.351 

Phyllonycteris 

2 

22.07 

(22.43-21.70) 

0.368 

2.358 

Erophylla 

2 

19.64 

(20.15-19.13) 

0.507 

3.650 

Brachyphylla 

2 

18.83 

(19.47-18.19) 

0.642 

4.826 

Desmodontinae 

3 

19.91 

(20.58-18.85) 

0.536 

4.663 

Diphylla 

1 

20.58 

Desmodus 

2 

20.30 

(20.78-19.82) 

0.476 

3.319 

Diaeni  us 

1 

18.85 

Mystacinidae 

1 

21.00 

Megadermatidae 

5 

20.09 

(21.41-16.04) 

1.017 

11.315 

Nycteridae 

1 

20.03 

Rhinolophidae 

7 

19.75 

(22.56-18.18) 

0.637 

8.532 

All  bats 

153 

18.84 

(28.54-  8.16) 

0.406 

26.629 

Craseonycteridae 

1 

18.64 

Natalidae 

1 

18.54 

Mormoopidae 

Myzapodidae 

2 

1 

18.48 

17.23 

(18.87-18.10) 

0.380 

2.910 

Thyropteridae 

1 

16.46 

Rhinopomatidae 

1 

14.62 

Vespertilionidae 

31 

13.98 

(21.09-  8.83) 

0.559 

22.239 

Emballonuridae 

12 

13.01 

(16.23-  9.55) 

0.580 

15.451 

Furipteridae 

1 

12.87 

Molossidae 

9 

1  1.66 

(15.78-10.16) 

0.570 

14.663 

Noctilionidae 

1 

8.16 

REPRODUCTIVE  PATTERNS 


Don  E.  Wilson 


“It  follows,  then,  that  an  ecologist  setting  out  to  learn  the  workings  of  some  part 
of  the  natural  world  must  study  the  strategies  of  individual  species.  The  question 
he  must  ask  himself  is:  What  are  the  tricks  used  to  turn  resources  into  babies?” 
This  quotation  from  Colinvaux  (1973)  may  be  appropriate  to  describe  the  fol¬ 
lowing  approach  to  the  problem  of  reproductive  strategies  in  phyllostomatid  bats. 

One  function  of  a  review  paper  is  to  provide  the  reader  with  a  summary  of 
available  data  and  pertinent  references  with  which  to  pursue  the  subject.  I  have 
attempted  to  do  this  at  the  species  level,  although  nothing  is  known  concerning 
reproduction  in  some  species  of  leaf-nosed  bats. 

Knowledge  of  bat  reproductive  patterns  has  undergone  a  spurt  in  growth  in 
recent  years,  as  has  been  the  case  in  many  other  fields.  Enough  information  is  now 
available  to  make  speculation  tantalizing,  but  not  enough  to  make  generalization 
rewarding.  Nevertheless,  some  general  patterns  seem  to  be  widespread  within 
the  primarily  tropical  phyllostomatids. 

Early  knowledge  of  bat  reproduction  was  based  mainly  on  temperate-zone 
species.  Reproductive  cycles  in  temperate  regions  usually  are  forced  into  a  tightly 
controlled  and  relatively  short  time-span  owing  to  rigors  of  the  climate.  Thus, 
most  temperate-zone  bats  produce  only  one  young  per  year,  and  populations  are 
highly  synchronized. 

Some  members  of  the  family  Phyllostomatidae,  such  as  Macrotus  and  Lep- 
tonycteris,  extend  northward  into  subtropical  and  temperate  zones.  Macrotus 
californicus  probably  has  the  most  distinctive  reproductive  pattern  in  the  family. 
Not  only  is  this  species  monestrous  and  monotocous,  but  it  has  a  rather  unique 
system  of  delayed  development  that  allows  the  embryo  to  stay  dormant  during 
unfavorable  times  of  the  year  (Bradshaw,  1961). 

A  variation  of  this  pattern  is  seen  in  Artibeus  jamaicensis  in  Panama  (Fleming, 
1971).  These  animals  have  young  in  March  or  April,  followed  by  a  postpartum 
estrus  and  a  second  pregnancy.  This  results  in  the  second  young  being  born  in 
July  or  August;  another  postpartum  estrus  follows.  The  embryos  from  the  second 
postpartum  estrus  implant  in  the  uterus,  but  undergo  delayed  development  until 
November,  when  they  resume  the  normal  pace  of  development  and  finish  the 
cycle  with  parturition  in  March  or  April. 

A  far  more  common  pattern  is  one  of  bimodal  or  seasonal  polyestry  similar 
to  that  of  Artibeus  jamaicensis,  but  without  delayed  development.  Fleming 
(1973)  and  Wilson  (1973)  have  discussed  this  pattern  for  Panamanian  and 
Costa  Rican  bats.  Members  of  the  genera  Glossophaga,  Carollia,  Uroderma,  and 
Artibeus  commonly  have  this  bimodal  pattern.  In  Panama,  this  pattern  involves 
birth  peaks  in  March-April  and  July-August.  In  Costa  Rica,  the  peaks  may  be 
shifted  to  February-March  and  June-July  (Fleming  et  al.,  1972).  In  Colombia, 


317 


318 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


the  pattern  seems  to  have  shifted  still  more  with  birth  peaks  occurring  in  January- 
February  and  May-June  for  some  species.  A  recent  paper  by  Taddei  (1976)  has 
confirmed  this  pattern  for  many  species  in  Brazil  as  well. 

In  many  of  the  polyestrous  species  there  is  so  much  asynchrony  within  a  given 
population  that  it  is  difficult  to  determine  if  the  individual  animals  are  producing 
more  than  one  young  per  year  or  if  they  simply  are  out  of  phase  with  each  other. 
The  presence  of  females  that  are  both  pregnant  and  lactating  is  one  simple  indica¬ 
tion  of  polyestry. 

It  may  be  possible  for  some  polyestrous  individuals  to  produce  three  young 
per  year,  as  is  the  case  with  Myotis  nigricans,  a  neotropical  vespertilionid  (Wilson 
and  Findley,  1970).  Variation  in  age  of  first  reproductive  activity,  copulation 
time,  fertilization,  gestation  period,  and  timing  of  postpartum  estrus  all  tend  to 
cause  asynchrony  in  a  population.  If  an  individual  bat  became  pregnant  at  the 
onset  of  copulatory  activity  and  proceeded  through  the  first  two  pregnancies  with 
little  or  no  delay,  there  would  be  sufficient  time  for  a  third  pregnancy  in  many 
cases.  I  suspect  that  two  per  year  is  a  more  common  occurrence. 

At  the  other  extreme  from  synchronized  monestrous  cycles  are  year-round 
continuous  reproductive  cycles  as  exemplified  by  the  vampire  Desmodus  rotun- 
dus.  Even  here,  it  is  likely  that  individual  bats  produce  only  two  young  per  year 
on  the  average,  and  asynchrony  within  populations  gives  the  appearance  of 
continuous  activity.  In  Colombia,  for  example,  pregnant,  lactating,  and  in¬ 
active  Artibeus  lituratus  can  be  taken  in  any  month  of  the  year.  Nevertheless, 
there  are  peak  periods  of  pregnancies  in  the  months  of  December  and  May. 

All  of  these  patterns  may  be  viewed  as  variations  on  a  single  theme.  Given  a 
year’s  time,  what  is  the  most  efficient  way  to  produce  offspring?  For  animals 
limited  by  the  rigorous  climates  of  the  temperate  zones,  this  results  in  a  single, 
population-wide  effort  at  the  time  of  maximum  food  availability.  For  tropical 
species,  it  often  might  be  possible  to  produce  two  litters  during  the  favorable 
period  of  food  abundance,  which  is  usually  extended  in  tropical  areas.  Most 
reproductive  patterns  in  tropical  areas  seem  to  be  correlated  with  seasonal  rain¬ 
fall  patterns.  The  dry  season  is  probably  the  most  stressful  time  of  year  for  many 
species,  and  reproductive  strategies  seem  geared  to  avoid  the  weaning  of  young 
during  this  season.  In  polyestrous  species,  the  weaning  of  young  from  the  first 
birth  peak  is  usually  timed  to  coincide  with  the  beginning  of  the  rainy  season,  a 
period  of  maximum  food  abundance.  Desmodus  rotundus  has  probably  been 
allowed  by  natural  selection  to  adopt  a  year-round,  asynchronous  cycle  due  to  the 
year-round  availability  of  its  food  source,  blood  from  domestic  cattle. 

Species  Accounts 

The  following  accounts  are  arranged  in  the  same  order  as  the  list  of  species 
given  by  Jones  and  Carter  (1976).  Each  account  consists  of  a  short  summary  or 
discussion,  to  be  used  in  conjunction  with  the  listing  of  available  data  from  the 
literature  presented  in  tabular  form.  Within  the  tables,  localities,  listed  by  state  or 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


319 


country,  are  arranged  from  North  to  South,  insofar  as  possible.  Dates,  listed  by 
month,  are  arranged  chronologically  from  January  to  December,  although  varying 
dates  in  a  single  reference  necessitate  some  departure  from  the  chronological 
scheme.  Numbers  under  the  columns  labeled  “Pregnant”,  “Lactating”,  and  “In¬ 
active,”  refer  to  number  of  specimens.  An  “X”  in  these  columns  means  the  num¬ 
ber  was  not  given.  In  the  references  column,  “USNM”  refers  to  records  from  the 
National  Museum  of  Natural  History  that  have  not  been  published  previously. 

The  following  list  includes  those  species  for  which  no  data  are  available.  I 
emphasize  these  species  here  and  in  the  species  accounts  in  hopes  that  it  will  spur 
efforts  to  gather  such  data:  Micronycteris  pusilla,  Micronycteris  behni,  Lonchor- 
hina  orinocensis,  Tonatia  brasiliense,  Tonatia  carrikeri,  Tonatia  venezuelae, 
Mimon  bennettii,  Mimon  koepckeae,  Phyllostomus  latifolius,  Lonchophylla 
hesperia,  Lonchophylla  thomasi ,  Anoura  werckleae ,  Scleronycteris  ega,  Licho- 
nycteris  degener,  Platalina  genovensium,  Sturnira  nana,  Vampyrops  aurarius, 
Vampyrops  nigellus,  Vampyrops  recifinus,  Chiroderma  doriae,  Phyllops 
falcatus,  Phyllops  haitiensis,  Ariteus  flavescens,  Ametrida  centurio,  Sphae- 
ronycteris  toxophyllum,  Brachyphylla  nana ,  Phyllonycteris  major. 

Micronycteris  megalotis 

Data  are  insufficient  from  any  one  locality  to  speculate  effectively  on  the  sea¬ 
sonal  reproductive  pattern  of  M.  megalotis.  The  data  are  not  inconsistent  with  a 
pattern  of  seasonal  breeding  in  harmony  with  the  rainfall  pattern.  In  the  northern 
part  of  the  range,  females  are  pregnant  during  the  beginning  of  the  rainy  season. 
In  the  southern  part  of  the  range,  however,  they  become  pregnant  earlier  in  the 
year  and  the  rainy  or  breeding  season  may  last  longer,  possibly  including  two 
breeding  cycles  per  female  per  year.  This  may  be  due  to  an  earlier  and  longer 
lasting  rainy  season  in  the  southern  portions  of  the  range.  See  Table  1 . 

Micronycteris  schmidtorum 

The  only  reference  to  this  species  appears  to  be  that  of  Mares  and  Wilson 
(1971),  who  reported  a  male  with  nonscrotal  testes  taken  in  February  in  Costa 
Rica. 

Micronycteris  minuta 

Data  available  (Table  1)  fit  a  pattern  of  breeding  initiated  at  the  beginning  of 
the  rainy  season.  Confirmation  of  this  pattern  must  await  information  relating  to 
other  seasons.  See  Table  1. 

Micronycteris  hirsuta 

Trinidad  dates  (Table  1)  are  from  the  appropriate  times  of  year  suggesting  at 
least  a  bimodal  reproductive  pattern.  Lack  of  data  from  later  in  the  year  pre¬ 
cludes  further  speculation. 


320 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1. — Reproductive  data  for  the  genus  Micronycteris. 


Place 

Date 

Pregnant  Lactating 

Inactive 

Reference 

Micronycteris  megalotis 

Veracruz 

Feb 

1 

Hall  and  Dalquest,  1963 

Dec 

1 

it 

Jun 

1 

Lackey,  1970 

Yucatan 

Apr 

1 

Jones  et  al.,  1973 

May 

1 

Birney  et  aL,  1974 

Michoacan 

May 

1 

Villa-R.,  1966 

Oaxaca 

May 

1 

it 

El  Salvador 

Mar 

1 

Burt  and  Stirton,  1961 

Nicaragua 

Mar 

2 

1 

Jones  et  al.,  1971a 

Apr 

2 

n 

Jun 

1  2 

1 

tt 

Aug 

2 

tt 

Costa  Rica 

Feb 

1 

Gardner  et  al.,  1970 

Panama 

May 

1 

Enders,  1935 

Trinidad 

Feb 

1 

Goodwin  and  Greenhall,  1961 

Mar 

1 

it 

Jun 

1 

it 

Colombia 

Jun 

1 

Thomas,  1972 

Venezuela 

Jul 

1 

USNM 

Aug 

3 

it 

Peru 

Aug 

2 

18 

Tuttle,  1970 

Brazil 

Jun 

3 

Peracchi  and  Albuquerque,  1971 

Micronycteris  minuta 

Costa  Rica 

Mar 

1 

Gardner  et  al.,  1970 

Trinidad 

Mar 

1 

Goodwin  and  Greenhall,  1961 

May 

1  4 

2 

it 

Peru 

Jul 

2 

Tuttle,  1970 

Micronycteris  hirsuta 

Trinidad 

Mar 

1 

Goodwin  and  Greenhall,  1961 

May 

2 

tt 

Peru 

Jul 

1 

1 

Tuttle,  1970 

Micronycteris  sylvestris 

Nayarit 

Mar 

1 

Jones,  1964  ft 

Veracruz 

Dec 

1 

Hall  and  Dalquest,  1963 

French  Guiana 

Feb 

X 

Brosset  and  Dubost,  1967 

Mar 

X 

it 

Micronycteris  brachyotis 

Goodwin  and  Greenhall  (1961)  reported  a  “breeding  male”  in  May  and  three 
others  in  June  from  Trinidad.  Rick  (1968)  found  one  pregnant  and  six  lactating 
females  in  July  in  Guatemala. 

Micronycteris  pusilla 

Nothing  is  known  about  reproduction  in  M.  pusilla. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


321 


Micronycteris  nicefori 

The  only  records  are  those  of  Goodwin  and  Greenhall  (1961),  reporting  two 
“breeding  males”  from  Trinidad  in  October,  and  Baker  and  Jones  (1975),  re¬ 
porting  a  lactating  female  from  Nicaragua  in  July. 

Micronycteris  sylvestris 

In  the  northern  part  of  the  range,  known  records  are  from  late  in  the  rainy 
season,  whereas  from  the  southern  portion  they  are  from  early  in  the  rainy  season. 
Data  from  other  times  of  the  year  are  necessary  before  speculating  further.  See 
Table  1. 

Micronycteris  behni 

Nothing  is  known  about  the  reproductive  pattern  of  M.  behni. 

Micronycteris  daviesi 

Tuttle  (1970)  collected  a  pregnant  female  in  August  in  Peru.  This  is  ap¬ 
parently  the  only  record  of  reproductive  activity  for  this  species. 

Macrotus  waterhousii 

In  Mexico,  M.  waterhousii  probably  has  a  single  young  per  year  (Table  2). 
The  available  evidence  suggests  May  as  the  most  likely  period  for  parturition. 
Additional  study  may  reveal  a  delayed  development  system  such  as  that  de¬ 
scribed  for  the  congeneric  M.  californicus  in  the  following  account.  Data  from  the 
Caribbean  populations  are  insufficient  for  any  meaningful  analysis. 

Macrotus  californicus 

In  addition  to  the  information  in  Table  2,  Bradshaw  (1961,  1962)  has  de¬ 
scribed  the  reproductive  strategy  of  M.  californicus  in  southern  Arizona.  A  good 
summary  of  the  reproductive  pattern  also  may  be  found  in  Anderson  (1969). 
Males  undergo  spermatogenesis  in  summer  and  autumn  and  inseminate  females  in 
autumn;  ovulation  and  fertilization  occur  immediately  following  copulation.  The 
single  embryo  undergoes  slow  growth  during  winter  until  March,  when  develop¬ 
ment  proceeds  at  a  more  rapid  rate  resulting  in  a  gestation  period  of  about  eight 
months.  Bradshaw  (1961)  coined  the  term  “delayed  development”  to  describe 
the  reproductive  pattern.  Parturition  occurs  in  June  and  young  are  foraging  by 
August.  Young-of-the-year  females  apparently  breed  during  the  first  autumn, 
but  males  are  not  reproductively  mature  until  the  following  year. 

Lonchorhina  aurita 

The  little  evidence  available  points  to  a  breeding  season  that  is  correlated  with 
the  beginning  of  the  rainy  season  (Table  2).  Panamanian  pregnancies  are  during 
the  dry  season  and  should  result  in  the  young  being  born  at  the  beginning  of  the 
rainy  season. 


322 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  2. —  Reproductive  data  for  the  genera  Macrotus,  Lonchorhina,  Macrophyllum, 

Tonatia,  andMimon. 


Place  Date  Pregnant  Lactating  Inactive  Reference 


Macrotus  waterhousii 

Sinaloa 

Jul 

X 

Jones  et  al.,  1972 

Jalisco 

Feb 

3 

X 

Watkins  et  al.,  1972 

Mar 

2 

X 

ft 

May 

X  X 

it 

Jul 

X 

n 

Sep 

X 

it 

Oct 

X 

it 

Tres  Marias  Is. 

May 

X 

Merriam,  1898 

Durango 

Jun 

2  1 

Jones,  1964  c 

Jamaica 

Dec 

7  2 

Osburn,  1865 

Dec 

4 

Goodwin,  1970 

Crooked  Is. 

Apr 

1 

Buden,  1975 

Cuba 

Mar 

4 

Anderson,  1969 

Caicos  Is. 

Feb 

2 

Buden,  1975 

Apr 

1 

Macrotus  californicus 

Miller,  1904 

California 

Mar 

1 

Cockrum,  1955 

Apr 

2* 

It 

Apr 

9 

USNM 

Apr 

60 

Grinnell,  1918 

May 

1 

Huey,  1925 

Baja  Calif. 

Jul 

5 

X 

Jones  et  al.,  1965 

Sonora 

Apr 

X 

Burt,  1938 

May 

4 

It 

Jul 

X 

It 

Aug** 

It 

Mar 

1 

3 

Cockrum  and  Bradshaw,  1963 

Apr 

6 

Lonchorina  aurita 

It 

Quintana  Roo 

Aug 

1 

Jones  et  al.,  1973 

Oaxaca 

Feb 

X 

Walker,  1975 

Mar 

8 

15 

Schaldach,  1965 

Guatemala 

Jan 

1 

Jones,  1966 

Panama 

Feb 

2 

Bloedel,  1955 

Mar 

2 

It 

Feb 

2 

Fleming  et  al.,  1972 

Mar 

2 

It 

Nov 

1 

It 

Trinidad 

Apr 

1 

Goodwin  and  Greenhall,  1961 

Peru 

Jul 

1 

Tuttle,  1970 

Aug 

1 

It 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


323 


Table  2. — Continued. 


Macrophyllum  macrophyllum 


El  Salvador 

Oct 

2 

Harrison,  1975 

Costa  Rica 

Mar 

X 

LaVal,  1977 

May 

X 

II 

Aug** 

II 

French  Guiana 

Oct 

X 

Brosset  and  Dubost,  1967 

Nov 

X 

Tonatia  bidens 

II 

Guatemala 

Feb 

1 

Carter  et  aL,  1966 

Honduras 

Aug** 

2 

2 

Valdez  and  LaVal,  1971 

Costa  Rica 

Jan 

1 

Gardner  et  aL,  1970 

Aug** 

LaVal,  1977 

Trinidad 

May 

2 

Goodwin  and  Greenhall,  1961 

Peru 

Apr 

2 

Gardner,  1976 

Jul 

2 

1 

ft 

Tonatia  minuta 

Honduras 

Aug 

1 

Valdez  and  LaVal,  1971 

Nicaragua 

Jul 

1 

II 

Costa  Rica 

Feb 

1 

LaVal,  1977 

Apr 

1 

It 

Panama 

Feb 

1 

Tonatia  silvicola 

Davis  et  al.,  1964 

Panama 

Mar 

2 

Fleming  et  al.,  1972 

Oct 

1 

It 

Nov 

1 

II 

Dec 

1 

It 

Colombia 

Jan 

1 

Thomas,  1972 

Peru 

Jul 

2 

Tuttle,  1970 

Aug 

2 

Mimon  cozumelae 

It 

Veracruz 

Apr 

2 

2 

Hall  and  Dalquest,  1963 

Yucatan 

Apr 

19 

Jones  et  al.,  1973 

Jul 

1 

II 

Campeche 

May 

X 

II 

Guatemala 

Mar 

1 

Rick,  1968 

Aug 

1  1 

II 

Honduras 

Jul 

1 

Valdez  and  LaVal,  1971 

Costa  Rica 

Apr 

X 

LaVal,  1977 

Aug 

1 

Mimon  crenulatum 

II 

Campeche 

Feb 

1 

Jones,  19646 

Costa  Rica 

Apr 

1 

LaVal,  1977 

Venezuela 

Mar 

2 

Goodwin  and  Greenhall,  1961 

Peru 

Jul 

2 

Tuttle,  1970 

•One  with  twins. 
••Young  taken. 


324 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Lonchorhina  orinocensis 

Nothing  is  known  about  reproduction  in  L.  orinocensis. 

Macrophyllum  macrophyllum 

Felten  (1956a)  postulated  that  this  species  breeds  in  the  dry  season.  The 
finding  of  pregnant  animals  during  the  late  rainy  season  in  French  Guiana  is 
unusual  when  compared  with  the  cycles  in  other  members  of  the  subfamily.  See 
Table  2. 

Tonatia  bidens 

Although  the  records  are  scattered,  I  suspect  that  this  species  breeds  more  than 
once  a  year  (Table  2).  Records  from  Honduras  suggest  a  bimodal  pattern  with 
subadult  animals  representing  the  earlier  breeding  cycle. 

Tonatia  brasiliense 

Nothing  is  known  about  the  reproductive  pattern  of  this  species. 

Tonatia  carrikeri 

Nothing  is  known  about  the  reproductive  pattern  of  T.  carrikeri,  although 
Gardner  (1976)  reported  two  reproductively  inactive  females  from  Peru  in  July. 

Tonatia  minuta 

This  species  also  appears  to  fit  the  bimodal  pattern,  although  additional  data 
are  obviously  necessary  to  confirm  this  hypothesis.  See  Table  2. 

Tonatia  silvicola 

Females  appear  to  give  birth  during  the  early  half  of  the  rainy  season;  there  is 
thus  far  no  evidence  of  more  than  one  young  per  year.  See  Table  2. 

Tonatia  venezuelae 

No  information  is  available  on  reproduction  in  this  species. 

Minion  bennettii 

Nothing  is  known  about  the  reproductive  pattern  of  this  bat. 

Mimon  cozumelae 

This  species  (Table  2)  apparently  produces  young  at  the  beginning  of  the  rainy 
season  and  the  available  data  suggest  only  a  single  young  per  year. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


325 


Mimon  crenulatum 

Records  from  Campeche  and  Venezuela  are  from  the  dry  season,  whereas 
Peruvian  records  are  from  the  rainy  season.  The  single  record  from  Costa  Rica 
was  taken  in  the  period  of  transition  between  dry  and  rainy  seasons.  See  Table  2. 

Mimon  koepckeae 

No  data  are  available  on  the  reproductive  pattern  of  this  species. 

Phyllostomus  discolor 

In  addition  to  records  listed  in  Table  3,  Mares  and  Wilson  (1971)  found  80 
per  cent  of  43  animals  in  1968  and  51  per  cent  of  69  animals  in  1970  to  be  re- 
productively  active  during  February  and  March  in  Costa  Rica.  Tamsitt  (1966) 
stated  that  in  Colombia  this  species  is  acyclic  or  continuous  in  its  breeding  habits. 
Most  of  the  above  data  suggest  this  pattern  for  other  areas  as  well;  however,  the 
lack  of  reproductive  activity  as  noted  by  Fleming  et  al.  (1972)  for  Costa  Rica 
seems  unusual.  Heithaus  et  al.  (1975)  suggested  that  P.  discolor  may  be  mon- 
estrous  in  Costa  Rica. 

Phyllostomus  hastatus 

Starrett  and  de  la  Torre  (1964)  reported  that  one  of  two  July-taken  males 
from  Costa  Rica  had  small,  inguinal  testes,  and  the  other  had  large,  scrotal  testes; 
both  were  in  an  early  stage  of  spermatogenesis  with  no  mature  sperm  in  the  testes. 

The  available  data  could  support  either  a  monestrous  (in  Nicaragua,  Panama, 
and  Trinidad)  or  polyestrous  (in  Colombia)  pattern.  In  fact,  this  may  be  a 
species  in  which  the  reproductive  strategy  varies  geographically.  See  Table  3. 

Phyllostomus  elongatus 

Additional  data  from  times  of  the  year  other  than  those  listed  in  Table  3  are 
needed  to  elucidate  the  pattern  of  this  species.  The  above  data  show  that  these 
animals  breed  during  the  middle  part  of  the  rainy  season. 

Phyllostomus  latifolius 

Nothing  is  known  about  the  reproductive  pattern  of  this  species. 

Phylloderma  stenops 

The  only  report  of  reproductive  activity  for  this  rare  species  is  that  of  LaVal 
(1977),  who  reported  a  pregnant  female  in  February  (embryo  length,  33  mm.) 
from  Costa  Rica.  Gardner  (1976)  reported  a  reproductively  inactive  female  from 
Peru  that  was  collected  in  May. 


326 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  3. — Reproductive  data  for  the  genera  Phyllostomus,  Trachops,  Chrotopterus,  and 

Vampyrum. 


Place 

Date 

Pregnant  Lactating 

Inactive 

Reference 

Phyllostomus  discolor 

Guatemala 

Mar 

1 

Jones,  1966 

El  Salvador 

Feb 

2 

4 

Felten,  1956  h 

Jun 

14 

29 

tt 

Aug 

11 

7 

tt 

Sep 

22 

43 

it 

Nov 

14 

X 

70 

it 

Dec 

13 

Burt  and  Stirton,  1961 

Nicaragua 

Mar 

2 

Jones,  1964a 

Costa  Rica 

Jan 

1 

Fleming  et  al.,  1972 

Mar 

1 

11 

ft 

Apr 

3 

If 

May 

6 

II 

Jul 

3 

II 

Dec 

11 

It 

Jul 

1 

Tamsitt  and  Valdivieso,  1961 

Trinidad 

Feb 

X 

Goodwin  and  Greenhall,  1961 

Mar 

X 

ft 

Jun 

X 

It 

Aug 

X 

X 

ft 

Sep 

X 

ft 

Oct 

X 

tt 

Colombia 

Feb* 

2 

2 

1 

Tamsitt  and  Valdivieso,  1964 

Mar 

3 

tt 

May 

1 

1 

tt 

Sep 

1 

ft 

Oct* 

3 

ft 

Venezuela 

Jul 

1 

2 

2 

Smith  and  Genoways,  1974 

Brazil 

Jul 

1 

Walker,  1975 

Phyllostomus  hastatus 

Nicaragua 

Mar 

2 

Jones  et  al.,  1971a 

Jun 

X 

ft 

Jul 

X 

tt 

Aug 

X 

tt 

Panama 

Apr 

1 

Fleming  et  al.,  1972 

May 

1 

tt 

Jun 

2 

tt 

Oct 

1 

tt 

Trinidad 

Mar 

X 

Goodwin  and  Greenhall,  1961 

Apr 

X 

X 

tt 

Jun 

X 

ft 

Sep 

X 

tt 

Nov 

X 

tt 

Venezuela 

Aug 

X 

USNM 

Colombia 

Mar 

1 

1 

Thomas,  1972 

May 

1 

tt 

Aug 

1 

tt 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


327 


•  Table  3. — Continued. 

Colombia 

Sep 

1 

2  1 

Thomas,  1972 

Oct 

2 

2 

tt 

Nov 

1 

tt 

Dec 

1  5 

tt 

Jul 

1 

7 

Arata  and  Vaughn,  1970 

Peru 

Jun 

1 

Tuttle,  1970 

Aug 

12 

8 

tt 

Brazil 

Aug 

1 

Phyllostomus  elongatus 

Peracchi  and  Albuquerque,  1971 

Colombia 

Jun 

1 

1 

Thomas,  1972 

Peru 

Jul 

6 

3 

Tuttle,  1970 

Aug 

1 

Trachops  cirrhosus 

tt 

Veracruz 

Apr 

1 

Hall  and  Dalquest,  1963 

Campeche 

Feb 

1 

Jones  et  al.,  1973 

Oaxaca 

Mar 

1 

Villa- R,  1966 

Chiapas 

Dec 

1 

ft 

Mar 

1 

Carter  et  al.,  1966 

Guatemala 

Mar 

4 

3 

Jones,  1966 

Apr 

6 

1 

ft 

El  Salvador 

Feb 

3 

Burt  and  Stirton,  1961 

Honduras 

Aug 

1 

Valdez  and  LaVal,  1971 

Nicaragua 

May 

4 

Carter  et  al.,  1966 

Costa  Rica 

Mar 

1 

ft 

Aug 

1  6 

Armstrong,  1969 

Panama 

Aug 

1 

Fleming  et  aL,  1972 

Oct 

1 

tt 

Nov 

1 

ft 

Trinidad 

Mar 

2 

Goodwin  and  Greenhall,  1961 

Peru 

Jul 

1 

Chrotopterus  auritus 

Tuttle,  1970 

Veracruz 

Apr 

1 

Hall  and  Dalquest,  1963 

Yucatan 

Apr 

1 

1 

Jones  et  al.,  1973 

Jul 

1  1 

tt 

Argentina 

Jul 

1 

Vampyrum  spectrum 

Villa-R.  and  Villa-C.,  1969 

Costa  Rica 

Aug 

1 

Gardner  et  aL,  1970 

Trinidad 

May 

1 

Goodwin  and  Greenhall,  1961 

‘Pregnant  and  Iactating. 


Trachops  cirrhosus 

Felten  (1956a)  stated  that  T.  cirrhosus  breeds  in  the  dry  season  in  El  Salvador, 
and  the  data  of  Burt  and  Stirton  (1961)  support  this.  This  species  may  have  an 
extended  season,  or  may  be  geographically  variable  with  regard  to  the  reproduc¬ 
tive  cycle.  Additional  data  on  other  seasons  from  any  of  the  above  localities  would 
be  useful.  See  Table  3. 


328 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Chrotopterus  auritus 

Data  are  insufficient  (Table  3)  to  allow  speculation  on  the  possible  reproduc¬ 
tive  pattern  of  C.  auritus  except  to  note  that  this  species  produces  young  during 
the  early  part  of  the  rainy  season. 

Vampyrum  spectrum 

The  only  information  available  other  than  that  in  Table  3  seems  to  be  Green- 
hall’s  (1968)  report  on  a  birth  in  captivity.  Again,  other  than  the  fact  that  V. 
spectrum  produces  young  during  the  rainy  season,  little  can  be  said  about  its  re¬ 
productive  cycle. 

Glossophaga  soricina 

Cockrum  (1955)  believed  G.  soricina  to  be  polyestrous,  with  the  young  born 
at  any  time  of  the  year  in  Mexico.  Fleming  (1973)  felt  that  this  species  is  sea¬ 
sonally  polyestrous  in  Panama,  with  bimodal  birth  peaks  occurring  in  March- 
April  and  July- August.  Tamsitt  (1966)  indicated  that  G.  soricina  is  acyclic  or 
continuously  breeding  in  Colombia.  Felten  (1956a)  noted  that  this  species  breeds 
throughout  the  year  in  El  Salvador.  Heithaus  et  al.  (1975)  suggested  bimodal 
polyestry  for  Costa  Rican  animals. 

This  is  one  of  the  few  species  of  phyllostomatid  bats  for  which  a  fair  amount  of 
reproductive  data  are  available  from  a  variety  of  localities  (Table  4).  The  data 
suggest  G.  soricina  is  polyestrous  in  most  areas.  Reproduction  may  be  somewhat 
geographically  variable  inasmuch  as  data  from  Panama  indicate  no  pregnancies 
during  the  period  August-December.  Also,  in  some  of  the  areas  where  these  bats 
appear  to  breed  continuously,  there  may  well  be  a  bimodal  pattern  for  individuals 
but  enough  asynchrony  within  the  population  to  allow  for  individuals  in  all  stages 
of  the  reproductive  cycle  to  be  collected  at  any  given  time. 

Rasweiler  (1972)  demonstrated  this  species  to  be  polyestrous  with  approxi¬ 
mately  a  24-day  cycle  in  captivity.  He  described  the  preimplantation  development 
and  histology  of  the  oviduct  in  some  detail. 

Glossophaga  alticola 

I  can  find  no  reproductive  information  for  G.  alticola  in  the  literature.  The 
National  Museum  of  Natural  History  has  two  specimens  taken  in  Oaxaca  in 
April,  one  of  which  was  pregnant  and  the  other  inactive. 

Glossophaga  commissarisi 

The  data  available  (Table  4)  are  not  inconsistent  with  a  pattern  of  bimodal 
polyestry.  Although  the  data  are  scanty,  the  information  from  Jalisco  supports 
this  hypothesis. 

Glossophaga  longirostris 

This  species  appears  to  breed  during  the  rainy  season,  but  the  data  are  in¬ 
conclusive  (Table  4). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


329 


Table  4. —  Reproductive  data  for  the  gem/s  Glossophaga. 


Place 

Date 

Pregnant 

Lactating 

Inactive 

Reference 

Glossophaga  soricina 

Sonora 

May 

3 

Cockrum,  1955 

Dec 

1 

2 

Cockrum  and  Bradshaw,  1963 

Chihuahua 

Jul 

2 

Anderson,  1972 

Durango 

Jun 

3 

Jones,  1964  c 

San  L.  Potosi 

Jun 

X 

Dalquest,  1953 

Sinaloa 

Jan 

X 

Jones  et  ai,  1972 

Mar 

X 

tt 

May 

X 

tt 

Aug 

X 

tt 

Sep 

X 

ft 

Oct 

X 

tt 

Nov 

X 

tt 

Dec 

X 

tt 

Nayarit 

Jan 

1 

Cockrum,  _1 955 

Feb 

5 

tt 

Aug 

1 

ft 

Jalisco 

Feb 

X 

Watkins  et  al„  1972 

Mar 

X 

tt 

Apr 

X 

tt 

Sep 

X 

tt 

Oct 

X 

tt 

Tres  Marias  Is. 

May 

X 

Merriam,  1898 

Colima 

Nov 

1 

2 

Villa-R.,  1966 

Dec 

X 

ft 

Queretaro 

Jan 

5 

Schmidly  and  Martin,  1973 

Dec 

6 

tt 

Puebla 

Jan 

1 

1 

LaVal,  1972 

Veracruz 

Mar 

X 

Hall  and  Dalquest,  1963 

Apr 

1 

7 

tt 

Sep 

4 

5 

tt 

Nov 

X 

ft 

Jun 

2 

Lackey,  1970 

Jul 

1 

tt 

Tabasco 

May 

X 

Villa-R.,  1966 

Jun 

X* 

X 

tt 

Jul 

X 

X 

ft 

Yucatan 

Feb 

X** 

x** 

Jones  et  al.,  1973 

Apr 

X*  * 

X** 

ft 

Jul 

X** 

X** 

ft 

Apr 

1 

4 

Birney  et  al.,  1974 

Aug 

X 

Pearse  and  Kellogg,  1938 

Oaxaca 

Mar 

3 

3 

USNM 

Apr 

4 

6 

tt 

Sep 

1 

Cockrum,  1955 

Chiapas 

Feb 

X 

Villa-R.,  1966 

Aug 

X*** 

Barlow  and  Tamsitt,  1968 

Guatemala 

Mar 

1 

Jones,  1966 

Aug 

2 

5 

tt 

330 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  4. — Continued. 


El  Salvador 

Jan 

22 

1 

28 

Felten,  1956ft 

Feb 

2 

22 

tt 

Mar 

2 

4 

5 

rt 

Apr 

1 

7 

rt 

Jun 

17 

ft 

Jul 

9 

12 

tt 

Aug 

17 

3 

5 

rt 

Sep 

5 

3 

it 

Oct 

3 

3 

rt 

Nov 

1 

1 

21 

rt 

Dec 

3 

3 

rt 

Sep 

1 

Burt  and  Stirton,  1961 

Nov 

1 

ft 

Jul 

7 

Starrett  and  de  la  Torre,  1964 

Honduras 

Aug 

1 

tt 

Costa  Rica 

Jul 

1 

If 

Aug 

1 

1 

tt 

Jul 

X 

Tamsitt  and  Valdivieso,  1961 

Aug 

X 

tt 

Panama 

Jan 

8 

6 

Fleming  et  at.,  1972 

Feb 

22 

ft 

Mar 

1 

2 

tt 

Apr 

1 

It 

May 

3 

tt 

Jun 

3 

1 

It 

Jul 

2 

5 

tt 

Aug 

3 

5 

tt 

Sep 

4 

rt 

Oct 

4 

tt 

Nov 

4 

rt 

Dec 

4 

tt 

Feb 

1 

Bloedel,  1955 

Jamaica 

Jan 

1 

4 

Goodwin,  1970 

Trinidad 

Jan 

X 

X 

Goodwin  and  Greenhall,  1961 

Feb 

X 

tt 

Mar 

X 

tt 

Apr 

X 

ft 

May 

X 

tt 

Jun 

X 

X 

ft 

Dec 

X 

tt 

Venezuela 

Aug 

1 

USNM 

Colombia 

Jan 

2 

Thomas,  1972 

Feb 

8 

2 

It 

Mar 

5 

3 

tt 

Apr 

4 

5 

ft 

May 

7 

tt 

Jun 

1 

2 

tt 

Jul 

1 

1 

If 

Aug 

1 

tt 

Sep 

2 

2 

tt 

Oct 

1 

6 

5 

tt 

Nov 

6 

It 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


331 


Table  4. — Continued. 


Colombia 

Dec 

5 

1 

11 

Thomas,  1972 

Jan 

1 

1 

Tamsitt  and  Valdivieso,  1964 

Mar 

1 

ft 

Apr 

8 

1 

ft 

Jun 

1 

ft 

Jul 

2 

tt 

Aug 

I 

tt 

Sep 

1 

1 

tt 

Oct 

2 

ft 

Nov 

7 

13 

ft 

Dec 

3 

1 

1 

tt 

Jul 

32 

23 

Arata  and  Vaughn,  1970 

Aug 

8 

13 

tt 

French  Guiana 

Feb 

X 

Brosset  and  Dubost,  1967 

Mar 

X 

it 

Oct 

X 

ft 

Peru 

Jun 

2 

Tuttle,  1970 

Jul 

1 

1 

ft 

Aug 

1 

ft 

Brazil 

Nov 

X 

Hamlett,  1935 

Dec 

X 

tt 

Jan 

X 

X 

Peracchi  and  Albuquerque,  1971 

Paraguay 

Oct 

1 

USNM 

Glossophaga  commissarisi 

Durango 

Jul 

1 

Baker  and  Greer,  1962 

Sinaloa 

Jan 

4 

Jones  et  ai,  1972 

Jul 

1 

4 

it 

Jalisco 

Feb 

1 

Watkins  et  al„  1972 

Apr 

2 

1 

tt 

May 

2 

it 

Jul 

3 

tt 

Sep 

1 

1 

tt 

Nov 

2 

tt 

Dec 

1 

tt 

Guatemala 

Feb 

1 

Jones,  1966 

Nicaragua 

Feb 

1 

3 

Jones,  1964a 

Glossophaga  longirostris 

Trinidad 

Feb 

X 

Goodwin  and  Greenhall,  1961 

Mar 

X 

tt 

Apr 

X 

it 

Aug 

X 

ft 

Sep 

X 

ft 

Jun 

X 

Goodwin,  1958 

Venezuela 

Jul 

2 

USNM 

Jul 

8 

6 

Smith  and  Genoways,  1974 

•Parturition. 
••Pregnant  or  lactating. 
•••Twins. 


332 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Monophyllus  redmani 

Goodwin  (1970)  felt  that  the  high  percentage  of  pregnancies  in  his  sample 
suggested  a  discrete  breeding  season  for  this  species.  Additional  data  from  other 
times  of  the  year  are  needed  in  order  to  verify  his  opinion.  See  Table  5. 

Monophyllus  plethodon 

Schwartz  and  Jones  (1967)  reported  pregnant  females  from  Dominica  in 
March  and  April,  perhaps  indicating  a  distinct  breeding  season  for  this  species. 

Leptonycteris  nivalis 

Davis  (1966)  reported  that  the  breeding  season  is  restricted  to  April,  May, 
and  June.  Easterla  (1972)  felt  that  young  probably  are  born  in  Mexico,  possibly 
in  June,  prior  to  the  time  bats  arrive  in  the  Big  Bend  area  of  Texas.  Records  from 
Veracruz  (Table  5)  indicate  a  second  pregnancy  of  the  year  for  this  migratory 
species. 

Leptonycteris  sanborni 

Cockrum  and  Ordway  (1959)  and  Hayward  and  Cockrum  (1971)  have  re¬ 
ported  on  reproduction  in  L.  sanborni  in  Arizona.  They  found  that  pregnant  fe¬ 
males  arrive  in  southern  Arizona  in  early  May  and  the  young  are  bom  shortly 
thereafter.  By  August  there  are  subadult  females  containing  embryos  10  mm.  in 
crown-rump  length;  however,  all  bats  have  left  for  Mexico  by  the  early  part  of 
October.  They  hypothesized  another  birth  peak  in  Mexico  in  early  November. 
In  January  females  have  small  embryos  and  in  February  they  begin  to  move  to 
the  northern  part  of  their  range  where  the  young  will  be  bom.  Hayward  and  Cock¬ 
rum  (1971)  suggested,  as  an  alternative  hypothesis,  that  delayed  development  as 
described  for  Macrotus  waterhousii  might  be  involved.  See  Table  5. 

Leptonycteris  curasoae 

Smith  and  Genoways  (1974)  found  a  large  colony  of  this  species  on  Margarita 
Island,  Venezuela,  which  in  July  was  estimated  to  contain  4000  females  nursing 
nearly  full-grown  young.  In  November,  seven  of  34  females  examined  were  preg¬ 
nant,  and  no  juveniles  were  present.  In  addition,  adult  males  with  large  (6  to  8 
mm.)  testes  were  present  in  November,  whereas  males  had  been  absent  in  July. 
This  appears  to  be  the  only  record  of  reproduction  for  this  species. 

Lonchophylla  Hesperia 

Nothing  is  known  about  the  reproductive  pattern  of  this  species. 

Lonchophylla  mordax 

Thomas  (1972)  collected  a  reproductively  inactive  female  of  this  species  in 
Colombia  in  January.  I  can  find  no  other  literature  records  relating  to  repro¬ 
duction  in  L.  mordax. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


333 


Lonchophylla  concava 

Although  there  are  few  reports  giving  reproductive  condition  (Table  5),  these 
are  from  sufficiently  distinct  times  of  the  year  as  to  suggest  the  possibility  of  more 
than  one  birth  peak  per  year. 

Lonchophylla  robusta 

Apparently,  reproductively  active  individuals  of  this  species  have  not  been 
recorded.  The  capture  of  inactive  females  in  several  months  of  the  year  (Table  5) 
suggests  an  asynchronous  reproductive  cycle. 

Lonchophylla  thomasi 

No  data  are  available  concerning  the  reproductive  pattern  of  this  species. 

Lionycteris  spurrelli 

The  only  published  record  of  reproductive  activity  in  L.  spurrelli  is  that  of 
Tuttle  (1970),  who  reported  a  pregnant  female  taken  in  August  in  Peru. 

Anoura  geoffroyi 

Alvarez  and  Ramirez-Pulido  (1972)  netted  12  males  but  no  females  at  the 
mouth  of  a  cave  in  Michoacan  and  suggested  that  this  species  may  form  sexually 
segregated  colonies.  The  data  of  Goodwin  and  Greenhall  (1961)  from  Trinidad 
support  this  notion  for  certain  times  of  the  year.  They  reported  20  males  and  25 
females  in  June;  29  males  and  one  female  in  October;  and  32  males  and  56  fe¬ 
males  in  November — all  from  the  same  caves.  Their  data  also  recommend  a 
discrete  breeding  season  occurring  late  in  the  rainy  season,  a  rather  unusual  pat¬ 
tern  for  phyllostomatids.  See  Table  5. 

Anoura  caudifer 

Pregnancy  records  (Table  5)  from  several  months  throughout  the  year  suggest 
an  asynchronous  reproductive  cycle.  Additional  data  from  other  months  of  the 
year  would  be  useful  in  discerning  the  true  reproductive  patterns. 

Anoura  cultrata 

Gardner  et  al.  (1970)  reported  a  pregnant  female  from  Costa  Rica  taken  in 
August.  They  also  collected  males  in  February,  May,  and  July  and  gave  testicular 
measurements. 

Anoura  werckleae 

Nothing  is  known  about  the  reproductive  pattern  of  this  species. 

Anoura  brevirostrum 

When  Carter  (1968)  described  A.  brevirostrum  from  Peru,  he  included  data 
from  a  lactating  female  and  two  inactive  females  taken  in  August. 


334 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  5. — Reproductive  data  for  the  genera  Monophyllus,  Leptonycteris,  Lonchophylla, 
Anoura,  Lichonycteris,  Hylonycteris,  Choeroniscus,  and  Choeronycteris. 


Place 

Date 

Pregnant  Lactating  Inactive 

Reference 

Jamaica 

Feb 

Monophyllus  redmani 

11 

Osburn,  1865 

Feb 

6  4 

McNab,  1976 

Jan 

2 

Goodwin,  1970 

Dec 

15  4 

n 

Caicos  Is. 

Jan 

1 

Homan  and  Jones,  1975 

Hispaniola 

Feb 

1 

II 

Dec 

2 

II 

Puerto  Rico 

Feb 

1 

It 

Texas 

Jun 

Leptonycteris  nivalis 

3 

Easterla,  1972 

Coahuila 

Jul 

20 

Baker,  1956 

Tamaulipas 

Aug 

12 

Alvarez,  1963 

Veracruz 

Sep 

6 

Hall  and  Dalquest,  1963 

Arizona 

Aug 

Leptonycteris  sanborni 

X 

Hoffmeister  and  Goodpaster,  1954 

Sonora 

Mar 

1 

Cockrum  and  Bradshaw,  1963 

Apr 

11 

II 

Sinaloa 

Feb 

1 

Jones  et  ai,  1972 

Jul 

1 

II 

Nov 

1 

II 

Jalisco 

Jan 

1 

Watkins  et  al.,  1972 

Jul 

1 

II 

Oct 

12 

II 

Morelos 

Sep 

1  2 

Villa-R.,  1966 

Mexico 

Nov 

1  2 

II 

Costa  Rica 

Mar 

Lonchophylla  concava 

1  1 

Davis  et  al.,  1964 

Aug 

1 

Gardner  et  al.,  1970 

Costa  Rica 

Mar 

Lonchophylla  robusta 

4 

Mares  and  Wilson,  1971 

Colombia 

Mar 

1 

Thomas,  1972 

Apr 

2 

II 

Jul 

1 

II 

Sep 

2 

II 

Peru 

Aug 

1 

Tuttle,  1970 

Zacatecas 

Jun 

Anoura  geoffroyi 

2 

Matson  and  Patten,  1975 

Sinaloa 

Jul 

6 

Jones  et  al.,  1972 

Colima 

Nov 

5 

Villa-R.,  1966 

Dec 

X 

II 

Guerrero 

Sep 

3 

II 

Oaxaca 

Jul 

3 

Baker  and  Womochel,  1966 

Nov 

X 

Schaldach,  1966 

Dec 

X 

II 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


335 


Nicaragua 
Costa  Rica 
Trinidad 
Peru 

Colombia 


French  Guiana 
Brazil 


Guatemala 
Costa  Rica 


Jalisco 

Tabasco 

Oaxaca 

Guatemala 
Costa  Rica 


Sinaloa 

Oaxaca 

Honduras 

Nicaragua 

Costa  Rica 

Trinidad 

Peru 


Table  5. —  Continued. 

Jul 

1 

1 

Jones  et  aL,  1971  a 

Mar 

2 

1 

Mares  and  Wilson,  1971 

Nov 

56 

Goodwin  and  Greenhall,  1961 

Jun 

2 

14 

Tuttle,  1970 

Anoura  caudifer 

Mar 

1 

Thomas,  1972 

May 

1 

rr 

Nov 

1 

tt 

Jun 

1 

Tamsitt  and  Valdivieso,  1966a 

Jan 

X 

Brosset  and  Dubost,  1967 

Feb 

X 

tt 

Feb 

1 

Kuhlhorn,  1953 

Jan 

2 

Lichonycteris  obscura 

Peracchi  and  Albuquerque,  1971 

Feb 

2 

1 

Carter  et  al„  1966 

Jan 

1 

Gardner  et  aL,  1970 

Mar 

1 

tt 

Hylonycteris  underwood  i 

Jul 

2 

Phillips  and  Jones,  1971 

Sep 

2 

2 

It 

May 

1 

Villa-R.,  1966 

Nov 

1  1 

ft 

Jul 

1 

Baker  and  Womochel,  1966 

Mar 

1 

Carter  et  al.,  1966 

Jan 

1 

LaVal,  1977 

Feb 

1 

tt 

Mar 

1 

tt 

Apr 

1 

tt 

Apr 

X 

Gardner  et  aL,  1970 

May 

X 

ft 

Jun 

X 

ft 

Jul 

X 

ft 

Aug 

1 

LaVal,  1972 

Oct 

1 

tt 

Nov 

1 

Choeroniscus  godmani 

ft 

Jul 

1 

Jones,  19646 

May 

1 

Schaldach,  1965 

Jul 

1 

Valdez  and  LaVal,  1971 

Mar 

1 

Jones  et  al.,  1971a 

Apr 

1 

tt 

Mar 

1 

Mares  and  Wilson,  1971 

Choeroniscus  intermedius 

Aug 

1 

Goodwin  and  Greenhall,  1961 

Jul 

2 

Tuttle,  1970 

336 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  5. — Continued. 


Choeronycteris  mexicana 


Arizona 

Jun 

1 

Campbell,  1934 

Jun 

X 

Barbour  and  Davis,  1969 

Jun 

X 

Walker,  1975 

Jul 

X 

n 

Aug 

35 

Hoffmeister  and  Goodpaster,  1954 

New  Mexico 

Jun 

2 

4 

Mumford  and  Zimmerman,  1962 

Jun 

4 

Mumford  et  al.,  1964 

Coahuila 

Mar 

1 

Baker,  1956 

Jun 

4 

tl 

Aug 

X 

It 

Sep 

X 

It 

Jun 

1 

Axtell,  1962 

Tamaulipas 

Aug 

X 

Alvarez,  1963 

Sonora 

Jul 

4 

Villa-R.,  1966 

Sinaloa 

Feb 

1 

Jones  et  al.,  1972 

Jalisco 

Jan 

1 

Watkins  et  al.,  1972 

Feb 

1 

It 

Mar 

3 

It 

Sep 

1 

It 

Oct 

1 

It 

Guerrero 

Feb 

2 

Villa-R.,  1966 

Scleronycteris  ega 

Nothing  has  been  recorded  about  reproduction  in  this  species. 

Lichonycteris  degener 

Nothing  is  known  about  the  reproductive  pattern  of  L.  degener. 

Lichonycteris  obscura 

This  species  is  reproductively  active  during  the  dry  season  in  Middle  America 
(Table  5),  but  until  data  are  available  from  other  months  of  the  year,  little  can 
be  said  of  the  overall  pattern. 

Hylonycteris  underwoodi 

The  data  from  Costa  Rica  (Table  5)  fit  the  bimodal  pattern  common  to  many 
other  species.  The  second  birth  peak  appears  to  be  later  in  the  rainy  season  than 
for  some  other  species. 

Platalina  genovensium 

Nothing  is  known  about  reproduction  in  this  species. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


337 


Choeroniscus  godmani 

Choeroniscus  godmani  (Table  5)  seems  to  fit  the  usual  pattern  of  weaning 
young  during  the  early  part  of  the  rainy  season,  but  the  lack  of  data  from  later 
in  the  year  makes  this  conclusion  tentative. 

Choeroniscus  minor 

The  only  apparent  report  of  reproductive  activity  for  this  species  is  that  of 
Tamsitt  et  al.  (1965),  who  reported  a  lactating  female  from  Colombia  in  Decem¬ 
ber.  Tuttle  (1970)  collected  a  juvenile  in  August  in  Pern. 

Choeroniscus  intermedius 

The  data  in  Table  5  are  too  few  to  provide  much  insight  into  the  reproductive 
pattern  of  this  species. 

Choeroniscus  inca 

Goodwin  and  Greenhall  (1961)  noted  a  pregnant  female  taken  in  February  in 
Trinidad. 

Choeroniscus  periosus 

The  only  record  is  that  of  Thomas  (1972),  who  captured  two  lactating  females 
in  Colombia  in  January. 

Choeronycteris  mexicana 

These  animals  are  pregnant  in  the  early  spring  in  Mexico  (Table  5),  and  those 
that  migrate  to  Arizona  and  New  Mexico  give  birth  in  June.  The  possibility  of  a 
second  period  of  parturition,  as  suggested  for  Leptonycteris  sanborni,  is  supported 
by  the  pregnancy  record  in  September  from  Jalisco. 

Musonycteris  harrisoni 

There  are  no  published  records  of  reproductive  activity  for  this  species,  but 
Alfred  L.  Gardner  has  kindly  made  available  to  me  his  unpublished  field  notes, 
which  record  one  inactive  and  two  pregnant  females  taken  in  September  in 
Colima. 

Carollia  castanea 

Pine  (1972)  suggested  that  C.  castanea  is  polyestrous,  but  cautioned  that  in  any 
one  locality  there  may  be  one  or  two  more  or  less  fixed  seasons.  This  caveat  is 
supported  by  Fleming  (1973),  who  suggested  that  in  Panama  C.  castanea  is 
bimodally  polyestrous,  with  birth  peaks  occuring  in  March-April  and  July-August. 
Thomas’  (1972)  data  show  that  females  are  pregnant  during  the  period  September- 
November  in  Colombia  corresponding  to  a  period  of  reproductive  quiescence  in 


338 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Panama  (Fleming,  1973).  These  differences  probably  reflect  contrasts  in  the  sea¬ 
sonality  of  the  rainfall  patterns  at  the  different  localities.  See  Table  6. 

Carollia  subrufa 

Felten  (1956a)  suggested  that  C.  subrufa  breeds  both  in  the  dry  and  wet 
seasons  in  El  Salvador.  Pine  (1972)  felt  that  they  either  breed  throughout  the 
year  or  that  possibly  there  is  a  period  of  inactivity  in  the  early  winter  months,  at 
least  in  some  areas.  The  data  from  El  Salvador,  the  most  extensive  for  any  one 
area,  would  seem  to  fit  a  bimodal  pattern  (Table  6). 

Carollia  brevicauda 

Pine  (1972)  suggested  that  C.  brevicauda  breeds  from  midwinter  to  early 
spring.  Three  records  of  females  both  pregnant  and  lactating  (Table  6)  attest  to 
the  presence  of  polyestry  in  Central  America.  This  species  may  exhibit  the  bi¬ 
modal  type  of  breeding  season  seen  for  other  Central  American  phyllostomatids; 
however,  data  from  late  in  the  year  are  needed  for  clarification  of  this  pattern. 

Carollia  perspicillata 

Fleming  (1973)  and  Heithaus  et  al.  (1975)  have  shown  that  C.  perspicillata 
fits  the  model  of  bimodal  polyestry,  and  the  data  summarized  here  support  this 
contention  (Table  6).  Birth  peaks  occur  in  the  periods  February-May  and  June- 
August  in  Panama,  and  somewhat  earlier  in  other  areas,  depending  on  seasonal 
rainfall  patterns  in  the  various  localities.  Several  of  the  data  sets  from  various 
localities  show  a  distinct  drop  in  reproductive  activity  during  the  latter  part  of 
the  rainy  season,  usually  in  the  period  from  October  to  December,  but  earlier  in 
Colombia.  Fleming  et  al.  (1972)  correlated  testis  size  with  spermatogenic 
activity  and  found  that  males  had  large  testes  just  preceding  those  times  when 
females  were  likely  to  be  sexually  active. 

Rhinophylla  pumilio 

The  data  in  Table  6  are  too  few  to  warrant  speculation  on  the  reproductive 
pattern  of  R.  pumilio. 

Rhinophylla  alethina 

Although  the  sample  (Table  6)  is  admittedly  small,  the  timing  of  the  repro¬ 
ductive  events  recorded  here  suggests  an  extended  or  possibly  asynchronous 
breeding  season.  Data  from  August-November  would  be  useful  for  clarifying  the 
pattern. 

Rhinophylla  fischerae 

The  lack  of  reproductive  activity  for  animals  taken  in  July  and  August  seems 
striking  when  compared  against  what  is  known  for  other  phyllostomatids.  See 
Table  6. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


339 


Table  6. —  Reproductive  data  for  the  genera  Carollia  and  Rhinophylla. 


Place 

Date 

Pregnant 

Lactating  Inactive 

Reference 

Carollia  castanea 

Honduras 

May 

3 

Pine,  1972 

Jul 

1 

n 

Nicaragua 

Feb 

X 

Jones  et  al.,  1971a 

Mar 

X 

X 

n 

Apr 

X 

n 

Jun 

X 

n 

Jul 

X 

X 

tt 

Aug 

X 

n 

Costa  Rica 

Feb 

5 

n 

Mar* 

1 

n 

Aug 

1 

ii 

Panama 

Jan 

5 

n 

Feb 

5 

n 

Mar 

5 

n 

Jan 

1 

2 

Fleming  et  al.,  1972 

Mar 

3 

3 

II 

Apr 

1 

II 

Jun 

1 

II 

Jul 

1 

II 

Aug 

4 

1 

II 

Oct 

1 

II 

Nov 

3 

II 

Dec 

2 

II 

Colombia 

Jan 

5 

2  5 

Thomas,  1972 

Feb 

1 

II 

Mar 

1 

2 

II 

Apr 

2 

1  7 

it 

May 

2 

II 

Jun 

2 

II 

Jul 

1 

II 

Sep 

1 

3 

II 

Oct 

2 

3 

II 

Nov 

3 

1 

II 

Dec 

2 

II 

French  Guiana 

Jan 

X 

Brosset  and  Dubost,  1967 

Feb 

X 

II 

Mar 

X 

II 

Peru 

Jul 

2 

Tuttle,  1970 

Aug 

1 

II 

Carollia  subrufa 

Puebla 

Jun 

3 

LaVal,  1972 

Guerrero 

May 

3 

Pine,  1972 

Dec 

1 

II 

Oaxaca 

May 

2 

3 

Villa-R.,  1966 

Chiapas 

Feb 

6 

ll 

May 

1 

Pine,  1972 

Jul 

2 

ll 

340 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  6. —  Continued. 


Chiapas 

Aug 

5 

1 

Oct 

1 

1 

Nov 

1 

Guatemala 

Feb 

6 

1 

Nov 

2 

El  Salvador 

Jan 

43 

7 

Feb 

1 

1 

Mar 

4 

1 

Sep 

5 

Oct 

3 

21 

Nov 

12 

Dec 

2 

Honduras 

Jul 

1 

Nicaragua 

Jul 

1 

Aug 

1 

1 

Panama 

Jan 

X 

Feb 

X 

Mar 

X 

Dec 

X 

Carollia 

brevicauda 

San  L.  Potosi 

Apr 

2 

Aug 

1 

Veracruz 

Feb 

2 

Mar 

19 

1 

Dec 

1 

Tabasco 

Apr 

2 

May 

1 

Campeche 

Jan 

4 

Quintana  Roo 

Apr 

1 

Aug 

1 

Oaxaca 

Feb 

1 

Mar 

2 

1 

Chiapas 

Jun 

1 

2 

3 

Jul 

6 

1 

2 

Nov 

2 

Guatemala 

Feb 

4 

4 

Mar 

1 

Aug 

1 

2 

Feb 

1 

Honduras 

Apr 

3 

2 

May* 

3 

1 

1 

Jun 

2 

4 

Nicaragua 

Jul* 

1 

Costa  Rica 

Mar 

1 

1 

1 

Apr* 

2 

Panama 

Jan 

1 

Feb 

10 

1 

Mar 

9 

Ecuador 

Mar 

1 

Peru 

Aug 

3 

Oct 

2 

Pine,  1972 

ft 

tt 

tt 

tt 

Felten,  1956c 

tt 

ft 

tt 

tt 

rr 

tt 

Pine,  1972 

ft 

tt 

Walker,  1975 

ft 

tt 

tt 

Pine,  1972 

tt 

tt 

ft 

tt 

tt 

tt 

ft 

If 

ft 

ft 

tt 

tt 

ft 

ft 

ft 

ft 

Jones,  1966 
Rick,  1968 
Pine,  1972 

ft 

ft 

tt 

tt 

ft 

tt 

tt 

tt 

ft 

tt 

tt 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


341 


Table  6. —  Continued. 


Carollia  perspicillata 


Puebla 

Jan 

4 

LaVal,  1972 

Veracruz 

May 

3 

Villa-R.,  1966 

Jun 

10 

1 

Lackey,  1970 

Jul 

1 

1 

n 

Campeche 

May 

2 

Jones  et  al.,  1973 

Jul 

1 

Pine,  1972 

Quintana  Roo 

Jul 

4 

II 

Jul 

5 

Jones  et  al.,  1973 

Aug 

1 

II 

Oaxaca 

Apr 

X 

Hahn,  1907 

Chiapas 

Aug 

1 

Pine,  1972 

Guatemala 

Mar 

1 

n 

Mar 

3 

1 

Jones,  1966 

El  Salvador 

Mar 

1 

Burt  and  Stirton,  1961 

Apr 

1 

2 

Felten,  1956c 

Oct 

4 

It 

Nov 

17 

n 

Dec 

4 

it 

Mar 

1 

Pine,  1972 

Honduras 

May 

2 

6 

3 

n 

Jun 

1 

n 

Jul 

2 

1 

it 

Nicaragua 

Feb 

8 

3 

ti 

Apr 

1 

n 

May 

1 

3 

it 

Jun 

2 

1 

it 

Jul 

2 

n 

Aug 

3 

n 

Costa  Rica 

Feb 

1 

5 

n 

Mar 

6 

1 

1 

n 

Apr 

2 

1 

2 

n 

Jul 

4 

1 

n 

Aug 

1 

1 

n 

Jan 

1 

3 

Fleming  et  al.,  1972 

Feb 

5 

5 

It 

Mar 

17 

3 

It 

Apr 

1 

10 

3 

It 

May 

12 

7 

II 

Jun 

3 

II 

Jul 

4 

7 

II 

Aug 

1 

4 

6 

II 

Sep 

1 

II 

Oct 

2 

II 

Nov 

3 

II 

Panama 

Jan 

1 

14 

It 

Feb 

15 

4 

II 

Mar 

28 

1 

8 

II 

Apr 

10 

16 

13 

II 

May 

5 

6 

4 

II 

Jun* 

6 

1 

II 

Jul 

10 

3 

4 

II 

342 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  6. — Continued. 


Panama 


Trinidad 


Venezuela 
French  Guiana 


Colombia 


Ecuador 


Aug 

20 

10 

15 

Sep 

2 

6 

18 

Oct 

4 

27 

Nov 

9 

Dec 

1 

19 

Mar 

1 

May 

2 

Feb 

10 

1 

Mar 

1 

2 

Apr 

1 

3 

4 

Jun 

2 

Jun 

X 

Feb 

X 

Mar 

X 

Apr 

X 

May 

X 

X 

Jun 

X 

X 

Jul 

X 

X 

Aug 

X 

X 

Sep 

X 

X 

Oct 

X 

X 

Jun 

7 

1 

8 

Jul 

X 

Aug 

X 

Sep 

X 

Oct 

X 

Nov 

X 

Jul 

1 

Aug 

34 

16 

50 

Sep 

20 

Jan 

1 

1 

Mar 

1 

3 

Apr 

1 

2 

Oct 

1 

1 

Jan 

9 

3 

4 

Feb 

1 

1 

Mar 

2 

2 

5 

Apr 

1 

1 

4 

May* 

3 

2 

2 

Jun 

3 

4 

8 

Jul 

2 

5 

Aug 

2 

4 

Sep 

1 

5 

Oct 

2 

5 

Nov 

1 

1 

6 

Dec 

4 

9 

Jun 

2 

Jul 

4 

3 

Mar 

2 

Jul 

1 

Aug 

1 

Sep 

1 

1 

Fleming  et  al.,  1972 

ft 

ft 

ft 

ft 

Enders,  1935 

Hall  and  Jackson,  1953 

Pine,  1972 

ft 

ft 

ft 

Hahn,  1907 

Goodwin  and  Greenhall,  1961 

ft 

ft 

ft 

ft 

ft 

ft 

ft 

ft 

Pirlot,  1963 

Brosset  and  Dubost,  1967 

ft 

ft 

ft 

rr 

Arata  and  Vaughn,  1970 

ft 

ft 

Tamsitt  and  Valdivieso,  1964 

ft 

ft 

ft 

Thomas,  1972 

ft 

rr 

tt 

rr 

rr 

rr 

rr 

rr 

rr 

rr 

rr 

Pine,  1972 

rr 

rr 

rr 

rr 

rr 


Bolivia 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


343 


Table  6.— Continued. 


Peru 

Aug 

2 

11 

Pine,  1972 

Jun 

12 

Tuttle,  1970 

Jul 

7 

tt 

Aug 

3 

12 

ft 

Brazil 

Jan 

X 

Peracchi  and  Albuquerque,  1971 

Sep 

X 

X 

tt 

Oct 

X 

Rhinophylla  pumilio 

ft 

Venezuela 

Dec 

1 

1  1 

Walker,  1975 

Peru 

Jun 

4 

Tuttle,  1970 

Jul 

5 

tt 

Colombia 

Jan 

1 

Thomas,  1972 

Apr 

1 

tt 

May 

1 

ft 

Jun 

1  2 

tt 

Jul 

1 

ft 

Dec 

1 

tt 

Rhinophylla  fischerae 

Peru 

Jul 

8 

Tuttle,  1970 

Aug 

1 

ft 

Aug 

7 

Carter,  1966 

*Pregnant  and  lactating. 


Sturnira  lilium 

Jones  (1966)  and  Jones  et  al.  (1973)  suggested  that  S.  lilium  probably  breeds 
through  the  year.  Actually,  the  data  presented  in  Table  7  support  the  model  of 
bimodal  polyestry  as  suggested  by  Fleming  et  al.  (1972).  Support  for  this  model 
is  provided  by  the  data  from  Costa  Rica  and  Colombia.  In  Costa  Rica,  birth  peaks 
occur  in  February-March  and  in  June-July  (Heithaus  et  al.,  1975);  in  Colombia, 
there  appears  to  be  much  less  synchrony  in  the  cycle. 

Sturnira  thomasi 

Genoways  and  Jones  (1975)  reported  two  lactating  females,  a  subadult,  and 
a  juvenile  from  Guadeloupe  in  July.  This  seems  to  be  the  only  record  of  repro¬ 
duction  available  for  this  species. 

Sturnira  tildae 

The  records  listed  in  Table  7  provide  no  basis  for  speculation  on  the  repro¬ 
ductive  habits  of  S.  tildae. 

Sturnira  magna 

Tuttle  (1970)  provided  testicular  measurements  on  three  Peruvian  males 
taken  in  July.  Gardner  (1976)  took  one  inactive  and  one  lactating  female  in 
May  and  another  inactive  female  in  July  in  Peru. 


344 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  7. — Reproductive  data  for  the  gcmrsSturnira. 


Place 

Date 

Pregnant 

Lactating 

Inactive 

Reference 

Sturnira  lilium 

Sonora 

Sep 

1 

Findley  and  Jones,  1965 

Sinaloa 

Apr 

1 

Cockrum  and  Bradshaw,  1963 

May 

1 

Jones  et  al.,  1972 

Jun 

1 

2 

II 

Aug 

2 

n 

Durango 

Jun 

3 

1 

Jones,  1964  c 

Jul 

1 

1 

Baker  and  Greer,  1962 

Jalisco 

Jan 

X 

Watkins  et  al.,  1972 

Mar 

X 

II 

Apr 

X 

X 

II 

Jun 

X 

X 

it 

Jul 

X 

X 

n 

Aug 

X 

it 

Sep 

X 

X 

ti 

Oct 

X 

it 

Nov 

2 

it 

Queretaro 

Jan 

1 

Spenrath  and  LaVal,  1970 

Puebla 

Jan 

4 

LaVal,  1972 

Veracruz 

Jun 

21 

Lackey,  1970 

Jul 

4 

1 

It 

Campeche 

Jan 

5 

Jones  et  al.,  1973 

Jul 

1 

It 

Quintana  Roo 

Aug 

1 

It 

Apr 

2 

Birney  et  al.,  1974 

Oaxaca 

Apr 

1 

1 

USNM 

Jul 

6 

Baker  and  Womochel,  1966 

Dec 

X 

Schaldach,  1966 

Chiapas 

May 

4 

Villa-R.,  1966 

Jun 

1 

It 

Guatemala 

Feb 

X 

X 

Jones,  1966 

Mar 

X 

X 

It 

Jun 

X 

X 

It 

Jul 

X 

X 

II 

Aug 

X 

X 

II 

May 

1* 

2 

Rick,  1968 

El  Salvador 

Jul 

1 

Starrett  and  de  la  Torre,  1964 

Nicaragua 

Jul 

1 

1 

II 

Costa  Rica 

Jan 

3 

1 

Fleming  et  aL,  1972 

Feb 

1 

5 

2 

II 

Mar 

7 

5 

II 

Apr 

5 

5 

II 

May 

4 

1 

3 

II 

Jun 

1 

2 

II 

Jul 

3 

2 

II 

Aug 

2 

6 

II 

Dec 

3 

II 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


345 


Dominica 


Martinique 
St.  Lucia 
St.  Vincent 
Colombia 


French  Guiana 


Peru 

Brazil 


Trinidad 

Peru 

Brazil 


Costa  Rica 


Jalisco 


Colima 

Queretaro 

Puebla 


Table  7. — Continued. 


Mar 

7 

Apr 

4 

Aug 

1  4 

Mar 

11 

2 

Aug 

2 

1 

Aug 

1 

Jul 

9 

3  4 

Aug 

1 

1  13 

Jan 

3 

3 

Feb 

3 

Mar 

3 

2 

Apr 

1 

1  2 

May 

2  3 

Jun 

1  3 

Jul 

2  2 

Aug 

2  1 

Sep 

4  4 

Oct 

4 

3  2 

Nov 

7 

Dec 

4 

1  3 

Jun 

X 

Jul 

X 

Aug 

X 

Jun 

8 

Jul 

1 

8 

Jul 

1 

Aug 

X 

Sturnira  tildae 

Mar 

1 

Jul 

2 

Jun 

1 

Jul 

1 

1 

Sturnira  mordax 

Feb 

2 

1 

May 

1 

May 

1 

Aug 

1 

Sturnira  ludovici 

Apr 

7 

5 

May 

2 

Jul 

1 

1 

Aug 

1  13 

Sep 

1 

Nov 

5 

Dec 

1 

Nov 

5 

5 

Sep 

12 

Nov 

2 

Jan 

11 

Jan 

1 

Jones  and  Phillips,  1976 

II 

n 

ti 

ti 

n 

Arata  and  Vaughn,  1970 

II 

Thomas,  1972 

II 

II 

II 

II 

II 

II 

II 

II 

II 

II 

II 

Brosset  and  Dubost,  1967 

II 

II 

Tuttle,  1970 

II 

USNM 

Peracchi  and  Albuquerque,  1971 

Goodwin  and  Greenhall,  1961 

Tuttle,  1970 

USNM 

II 

Gardner  et  al.,  1970 

II 

LaVal,  1977 
Armstrong,  1969 

Watkins  et  al.,  1972 

II 

II 

II 

II 

II 

II 

Jones  and  Phillips,  1964 
Villa-R.,  1966 

II 

Schmidly  and  Martin,  1973 
LaVal,  1972 


346 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  7. — Continued. 


Oaxaca 

Jul 

4 

Baker  and  Womochel,  1966 

Chiapas 

Aug 

1 

Villa-R.,  1966 

Costa  Rica 

Jul 

1 

Starrett  and  de  la  Torre,  1964 

Colombia 

Jan 

2 

Thomas,  1972 

Feb 

1 

It 

Mar 

4 

1 

ft 

Apr 

4 

1 

ft 

May* 

12 

9  6 

ft 

Jun 

1 

ft 

Jul 

1  1 

ft 

Aug 

7 

1 

ft 

Sep 

1 

ft 

Oct 

4 

2 

ft 

Nov 

2 

tt 

Dec 

4 

5 

tt 

Peru 

Jun 

1 

Tuttle,  1970 

Sturnira  erythromos 

Colombia 

May 

2 

Thomas,  1972 

Dec 

2 

it 

Peru 

Jun 

1 

Gardner  and  O’Neill,  1969 

Aug 

10 

5 

tt 

*Pregnant  and  lactating. 


Sturnira  mordax 

The  presence  of  pregnant  females  both  in  February  and  in  August  suggests 
polyestry  for  S.  mordax  (Table  7). 

Sturnira  bidens 

The  only  published  record  of  reproductive  activity  for  this  species  is  that  of 
Gardner  and  O’Neill  (1969),  who  reported  three  pregnant  females  and  one  in¬ 
active  female  from  Peru  in  August. 

Sturnira  nana 

Nothing  is  known  about  the  reproductive  pattern  of  S.  nana. 

Sturnira  aratathomasi 

Thomas  and  McMurray  (1974)  reported  pregnant  females  from  Colombia  in 
February  and  August.  These  pregnancy  dates  are  not  inconsistent  with  those  for 
other,  more  common  species  of  the  genus  Sturnira  and  may  represent  the  familiar 
bimodal  pattern. 

Sturnira  ludovici 

Sturnira  ludovici  appears  to  me  to  be  another  species  with  a  bimodal  polyes- 
trous  pattern  (Table  7).  The  data  from  Colombia  are  strikingly  similar  to  those 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


347 


presented  for  S.  lilium.  Starrett  and  de  la  Torre  (1964)  presented  data  on  testis 
size  and  spermatogenesis  from  two  males  from  Costa  Rica. 

Sturnira  erythromos 

Speculation  on  the  reproductive  pattern  of  S.  erythromos  must  await  further 
data.  See  Table  7. 

Uroderma  bilobatum 

Davis  (1968)  suggested  that  U.  bilobatum  seemingly  lacks  a  restricted  breed¬ 
ing  season  based  on  his  examination  of  58  females  from  a  variety  of  localities 
from  Oaxaca  to  Venezuela.  Of  these,  three  were  pregnant  in  January,  five  in 
February,  and  one  each  in  May,  July,  and  November  (Table  8).  Fleming  (1973) 
pointed  out  that  in  Panama  this  species  is  another  example  of  bimodal  polyestry 
and  much  of  the  above  data  are  in  agreement  with  that  conclusion.  Again,  the  in¬ 
formation  from  Colombia  shows  that  the  timing  of  reproductive  peaks  is  quite 
different  from  that  in  Panama,  with  the  second  major  pregnancy  period  in  Colom¬ 
bia  occurring  in  the  late  rainy  season.  Fleming  et  al.  (1972)  presented  data  on 
testis  size  and  spermatogenesis,  showing  that  males  undergo  active  spermato¬ 
genesis  in  a  bimodal  fashion  also. 

Uroderma  magnirostrum 

Although  the  data  are  few  and  from  widely  scattered  localities  (Table  8),  I 
suspect  U.  magnirostrum  will  prove  to  have  a  polyestrous  pattern  like  that  of  its 
congener,  U.  bilobatum. 

Vampyrops  infuscus 

The  only  report  of  reproduction  in  this  species  appears  to  be  that  of  Marinkelle 
(1970),  who  collected  one  pregnant  female  and  three  lactating  females  in 
Colombia  in  March. 

Vampyrops  vittatus 

Pregnancies  occur  in  the  early  part  of  the  rainy  season  in  Costa  Rica  (Table  8), 
but  data  from  other  seasons  are  lacking. 

Vampyrops  dorsalis 

The  Colombian  data  (Table  8)  show  V.  dorsalis  to  fit  the  pattern  of  bimodal 
polyestry  common  to  several  other  species  of  Colombian  phyllostomatids. 

Vampyrops  aurarius 

No  data  are  available  about  reproduction  in  this  species. 

Vampyrops  nigellus 

Nothing  is  known  about  the  reproductive  pattern  of  V.  nigellus. 


348 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  8. —  Reproductive  data  for  the  genera  Uroderma,  Vampyrops,  and  Vampyrodes. 


Place 

Date 

Pregnant 

Lactating 

Inactive 

Reference 

Uroderma  bilobatum 

Veracruz 

Jun 

1 

Lackey,  1970 

Jul 

1 

it 

Chiapas 

May 

X 

Villa-R.,  1966 

Aug 

1 

ft 

Guatemala 

Feb 

1 

Jones,  1966 

El  Salvador 

Jan 

4 

Felten,  1956a 

Jan 

3 

Burt  and  Stirton,  1961 

May 

1 

tt 

Honduras 

Jul 

72 

1 

12 

Baker  et  aL,  1975 

Nicaragua 

Feb 

4 

Jones,  1964  a 

Aug 

2 

Davis  et  aL,  1964 

Panama 

Jan 

8 

Davis,  1968 

Mar 

X 

Bloedel,  1955 

Jan 

16 

1 

Fleming  et  al.,  1972 

Feb 

7 

tt 

Mar 

1 

11 

3 

tt 

Feb 

X 

X 

Walker,  1975 

Mar 

X 

X 

tt 

Apr 

X 

X 

tt 

Apr 

10 

15 

Fleming  et  al.,  1972 

May 

12 

3 

2 

tt 

Jun 

4 

1 

ft 

Jul 

4 

9 

6 

tt 

Aug 

1 

2 

tt 

Sep 

24 

ft 

Oct 

10 

tt 

Nov 

1 

tt 

Dec 

2 

tt 

Trinidad 

Feb 

1 

1 

Goodwin  and  Greenhall,  1961 

May 

3 

3 

2 

tt 

Colombia 

Jan 

1 

Tamsitt  and  Valdivieso,  1964 

Mar 

1 

1 

ft 

Jul 

3 

ft 

Sep 

1 

tt 

Nov 

1 

1 

tt 

Nov 

1 

Thomas,  1972 

Peru 

Aug 

1 

2 

Tuttle,  1970 

Brazil 

Jul 

3 

USNM 

Uroderma  magnirostrum 

El  Salvador 

Jun 

1 

Davis,  1968 

Nicaragua 

Mar 

1 

Jones  et  al.,  1971a 

Jul 

1 

Davis,  1968 

Bolivia 

Sep 

10 

7 

It 

Brazil 

Jun 

1 

USNM 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


349 


Table  8. — Continued. 


Vampyrops  vittatus 

Costa  Rica 

Mar 

X 

LaVal,  1977 

Apr 

2 

Davis  et  al.,  1964 

Jan 

1 

Gardner  et  aL,  1970 

May 

4 

tt 

Jun 

1 

1 

1 

tt 

Jul 

1 

tt 

Jul 

1 

Tamsitt  and  Valdivieso,  1961 

Colombia 

May 

1 

3 

Thomas,  1972 

Oct 

1 

It 

Dec 

1 

tt 

Peru 

Jun 

4 

Tuttle,  1970 

Aug 

9 

tt 

Vampyrops  dorsalis 

Colombia 

Aug 

3 

18 

Arata  and  Vaughn,  1970 

Jan 

8 

8 

Thomas,  1972 

Feb 

2 

5 

tt 

Mar* 

12 

3 

ft 

Apr 

4 

2 

tt 

May 

5 

2 

10 

tt 

Jun 

2 

5 

8 

ft 

Jul* 

2 

2 

8 

tt 

Aug 

9 

tt 

Sep 

7 

tt 

Oct 

1 

4 

It 

Nov 

3 

ft 

Dec 

7 

2 

ft 

Vampyrops  brachycephalis 


Venezuela 

Feb 

1 

Rouk  and  Carter,  1972 

Jul 

1 

tt 

Oct 

1  3 

tt 

Colombia 

Jul 

1 

tt 

Peru 

Aug 

2 

5 

tt 

Vampyrops  helleri 

Tabasco 

May 

1 

Villa-R.,  1966 

Chiapas 

May 

1 

ft 

Jul 

1 

Davis  et  al.,  1964 

Guatemala 

May 

1 

Rick,  1968 

El  Salvador 

Jun 

2 

LaVal,  1969 

Honduras 

Aug 

1 

ft 

Nicaragua 

Mar 

X 

Jones  et  al.,  1971  a 

Apr 

X 

tt 

Jun 

X 

It 

Jul 

X 

ft 

Aug 

X 

tt 

Costa  Rica 

Mar 

1 

Mares  and  Wilson,  1971 

Aug 

1 

Starrett  and  de  la  Torre,  1964 

Panama 

Jan 

1 

Fleming  et  al.,  1972 

350 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  8. — Continued. 


Panama 

Apr 

1 

1 

Fleming  et  al.,  1972 

Jul 

1 

ft 

Sep 

1 

tt 

Oct 

1 

tt 

Nov 

1 

tt 

Dec 

1 

tt 

Colombia 

Aug 

2 

2 

9 

Arata  and  Vaughn,  1970 

Jan* 

3 

13 

6 

Thomas,  1972 

Feb 

2 

7 

tt 

Mar 

7 

3 

ft 

Apr* 

3 

5 

8 

tt 

May 

1 

9 

5 

tt 

Jun* 

2 

7 

tt 

Jul 

6 

tt 

Aug 

6 

tt 

Sep 

14 

tt 

Oct 

7 

3 

tt 

Nov 

5 

1 

tt 

Dec 

5 

2 

tt 

French  Guiana 

Aug 

X 

ft 

Sep 

X 

tt 

Peru 

Jul 

1 

Tuttle,  1970 

Aug 

2 

2 

tt 

Vampyrodes  caraccioli 

Veracruz 

Apr 

1 

Villa-R.,  1966 

Chiapas 

Jun 

1 

Jones,  1964  b 

Jul  i 

1 

Davis  et  al.,  1964 

Honduras 

May 

1 

tt 

Aug 

14 

2 

Valdez  and  LaVal,  1971 

Nicaragua 

Jul 

2 

Jones  et  al.,  1971a 

Aug 

1 

1 

ft 

Panama 

Jan 

2 

Fleming  et  al.,  1972 

Apr 

1 

tt 

Tobago 

Sep 

1 

Goodwin  and  Greenhall,  1961 

Colombia 

Jan 

4 

2 

1 

Thomas,  1972 

Feb 

1 

1 

ft 

Mar* 

1 

2 

tt 

Apr 

1 

1 

ft 

May 

1 

tt 

Jun 

6 

3 

tt 

Jul 

1 

7 

tt 

Aug 

1 

5 

Thomas,  1972 

Sep 

2 

tt 

Oct 

2 

I 

tt 

Nov 

4 

tt 

Peru 

Jun 

2 

Tuttle,  1970 

Jul 

1 

ft 

Pregnant  and  lactating. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


351 


Vampyrops  brachycephalis 

It  is  fruitless  to  speculate  on  the  reproductive  pattern  of  V.  brachycephalis 
on  the  basis  of  the  few  known  records  (Table  8). 

Vampyrops  heller i 

Jones  et  al.  (1971  a)  suggested  that  Nicaraguan  V.  helleri  probably  breed 
throughout  much  of  the  year.  Fleming  et  al.  (1972)  thought  that  this  species 
might  be  bimodally  polyestrous  based  on  their  evidence  from  Panama.  Thomas’ 
(1972)  work  in  Colombia,  by  far  the  most  extensive,  indicated  a  single  period 
of  nonpregnancies  from  July  through  September.  This  is  also  suggestive  of  a  bi- 
modal  polyestrous  pattern.  See  Table  8. 

Vampyrops  lineatus 

Peracchi  and  Albuquerque  (1971)  reported  pregnant  females  in  January, 
March,  and  December  in  Brazil. 

Vampyrops  recifinus 

Nothing  is  known  about  the  reproductive  pattern  of  V.  recifinus. 

Vampyrodes  caraccioli 

The  data  from  Colombia  (Table  8)  suggest  the  familiar  pattern  of  two  sequen¬ 
tial  breeding  periods  followed  by  a  quiescent  period,  as  indicated  here  by  fewer 
pregnancies  during  the  July-September  period. 

Vampyressa  pusilla 

Although  the  data  are  not  complete  (Table  9),  they  suggest  a  pattern  of  bi- 
modal  polyestry.  Panamanian  females  have  been  recorded  as  preg¬ 
nant  and  lactating  during  the  early  part  of  the  rainy  season,  whereas  records 
from  Colombia  indicate  the  mid-rainy  season  break  seen  in  other  species  in  this 
area. 

Vampyressa  melissa 

Nothing  is  known  about  the  reproductive  pattern  of  this  species,  although 
Gardner  (1976)  reported  three  reproductively  inactive  females  from  Peru  in 
May. 

Vampyressa  nymphaea 

Colombian  samples  (Table  9)  are  substantial,  and  indicate  the  familiar  pattern 
of  two  periods  of  activity  followed  by  a  quiescent  period.  The  time  of  inactivity 
seems  to  be  slightly  later  in  V.  nymphaea  than  in  other  species. 


352 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  9. —  Reproductive  data  for  the  genera  Vampyressa,  Chiroderma,  and  Ectophylla. 


Place  Date  Pregnant  Lactating  Inactive  Reference 


Vampyressa  pusilla 

Campeche 

Feb 

1 

Jones  et  al.,  1973 

Chiapas 

Jul 

1 

Davis  et  aL,  1964 

Guatemala 

Jul 

1 

Rick,  1968 

Honduras 

Aug 

1 

Valdez  and  LaVal,  1971 

Nicaragua 

Mar 

4 

Jones  et  al.,  1971a 

Jul 

1 

Starrett  and  de  la  Torre,  1964 

Costa  Rica 

Feb 

2 

Mares  and  Wilson,  1971 

Mar 

1 

rr 

Jul 

1 

Armstrong,  1969 

Aug 

1 

rr 

Panama 

Jan 

1 

Fleming  et  al.,  1972 

Mar 

1 

rr 

Apr 

1 

2 

Apr 

1 

Hall  and  Jackson,  1953 

Colombia 

Mar 

1 

Thomas,  1972 

Apr 

1 

1 

tt 

May 

2 

2 

rr  • 

Jul 

1 

1 

rr 

Aug 

1 

1 

rr 

Nov 

1 

1 

rr 

Aug 

1 

3 

Arata  and  Vaughn,  1970 

Vampyressa  nymphaea 

Nicaragua 

Feb 

1 

Jones  et  al.,  1971a 

Costa  Rica 

Apr 

2 

Gardner  et  al.,  1970 

Panama 

May 

1 

Hall  and  Jackson,  1953 

Colombia 

Jan 

29 

1 

2 

Thomas,  1972 

Feb 

4 

8 

rr 

Mar* 

9 

25 

1 

rr 

Apr* 

8 

3 

2 

rr 

May 

4 

1 

2 

rr 

Jun 

6 

5 

4 

rr 

Jul* 

15 

40 

13 

rr 

Aug* 

17 

13 

4 

rr 

Sep 

2 

rr 

Oct 

1 

rr 

Nov 

6 

rr 

Dec 

12 

1 

rr 

Chiroderma  villosum 

Chiapas 

May 

1 

Davis  et  al,,  1964 

Jul 

3 

2 

rr 

Dec 

2 

3 

rr 

Nicaragua 

Mar 

4 

1 

Jones  et  al.,  1971a 

Jul 

4 

rr 

Panama 

Mar 

1 

Fleming  et  al.,  1972 

Apr 

1 

3 

rr 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


353 


Table  9. —  Continued. 


Trinidad 

Aug 

1 

Goodwin  and  Greenhall,  1961 

Sep 

1 

It 

Colombia 

Jan 

1 

Thomas,  1972 

Peru 

Aug 

1 

Tuttle,  1970 

Chiroderma  salvini 

Chihuahua 

Jul 

2 

Anderson,  1972 

Sinaloa 

Jan 

1 

Jones  et  al.,  1972 

Jalisco 

Feb 

2 

Watkins  et  al.,  1972 

Colima 

Sep 

2 

Villa-R.,  1966 

Honduras 

Jul 

1 

Carter  et  al,  1966 

Jul 

1 

1 

LaVal,  1969 

Aug 

1 

1 

It 

Colombia 

Jan 

2 

1 

Thomas,  1972 

Mar* 

1 

2 

n 

Apr* 

1 

1 

n 

May 

1 

n 

Jun 

2 

1 

tt 

Jul 

3 

3 

tr 

Oct 

1 

n 

Nov 

1 

it 

Dec 

3 

n 

Chiroderma  trinitatum 

Panama 

Feb 

2 

Fleming  et  al,  1972 

May 

1 

1 

It 

Sep 

1 

It 

Trinidad 

Mar 

1 

Goodwin  and  Greenhall,  1961 

Colombia 

Jul 

1 

Thomas,  1972 

Peru 

Jul 

1 

Tuttle,  1970 

Brazil 

Jun 

1 

USNM 

Jul 

1 

It 

Ectophylla  macconnelli 

Colombia 

Jan 

1 

1 

Thomas,  1972 

Peru 

Aug 

1 

Tuttle,  1970 

‘Pregnant  and  lactating. 


Vampyressa  brocki 

The  only  published  record  of  reproductive  activity  in  V.  brocki  is  that  of 
Baker  et  al.  (1972),  who  reported  one  lactating  and  two  pregnant  females  from 
Colombia  that  were  taken  in  June  and  July. 

Vampyressa  bidens 

Davis  (1975)  reported  two  of  14  females  pregnant  in  December  in  Peru.  This 
appears  to  be  the  only  published  record  of  reproductive  activity  for  this  species. 


354 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Chiroderma  doriae 

Nothing  is  known  about  the  reproductive  pattern  of  C.  doriae. 

Chiroderma  improvisum 

No  information  is  available  on  reproduction  in  this  species. 

Chiroderma  villosum 

Although  the  records  listed  in  Table  9  are  diverse,  they  are  too  insufficient  to 
have  much  predictive  value.  Davis  et  al.  ( 1 964)  suggested  that  this  species  breeds 
throughout  the  year  on  the  basis  of  their  specimens  from  Chiapas.  These  data 
also  fit  the  pattern  of  bimodal  polyestry  fairly  well,  but  unfortunately  there  are  no 
records  from  late  in  the  rainy  season. 

Chiroderma  salvini 

This  species  is  obviously  polyestrous  in  Colombia,  and  when  further  data  are 
available,  may  prove  to  have  a  bimodal  pattern  similar  to  that  found  in  other 
species  of  Colombian  phyllostomatids.  See  Table  9. 

Chiroderma  trinitatum 

Analysis  of  the  reproductive  pattern  of  C.  trinitatum  must  await  further  data 
(see  Table  9).  Pregnancy  records  are  all  from  early  in  the  rainy  season  and  late 
the  dry  season. 

Ectophylla  alba 

Gardner  et  al.  (1970)  reported  a  pregnant  female  in  March  and  a  lactating 
female  in  April  from  Costa  Rica.  LaVal  (1977)  recorded  pregnant  females  in 
February  and  August  in  Costa  Rica.  He  also  found  one  lactating  female  in  March, 
and  postlactating  animals  in  September  and  November. 

Ectophylla  macconnelli 

In  addition  to  the  records  shown  in  Table  9,  A.  L.  Gardner  (personal  com¬ 
munication)  collected  a  lactating  female  in  May  and  a  pregnant  female  in  July 
from  Peru. 

Artibeus  cinereus 

The  records  for  Colombia  (Table  10)  are  in  accord  with  the  pattern  of  bi¬ 
modal  polyestry  as  suggested  for  several  other  Colombian  species.  Larger  samples 
would  help  to  define  pregnancy  and  birth  peaks. 

Artibeus  glaucus 

I  can  find  no  published  records  of  reproductive  activity  for  A.  glaucus,  but 
there  is  a  USNM  specimen  from  Venezuela  recorded  as  lactating  in  August. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


355 


Alfred  L.  Gardner  (personal  communication)  has  collected  inactive  females  in 
Peru  in  April  and  May. 

Artibeus  watsoni 

In  addition  to  the  data  in  Table  10,  Davis  (1970)  recorded  pregnant  females 
from  the  months  of  February,  March,  April,  July,  August,  and  November  from 
throughout  the  range  of  A.  watsoni  (southern  Mexico-Panama).  Fleming  (1973) 
suggested  that  this  species  provides  an  example  of  bimodal  polyestry  in  Panama. 
The  few  data  from  Nicaragua  also  fit  this  pattern. 

Artibeus  phaeotis 

Davis  (1970)  reported  pregnant  females  in  January,  February,  April,  June, 
July,  and  August,  and  inactive  females  from  all  other  months  except  November 
from  throughout  the  range  of  A.  phaeotis  (Sinaloa  to  Panama).  Fleming  (1973) 
reported  A.  phaeotis  as  seasonally  polyestrous  in  Panama,  and  the  data  from 
Mexico  seem  to  support  this.  Heithaus  et  al.  (1975)  suggested  bimodal  polyestry 
as  the  pattern  in  Costa  Rica  as  well.  See  Table  10. 

Artibeus  toltecus 

Davis  (1969)  recorded  pregnant  females  in  each  month  from  January  through 
August  in  Mexico  and  Central  America  (Table  10).  Davis  et  al.  (1964)  sug¬ 
gested  an  extended  breeding  season  for  A.  toltecus  and  mentioned  the  possibility 
of  their  having  two  births  per  year.  The  data  support  this  assertion. 

Artibeus  aztecus 

Davis  (1969)  mentioned  three  pregnant  and  two  inactive  females  taken  in 
March  and  April  in  either  southern  Mexico,  Guatemala,  or  Honduras.  The  data 
in  Table  10  from  northern  Mexico  suggest  that  these  bats  are  pregnant  during 
the  summer  months.  Additional  information  from  other  times  of  the  year  would 
be  useful  in  clarifying  the  reproductive  cycle. 

Artibeus  hirsutus 

Anderson  (1960)  suggested  that  A.  hirsutus  lacks  a  restricted  breeding  season. 
In  support  of  this  claim,  Findley  and  Jones  (1965)  found  in  Sonora  that  two  of 
the  lactating  females  had  placental  scars  while  a  third  had  sperm  in  the  uterus. 
They  also  found  three  males  with  sperm  and  eight  without  in  the  same  sample. 
As  they  pointed  out,  spermatogenesis,  copulation,  lactation,  and  parturition  all 
were  occurring  at  the  same  time.  See  Table  10. 

Artibeus  inopinatus 

Reproductive  information  for  A.  inopinatus  seems  to  be  lacking,  but  in  the 
description  of  the  species,  Davis  and  Carter  (1964)  mentioned  five  young  animals 
taken  in  August  in  Honduras.  Two  of  the  young  bats  appeared  to  be  about  one 


356 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  10. — Reproductive  data  for  the  genus  Artibeus. 


Place  Date  Pregnant  Lactating  Inactive  Reference 


Artibeus  cinereus 

Trinidad 

Sep 

X 

Goodwin  and  Greenhall,  1961 

Oct 

X 

II 

Venezuela 

Jul 

1 

2 

USNM 

Aug 

1 

II 

Colombia 

Jan 

9 

1*  2 

Thomas,  1972 

Feb 

1 

II 

Mar 

1 

II 

Apr 

2 

3 

It 

May 

1 

1  1 

It 

Jul 

1 

2 

II 

Aug 

1 

1  2 

II 

Sep 

1  4 

II 

Oct 

1 

2 

II 

Nov 

2 

2 

II 

Dec 

3 

6 

II 

Aug 

1 

5  9 

Arata  and  Vaughn,  1970 

Peru 

Jul 

1 

Tuttle,  1970 

Brazil 

Jun 

1 

USNM 

Jul 

1 

4 

II 

Artibeus  watsoni 

Guatemala 

Mar 

1 

Jones,  1966 

Nicaragua 

Feb 

1 

1 

Jones  et  al.,  1971a 

Aug 

1 

1 

II 

Panama 

Jan 

1 

Fleming  et  al,  1972 

Feb 

1 

II 

Apr 

2 

II 

Jun 

1 

II 

Aug 

2 

II 

Dec 

1 

3 

II 

Artibeus  phaeotis 

Sinaloa 

Jul 

4 

Jones  et  al.,  1972 

Oct 

1 

II 

Jalisco 

Jan 

6 

Watkins  et  al.,  1972 

Apr 

1 

II 

Jun 

11 

II 

Aug 

1 

II 

Campeche 

Jan 

2 

Jones  et  al.,  1973 

Feb 

7 

II 

Mar 

1 

1 

II 

Quintana  Roo 

Aug 

2 

II 

Apr 

1 

Birney  et  al.,  1974 

So.  Mexico 

Jan 

X 

Villa-R.,  1966 

Feb 

X 

II 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


357 


Table  10. — Continued. 


So.  Mexico 

Sep 

X 

Villa-R.,  1966 

Oct 

X 

ft 

Guatemala 

Mar 

1 

Jones,  1966 

Apr 

1 

Murie,  1935 

May* 

2 

Rick,  1968 

Panama 

Jan 

1 

Fleming  et  al.,  1972 

Feb 

9 

1 

ft 

Mar 

2 

ft 

Apr 

1 

1 

It 

Jun 

1 

ft 

Aug 

1 

2 

ft 

Artibeus  toltecus 

Tamaulipas 

Jul 

1 

de  la  Torre,  1954 

Sinaloa 

Jan 

X 

Jones  et  al.,  1972 

May 

X 

X 

ft 

Oct 

X 

ft 

Jalisco 

Jan 

9 

X 

Watkins  et  al.,  1972 

Feb 

7 

it 

Mar 

1 

ft 

Apr 

5 

ft 

Jun 

10 

X 

ft 

Jul 

1 

X 

ft 

Aug 

X 

ft 

Sep 

X 

ft 

Puebla 

Jan 

2 

1 

LaVal,  1972 

Chiapas 

May 

3 

Davis  et  al.,  1964 

Jun 

4 

ft 

Aug 

1 

ft 

El  Salvador 

Jan 

7 

Burt  and  Stirton,  1961 

Nicaragua 

Apr 

8 

Jones  et  al.,  1971  a 

Jun 

1 

Artibeus  aztecus 

it 

Tamaulipas 

Jul 

1 

Alvarez,  1963 

Aug 

1 

ft 

Durango 

Jul 

1 

Baker  and  Greer,  1962 

Sinaloa 

Jul 

18 

5 

Jones  et  al.,  1972 

Queretaro 

Jan 

5 

Schmidly  and  Martin,  1973 

Mexico 

Sep 

1 

Villa-R.,  1966 

Artibeus  hirsutus 

Chihuahua 

Jul 

1 

Anderson,  1972 

Sonora 

Apr 

1 

Cockrum  and  Bradshaw,  1963 

May 

2 

Cockrum,  1955 

May 

8 

Anderson,  1960 

Sep 

15 

6  4 

Findley  and  Jones,  1965 

Sinaloa 

Jun 

1  1 

Jones  et  al.,  1972 

Jul 

1 

ft 

Aug 

5 

2 

ft 

Dec 

1 

ft 

Jalisco 

Feb 

2 

2 

Watkins  et  al.,  1972 

Jun 

5 

X  15 

ft 

Aug 

1 

X  1 

ft 

358  SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  10. — Continued. 


Guerrero 

May 

2 

11 

Anderson,  1960 

Artibeus  jamaicensis 

Tamaulipas 

Mar 

6 

de  la  Torre,  1954 

May 

1 

Alvarez,  1963 

San  L.  Potosf 

Jun 

3 

Cockrum,  1955 

Sinaloa 

Jan 

X 

Jones  et  al.,  1972 

Feb 

X 

tt 

Apr 

X 

tt 

May 

X 

n 

Jun 

X 

it 

Jul 

X 

it 

Sep 

X 

it 

Nov 

X 

it 

Jun 

3 

Anderson,  1960 

Nayarit 

Apr 

X 

Villa-R.,  1966 

Jalisco 

Jul 

1 

Anderson,  1960 

Jan 

X 

Watkins  et  al.,  1972 

Feb 

X 

It 

Mar 

X 

ft 

Apr 

X 

X 

It 

May 

X 

X 

tr 

Jun 

X 

tt 

Jul 

X 

it 

Oct 

X 

rt 

Guerrero 

Feb 

1 

Villa-R.,  1966 

Morelos 

Jul 

4 

Novick,  1960 

Queretaro 

Jan 

13 

Schmidly  and  Martin,  1973 

Puebla 

Jan 

2 

LaVal,  1972 

Veracruz 

Feb 

X 

Hall  and  Dalquest,  1963 

Jul 

6 

3 

Webb  etal.,  1967 

Aug** 

1 

Barlow  and  Tamsitt,  1968 

Yucatan  Pen. 

Apr 

1 

Bowles,  1973 

May 

2 

ft 

Feb 

1 

1 

Jones  et  al.,  1973 

Apr 

1 

ft 

May 

1 

It 

Jul 

X 

X 

X 

tt 

Aug 

X 

tt 

Mar 

4 

X 

Birney  et  al,  1974 

Apr 

3 

X 

tt 

Isla  Cozumel 

Aug 

5 

6 

Jones  and  Lawlor,  1965 

Oaxaca 

Apr 

1 

USNM 

Guatemala 

Jan 

6 

Jones,  1966 

Feb 

4 

1 

tt 

Mar 

1 

2 

tt 

Aug 

4 

It 

May 

1 

Rick,  1968 

El  Salvador 

Dec 

16 

Burt  and  Stirton,  1961 

Costa  Rica 

Jan 

2 

2 

Fleming  et  al.,  1972 

Feb 

16 

5 

14 

It 

Mar 

1 

20 

2 

It 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


359 


Table  10. — Continued. 


Costa  Rica 

Apr 

7 

12 

4 

Fleming  et  al.,  1972 

May 

10 

1 

2 

ft 

Jun 

1 

1 

tt 

Jul 

2 

9 

tt 

Sep 

1 

1 

it 

Oct 

51 

tt 

Nov 

1 

15 

tt 

Dec 

1 

1 

16 

it 

Aug* 

X 

X 

4 

Tamsitt  and  Valdivieso,  1961 

Panama 

Jan 

41 

7 

Fleming  et  al.,  1972 

Feb 

15 

2 

tt 

Mar 

23 

42 

11 

tt 

Apr* 

12 

18 

tt 

May* 

22 

5 

ft 

Jun 

14 

4 

4 

tt 

Jul 

10 

21 

21 

ft 

Aug 

19 

15 

tt 

Sep 

21 

26 

ft 

Oct 

4 

20 

ft 

Nov 

15 

tt 

Dec 

1 

4 

ft 

Jamaica 

Dec 

6 

4 

Goodwin,  1970 

Feb 

6 

1 

2 

McNab,  1976 

Providencia 

Jan 

4 

5 

Tamsitt  and  Mejia,  1962 

Puerto  Rico 

Feb 

X 

Fenton,  1969 

Feb 

X 

Tamsitt,  1970 

Mar 

X 

X 

tt 

Apr 

X 

tt 

Jun 

X 

tt 

Jun 

X 

X 

Anthony,  1918 

Aug 

2 

Tamsitt  and  Valdivieso,  1970 

Virgin  Is. 

Apr** 

X 

Barlow  and  Tamsitt,  1968 

Jul** 

X 

ft 

Trinidad 

Feb 

X 

X 

Goodwin  and  Greenhall,  1961 

Mar 

X 

X 

tt 

Apr 

X 

X 

tt 

May 

X 

X 

tt 

Jun 

X 

tt 

Jul 

X 

X 

tt 

Sep 

X 

tt 

Jun 

X 

Jones,  1946 

Colombia 

Jun 

1 

Tamsitt  and  Valdivieso,  1963  6 

Jul 

1 

1 

6 

Arata  and  Vaughn,  1970 

Aug 

3 

14 

ft 

Jan 

14 

4 

8 

Thomas,  1972 

Feb 

3 

ft 

Mar* 

1 

3 

1 

ft 

Apr* 

3 

tt 

May 

2 

5 

ft 

Jun 

1 

2 

2 

tt 

Jul* 

2 

10 

13 

tt 

Aug 

1 

3 

16 

tt 

360 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  10. — Continued. 


Colombia 

Sep 

1 

15 

Thomas,  1972 

Oct 

1 

2 

tt 

Nov 

3 

2 

tt 

Dec 

9 

5 

tt 

Venezuela 

Jul 

4 

2 

1 

Smith  and  Genoways,  1974 

Peru 

Jun 

14 

Tuttle,  1970 

Jul 

4 

It 

Aug 

3 

tt 

Artibeus  lituratus 

Tamaulipas 

Mar 

2 

de  la  Torre,  1954 

May 

10 

Alvarez,  1963 

Durango 

Jun 

2 

3 

Jones,  1964  c 

Sinaloa 

Feb 

X 

Jones  et  al.,  1972 

Apr 

X 

It 

Jun 

X 

tt 

Jul 

X 

X 

ft 

Oct 

X 

tt 

Nov 

13 

Anderson,  1960 

Jalisco 

Mar 

2 

X 

Watkins  et  al.,  1972 

Apr 

2 

X 

tt 

Jun 

3 

tt 

Jul 

1 

X 

ft 

Aug 

X 

tt 

Sep 

X 

ft 

Oct 

X 

ft 

Morelos 

May 

1 

Cockrum,  1955 

Queretaro 

Jan 

1 

Spenrath  and  LaVal,  1970 

Veracruz 

Feb 

1 

Hall  and  Dalquest,  1963 

Yucatan  Pen. 

Jan 

2 

Jones  et  al.,  1973 

Feb 

1 

tt 

Apr 

2 

tt 

Jul 

1 

tt 

Oaxaca 

Apr 

5 

Villa-R.,  1966 

Guatemala 

Feb 

2 

Jones,  1966 

Mar 

3 

1 

ft 

May 

2 

Rick,  1968 

El  Salvador 

Jul 

1 

Starrett  and  de  la  Torre,  1964 

Costa  Rica 

Jul 

1 

ft 

Jan 

1 

1 

Fleming  et  al.,  1972 

Feb 

1 

ft 

Apr 

2 

tt 

May 

1 

1 

ft 

Jul 

2 

2 

ft 

Sep 

3 

ft 

Oct 

1 

ft 

Nov 

3 

tt 

Dec 

10 

tt 

Panama 

Jan 

9 

3 

tt 

Mar 

1 

2 

ft 

Apr 

2 

2 

1 

ft 

May 

1 

tt 

BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


361 


Table  10. — Continued. 


Panama 

Aug 

1 

Fleming  et  al.,  1972 

Sep 

6 

n 

Oct 

1 

n 

Mar 

3 

1 

Bloedel,  1955 

Apr 

1 

Hall  and  Jackson,  1953 

May 

2 

ft 

Trinidad 

Feb 

X 

Goodwin  and  Greenhall,  1961 

Mar 

X 

II 

Apr 

X 

X 

It 

May 

X 

X 

ft 

Jun 

X 

X 

It 

Jul 

X 

X 

It 

Aug 

X 

II 

Sep 

X 

It 

Oct 

X 

II 

Venezuela 

Aug 

1 

USNM 

Jul 

1 

1 

Smith  and,Genoways,  1974 

French  Guiana 

Aug 

X 

Brosset  and  Dubost,  1967 

Sep 

X 

ft 

Colombia 

Jan 

9 

1 

Tamsitt  and  Valdivieso,  1965  a 

Feb 

1 

II 

Mar* 

1 

1 

It 

Apr 

1 

It 

May 

3 

1 

3 

It 

Jun* 

4 

5 

6 

It 

Jul 

1 

1 

II 

Aug 

1 

1 

II 

Sep* 

4 

It 

Oct* 

4 

1 

9 

It 

Nov* 

8 

2 

8 

II 

Jan 

18 

24 

21 

Thomas,  1972 

Feb 

14 

12 

It 

Mar 

9 

5 

10 

It 

Apr* 

13 

4 

9 

It 

May* 

22 

10 

16 

It 

Jun 

8 

13 

18 

It 

Jul* 

7 

4 

13 

It 

Aug* 

3 

9 

23 

It 

Sep 

3 

7 

19 

It 

Oct 

23 

4 

32 

II 

Nov 

33 

1 

27 

It 

Dec 

30 

7 

33 

It 

Jul 

5 

10 

32 

Arata  and  Vaughn,  1970 

Aug 

3 

II 

Peru 

Jul 

2 

Tuttle,  1970 

Brazil 

Jun 

1 

USNM 

Jul 

2 

6 

II 

Jul 

X 

Peracchi  and  Albuquerque,  1971 

Aug 

X 

II 

’Pregnant  and  lactating. 
’’Twins. 


362 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


month  old  and  the  others  were  older,  but  still  in  subadult  pelage.  Baker  and  Jones 
(1975)  also  recorded  August-taken  young  from  Nicaragua. 

Artibeus  concolor 

The  only  record  of  reproduction  in  A.  concolor  is  that  of  Thomas  (1972),  who 
collected  a  pregnant  female  in  February  in  Colombia. 

Artibeus  jamaicensis 

Artibeus  jamaicensis  is  one  of  the  few  species  for  which  adequate  information 
on  reproduction  is  available  (Table  10).  Goodwin  (1970)  reported  that  breed¬ 
ing  is  generally  synchronized  in  Jamaica;  Tamsitt  and  Mejia  (1962)  discussed  a 
restricted  season  on  Providencia;  and  Felten  (1956a)  suggested  that  breeding 
occurs  in  the  dry  season  in  El  Salvador.  On  the  other  hand,  Tamsitt  (1966)  and 
Jones  et  al.  (1973)  argued  for  continuous  or  acyclic  breeding  behavior  in  Colom¬ 
bia  and  the  Yucatan  Peninsula,  respectively.  Fleming  et  al.  (1972)  and  Fleming 
(1973)  have  shown  this  species  to  be  seasonally  polyestrous  in  Panama  and 
Costa  Rica.  Fleming  et  al.  (1972)  also  presented  data  on  testis  size  correlated 
with  spermatogenic  activity  in  males.  Heithaus  et  al.  (1975)  supported  the  case 
for  bimodal  polyestry  in  Costa  Rica,  pointing  out  that  the  two  birth  peaks  occur 
at  times  of  peak  flower  and  fruit  availability. 

Fleming  (1971)  has  shown  that  A.  jamaicensis  has  a  unique  seasonally  poly¬ 
estrous  cycle  in  Panama.  A  peak  in  parturition  occurs  in  March  and  April,  fol¬ 
lowed  by  postpartum  estrous  and  a  second  peak  in  parturition  in  July  and  August. 
“Blastocysts  conceived  after  the  second  birth  implant  in  the  uterus  but  are  dor¬ 
mant  from  September  to  mid-November,  when  normal  development  again  re¬ 
sumes”  (Fleming,  1971:402).  Embryos  then  develop  and  young  are  bom  during 
the  March-April  birth  peak. 

Artibeus  lituratus 

In  the  northern  part  of  its  distribution,  A.  lituratus  produces  only  one  young 
per  year,  but  farther  southward,  the  period  of  reproductive  activity  is  extended 
(Table  10).  In  Costa  Rica  and  Panama,  these  bats  probably  are  on  a  bimodal 
cycle  with  a  quiescent  period  after  the  second  birth  peak  in  the  rainy  season 
(Heithaus  et  al.,  1975).  In  Colombia,  breeding  proceeds  throughout  the  year 
(Tamsitt  and  Valdivieso,  1963a).  Tamsitt  (1966)  noted  that  A.  lituratus  is  an 
acyclic  or  continuous  breeder  in  Colombia.  Tamsitt  and  Valdivieso  (19656) 
studied  the  reproductive  cycle  of  males  in  Colombia  and  their  data,  based  on 
presence  of  sperm,  length  and  tubule  diameter  of  the  testes,  and  diameter  of  the 
epididymides,  indicate  that  males  are  capable  of  reproductive  activity  at  any  time 
of  the  year,  and  that  the  reproductive  pattern  is  acyclic  without  any  suggestion  of 
seasonal  variation. 

Thomas  (1972)  presented  a  much  more  extensive  sample  from  Colombia  and, 
although  he  has  confirmed  year-round  activity  with  the  presence  of  pregnant, 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


363 


lactating,  and  inactive  females  in  every  month  of  the  year,  his  data  indicate  bi- 
modal  activity  peaks.  Pregnancy  peaks  occur  in  December  and  May,  with  lacta¬ 
tion  peaks  lagging  about  a  month  behind,  as  would  be  expected. 

Enchisthenes  hartii 

Gardner  et  al.  (1970)  suggested  that  E.  hartii  is  reproductively  active  through¬ 
out  the  year  in  Costa  Rica.  The  only  inactive  animals  they  found  were  subadults, 
one  in  May  and  three  in  July.  This  species  may  be  found  to  undergo  a  period  of 
reproductive  inactivity  when  data  become  available  from  later  in  the  year.  See 
Table  1 1. 

Ardops  nichollsi 

I  can  find  no  records  other  than  those  of  Jones  and  Schwartz  (1967)  who  re¬ 
ported  four  pregnant  females  in  March  and  one  lactating  and  two  pregnant  fe¬ 
males  in  April  from  Dominica. 

Phyllops  falcatus 

No  information  is  available  on  reproduction  in  this  species. 

Phyllops  haitiensis 

Nothing  is  known  about  the  reproductive  pattern  of  P.  haitiensis. 

Ariteus  flavescens 

No  data  are  available  concerning  the  reproductive  pattern  of  this  species. 

Stenoderma  rufum 

Tamsitt  and  Valdivieso  (1966  b)  described  parturition  in  S.  rufum.  This 
species  seems  to  be  polyestrous  on  Puerto  Rico,  but  data  from  the  period  Septem¬ 
ber  through  December  are  needed  in  order  to  clarify  their  reproductive  pattern. 
See  Table  11. 

Pygoderma  bilabiatum 

Peracchi  and  Albuquerque  (1971)  reported  a  pregnant  female  collected  in 
August  in  Brazil. 

Ametrida  centurio 

Nothing  is  known  about  the  reproductive  pattern  of  this  species. 

Sphaeronycteris  toxophyllum 

Nothing  has  been  published  about  the  reproductive  pattern  of  S.  toxophyllum. 


364 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1 1. — Reproductive  data  for  the  genera  Enchisthenes,  Stenoderma,  and  Centurio. 


Place 

Date 

Pregnant  Lactating  Inactive 

Reference 

Enchisthenes  hartii 

Honduras 

Aug* 

LaVal,  1969 

Costa  Rica 

Jan 

1 

Gardner  et  al.,  1970 

May 

1 

X 

tt 

Jun 

1 

X 

tt 

Jul 

X 

tt 

Aug 

1 

Armstrong,  1969 

Aug 

1 

LaVal,  1977 

Colombia 

Apr 

7 

5 

Thomas,  1972 

May 

12 

13 

ft 

Jul 

1  3 

ft 

Aug 

7 

ft 

Sep 

1 

ft 

Peru 

Nov 

1 

Gardner,  1976 

Stenoderma  rufum 

Puerto  Rico 

Feb 

X 

X 

Tamsitt,  1970 

Mar 

X 

tt 

May 

X 

tt 

Jul 

X 

rr 

Aug 

X 

tt 

Nov 

X 

tt 

Jul 

6 

1  9 

Jones  et  al.,  1971  b 

Jul 

1 

Genoways  and  Baker,  1972 

Aug 

1 

tt 

Aug 

1 

Tamsitt  and  Valdivieso,  19666 

Centurio  senex 

Tamaulipas 

Jun 

1 

Alvarez,  1963 

Jalisco 

Mar 

2 

1 

Watkins  et  al.,  1972 

Aug 

1 

tt 

Apr 

1 

Jones,  19646 

Veracruz 

Apr 

5 

Villa-R.,  1966 

Yucatan  Pen. 

Jan 

1 

Jones  et  al.,  1973 

Feb 

1 

tt 

Jul 

1 

ft 

Oaxaca 

Mar 

1 

Villa-R.,  1966 

Mar 

1 

1 

USNM 

Chiapas 

Apr 

1 

Davis  et  al.,  1964 

Jul 

1 

tt 

Honduras 

Aug 

2 

LaVal,  1969 

Nicaragua 

Feb 

1 

Jones  et  al.,  1971  a 

Mar 

2 

1 

ft 

Costa  Rica 

Mar 

1 

Mares  and  Wilson,  1971 

Trinidad 

Jan 

1 

Goodwin  and  Greenhall,  1961 

Oct 

1 

tt 

Pregnant  and  lactating. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


365 


Centurio  senex 

Although  there  are  a  fair  number  of  records  for  C.  senex  (Table  11),  the  data 
from  any  given  area  are  too  few  to  decipher  reproductive  patterns.  Pregnancies 
from  February  and  July  on  the  Yucatan  Peninsula  suggest  the  possibility  of 
either  polyestry  or  asynchrony. 

Brachyphylla  cavernarum 

Anthony  (1918)  reported  lactating  females  in  July  in  Puerto  Rico,  and  Nellis 
(1971)  found  a  lactating  female  in  April  on  St.  Croix.  Walker  (1975)  mentioned 
pregnant  females  in  February  on  Puerto  Rico,  March  on  St.  Croix,  and  a  lactating 
female  in  April  on  Puerto  Rico. 

Buden  (1977)  collected  12  females,  all  of  which  were  pregnant,  in  March  on 
the  Island  of  Caicos  in  the  West  Indies.  All  fetuses  were  24  to  34  mm.  in  length, 
suggesting  a  synchronized  cycle.  The  females  lactating  in  July  (Anthony,  1918) 
suggest  the  possibility  of  a  second  period  of  parturition  as  well. 

Brachyphylla  nana 

Nothing  is  known  about  the  reproductive  pattern  of  this  species. 

Erophylla  bombifrons 

Although  the  data  are  sparse  (Table  12),  they  suggest  a  restricted  breeding 
season.  Females  are  pregnant  from  February  to  June  and  lactating  in  July.  This 
would  result  in  the  production  of  young  early  in  the  rainy  season,  a  time  when 
resources  should  be  most  plentiful. 

Erophylla  sezekorni 

Buden  (1976)  summarized  data  based  on  91  pregnant  or  lactating  females  and 
immatures.  He  suggested  a  gestation  period  during  the  first  part  of  the  year  with 
parturition  in  early  summer.  He  found  females  carrying  small  fetuses  in  February 
and  larger  fetuses  in  April  and  May.  Lactating  females  were  taken  in  June  and 
many  juveniles  in  July.  Immature  animals  approaching  adult  size  were  taken  in 
August.  Thus,  the  pattern  appears  identical  to  that  described  above  for  E.  bombi¬ 
frons.  See  Table  12. 

Phyllonycteris  poeyi 

Miller  (1904)  reported  that  all  of  the  females  he  examined  from  Cuba  were 
pregnant  in  June. 

Phyllonycteris  major 

Nothing  is  known  about  reproductive  patterns  of  P.  major,  a  bat  which  is  likely 
extinct. 


366 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


■ 


Table  12. —  Reproductive  date  for  the  genus  Erophylla. 


Place 

Date 

Pregnant  Lactating  Inactive 

Reference 

Erophylla  bombifrons 

Hispaniola 

Feb 

1 

Buden,  1976 

Puerto  Rico 

Jul 

1 

ft 

Apr 

X 

Barlow  and  Tamsitt,  1968 

Jun 

X 

Valdivieso  et  al.,  1968 

Mar 

X 

Walker,  1975 

Apr 

X 

tt 

May 

X 

tt 

Jul 

X 

tt 

Erophylla  sezekorni 

Cuba 

Feb 

11 

Buden,  1976 

Bahamas 

Apr 

11 

tt 

May 

6 

tt 

Jun 

4 

tt 

Jul 

1* 

rt 

Jun 

8  2 

Blake,  1885 

♦Plus  many  immatures. 


Phyllonycteris  aphylla 

The  only  record  of  reproductive  activity  in  this  species  is  that  of  Goodwin 
(1970),  who  reported  a  pregnant  female  taken  in  January  on  Jamaica. 

Desmodus  rotundus 

More  is  known  about  the  reproduction  of  D.  rotundus  than  about  any  other 
phyllostomatid  (Table  13).  DeVerteiul  and  Urich  (1936)  apparently  were  the 
first  to  suggest  that  D.  rotundus  breeds  year-round,  based  on  their  work  on 
Trinidadian  populations.  Wimsatt  and  Trapido  (1952)  confirmed  this  in  Panama 
by  presenting  data  on  both  males  and  females,  and  suggested  a  gestation  period  of 
five  to  six  months.  Burt  and  Stirton  (1961)  reported  continuous  breeding  in  El 
Salvador.  Goodwin  and  Greenhall  (1961)  recorded  the  same  thing  for  popula¬ 
tions  on  Trinidad  and  reported  pregnant  females,  lactating  females,  and  young 
animals  in  every  month,  although  the  highest  incidence  of  young  was  in  April  and 
May  and  again  in  October  and  November.  They  also  suggested  that  males  may 
roost  separately  from  females  when  the  young  are  born. 

Crespo  et  al.  (1961)  gave  a  detailed  account  of  reproduction  in  vampires 
based  on  their  work  in  Argentina  during  September  and  November.  They  found 
that  in  males  both  testes  are  active  and  coincide  in  their  activity  rhythm.  Sexually 
active  males  with  well-developed  epididymides  containing  spermatozoa  and  in¬ 
active  males  with  small  epididymides  and  no  spermatozoa  were  found  in  the 
same  population  at  the  same  time  of  year.  Sexually  active  males  were  present  in 
September  and  November.  In  some  instances,  adult  males  have  epididymides  with 
few  spermatozoa  mixed  with  resting  cells,  which  could  be  interpreted  as  the 
beginning  of  a  new  cycle  of  activity. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


367 


For  females,  Crespo  et  al.  (1961)  found  that  both  ovaries  are  functional,  with 
only  a  slight  difference  in  degree  of  development  of  follicles.  Ovaries  are  in  a 
periovarian  capsule,  and  the  fallopian  tubes  begin  in  the  walls  of  the  capsules. 
There  is  always  only  one  embryo,  which  occupies  one  uterine  horn  first  but,  with 
development  extends  into  both  horns  and  the  body  of  the  uterus,  obliterating  the 
partitioning  of  the  uterus.  At  the  end  of  a  pregnancy,  the  ovary  without  the  corpus 
luteum  is  in  early  proestrous  and  will  produce  the  next  ovum.  One  postpartum 
specimen  had  a  corpus  luteum  in  one  ovary  and  a  corpus  albicans  representing  a 
previous  pregnancy  in  the  other  ovary. 

In  September  and  November,  there  are  proestrous  immature  animals  bearing 
primary  and  secondary  follicles.  None  of  the  animals  examined  had  vaginal  plugs 
or  sperm  in  the  uteri. 

Hall  and  Dalquest  (1963)  mentioned  that  these  animals  seem  to  have  no 
regular  breeding  season  in  Veracruz.  They  found  a  few  young  in  various  stages 
of  development,  pregnant  females,  and  inactive  females  in  all  of  the  colonies 
examined.  Dalquest  (1955)  had  earlier  pointed  this  out  for  San  Luis  Potosf 
populations,  and  suggested  that  young  are  bom  in  all  months  of  the  year. 

Villa-R.  (1966)  found  pregnant  females,  lactating  females,  and  newborn 
young  at  all  times  of  the  year  during  1 5  years  work  in  Mexico. 

Greenhall  (1965)  described  mating  behavior  (including  copulation),  preg¬ 
nancy,  and  young  animals  in  captivity.  Schmidt  and  Manske  (1973)  found  a 
gestation  period  of  seven  months  and  lactation  period  of  three  to  nine  months  for 
captive  animals.  Linhart  (1971)  compiled  a  useful  bibliography  of  vampire  bats. 

Diaemus  youngii 

The  only  recorded  reproductive  information  for  this  species  is  that  of  Goodwin 
and  Greenhall  (1961)  for  Trinidad.  They  found  two  lactating  females  in  August 
and  in  October  they  took  one  immature  male,  four  pregnant  females,  one  lactating 
female,  and  two  inactive  females. 

Diphylla  ecaudata 

Dalquest  (1955)  reported  that  D.  ecaudata  seems  to  have  a  well-defined  breed¬ 
ing  season  and  may  have  a  single  young  per  year  in  eastern  Mexico.  Felten 
(1956a),  however,  felt  that  they  breed  in  both  dry  and  wet  seasons  in  El  Salvador 
and  postulated  two  litters  per  year.  From  the  scatter  in  the  records  listed  in  Table 
13,  I  am  inclined  to  agree  with  Felten. 

Summary 

The  three  most  obvious  reproductive  strategies  found  in  the  family  Phyl- 
lostomatidae  are  summarized  in  Fig.  1.  The  most  critical  environmental  param¬ 
eter  is  the  seasonality  of  the  rainfall  pattern.  Although  a  great  deal  of  geographic 
variation  exists,  the  pattern  of  a  dry  season  during  the  months  of  January  to  April 
or  May  is  common  in  Middle  America  and  in  many  areas  in  northern  South 
America.  In  tropical  Mexico,  the  rains  often  begin  as  late  as  June,  but  as  one 


368 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  13. —  Reproductive  data  for  the  genera  Desmodus  and  Diphylla. 


Place  Date  Pregnant  Lactating  Inactive  Reference 


Tamaulipas 

Mar 

May 

Jun 

Aug 

Chihuahua 

May 

Durango 

Jun 

Sinaloa 

Jan 

Mar 

May 

Dec 

Nayarit 

Jan 

Jalisco 

Jan 

Feb 

Mar 

Apr 

May 

Jun 

Jul 

Aug 

Sep 

Colima 

Mar 

May 

Jul 

Zacatecas 

Oct 

Michoacan 

Jul 

Guerrero 

Jun 

Aug 

Sep 

Nov 

Queretaro 

May 

Jun 

Dec 

Puebla 

Jan 

Morelos 

Jan 

Mexico 

Jan* 

Veracruz 

Feb 

Jun 

Jul 

Yucatan  Pen. 

Jan 

Feb 

Mar 

Apr 

Jun 

Jul 

Aug 

Apr 

Desmodus  rotundus 

1 
2 

1  5 

9 
4 

1 
1 
1 
1 
1 
1 

X 
X 
X 
X 
X 
X 
X 

X 
X 

52  1 

36  9 

2  23 

1 
1 

22 
16 
21 
10 
1 
1 

3 
2 
1 
1 

X 

4 
1 
1 
1 
2 

4  X 

1 

1  X 
X 

2 


Alvarez,  1963 

tt 

tt 

tt 

Anderson,  1972 
Jones,  1964c 
Jones  et  al.,  1972 

ft 

tt 

tt 

Cockrum,  1955 
Watkins  et  al.,  1972 

ft 

tt 

ft 

tt 

tt 

tt 

tt 

tt 

Burns  and  Crespo,  1975 

tt 

ft 

Cockrum,  1955 
Hall  and  Villa-R„  1949 
Forment  et  al.,  1971 

tt 

tt 

tt 

Schmidly  and  Martin,  1973 

ft 

tt 

LaVal,  1972 
Burns,  1970 

ft 

Hall  and  Dalquest,  1963 
Lackey,  1970 

ft 

Jones  et  al,  1973 

ft 

tt 

tt 

tt 

ft 

ft 

Birney  et  al.,  1974 


39 

23 

1 


7 

31 

43 

12 

3 

1 

2 


12 

1 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


369 


Table  13. — 

■Continued. 

Guatemala 

Mar 

3 

Jones,  1966 

El  Salvador 

Feb 

X 

Felten,  1956c 

Mar 

5 

II 

May 

2 

II 

Jul 

1 

tl 

Aug 

1 

II 

Oct 

1 

It 

Nov 

X 

It 

Costa  Rica 

Jan 

4 

6 

Fleming  et  al„  1972 

Feb 

7 

2 

15 

It 

Mar 

1 

1 

5 

It 

Apr 

1 

2 

II 

May 

2 

2 

II 

Jul 

1 

4 

II 

Aug 

4 

II 

Oct 

1 

II 

Nov 

1 

2 

It 

Dec 

4 

II 

Panama 

Apr 

2 

1 

II 

May 

1 

1 

II 

Feb 

6 

4 

Wimsatt  and  Trapido,  1952 

Apr 

5 

II 

May 

1 

3 

II 

Jul 

1 

2 

It 

Nov 

2 

1 

II 

Trinidad 

Jan 

X 

DeVerteiul  and  Urich,  1936 

Jun 

X 

It 

Nov 

X 

II 

Dec 

X 

It 

Colombia 

Nov 

X 

Tamsitt  and  Valdivieso,  1963  6 

Jul 

3 

3 

12 

Arata  and  Vaughn,  1970 

Apr 

1 

Thomas,  1972 

May 

1 

It 

Oct 

1 

1 

1 

II 

Venezuela 

Apr 

1 

6 

Pirlot  and  Leon,  1965 

Brazil 

Jan 

X 

X 

Peracchi  and  Albuquerque,  1971 

Diphylla  ecaudata 

Tamaulipas 

Nov 

2 

1 

Alvarez,  1963 

San  L.  Potosi 

Mar 

3 

Dalquest,  1953 

Jul 

X 

II 

Yucatan 

Nov 

1 

2 

Hatt,  1938 

May 

1 

Birney  et  aL,  1974 

Mexico 

Aug 

1 

Villa-R.,  1966 

Oct 

7 

II 

Nov 

2 

2 

II 

El  Salvador 

Aug 

1 

Felten,  1956c 

Honduras 

Jul 

2 

1 

3 

Valdez  and  LaVal,  1971 

Nicaragua 

Apr 

2 

Jones  et  at .,  1971  a 

370 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


ASEASONAL  POLYESTRY 


LACTATION  LACTATION 

GESTATION  PARTURITION  GESTATION 


BIMODAL  POLYESTRY 


GESTATION  PARTURITION 

MONESTRY 


LACTATION 

GESTATION  PARTURITION 


DESMODUS 


PARTURITION 


GLOSSOPHAGA 
CAROLL1A 
URODERMA 
ART  I  BE  US 

LACTATION 


MAC ROT US 
LEPTONYCTERIS 


GESTATION  PARTURITION  LACTATION 


maximum 

stress 

MAXIMUM  FRUIT 

MAXIMUM  INSECTS 

dry 

rains 

begin 

heavy 

rains 

JAN 

FEB  MAR 

APR 

MAY 

JUN  JUL  AUG  SEP 

OCT  NOV  DEC 

Fig.  1. — Summary  of  the  three  common  reproductive  patterns  and  of  the  environmental 
events  affecting  them. 


moves  southward,  the  length  of  the  dry  season  generally  decreases,  and  in  some 
areas  may  be  only  a  month  or  less  in  duration.  Also,  annual  variations  occur  in 
any  given  locality.  Nevertheless,  for  purposes  of  this  discussion,  the  pattern  rep¬ 
resented  in  Fig.  1  may  be  taken  as  representative. 

This  environmental  seasonality  affects  reproductive  cycles  of  bats  through  the 
food  supply.  The  time  of  maximum  abundance  of  a  wide  variety  of  both  fruits  and 
insects  is  just  after  the  beginning  of  the  rainy  season.  Thereafter,  a  general  de¬ 
cline  is  seen,  culminating  in  a  period  of  minimal  abundance  during  the  dry  season. 

The  critical  time  for  most  bat  populations  seems  to  be  the  period  of  weaning 
of  the  young  (Wilson  and  Findley,  1970;  Fleming  et  al.,  1972).  Thus,  although 
it  may  be  possible  for  females  to  undergo  gestation  and  lactation  during  the  stress¬ 
ful  time  of  year,  the  young  are  usually  weaned  during  the  most  energetically 
favorable  periods. 

In  monestrous  species  of  the  Phyllostomatidae,  there  is  a  distinct  period  of 
reproductive  activity  culminating  with  weaning  of  the  young  shortly  after  the  be¬ 
ginning  of  the  rainy  season  when  food  is  plentiful.  This  pattern  is  seen  in  some 
species  at  the  northern  limit  of  the  range  of  the  family,  where  the  time  of  maxi¬ 
mum  food  availability  is  fairly  short.  The  nectarivorous  bats  of  the  genus 
Leptonycteris  show  this  pattern  in  the  southwestern  United  States,  where  they 
migrate  northward  from  Mexico  and  have  their  young  in  May  or  June.  These 
young  are  weaned  in  July  or  August,  the  peak  of  the  rainy  season  and  the  period 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


371 


Table  14. — Reproductive  patterns  of  the  20  species  for  which  adequate  data  exist. 


Macrotus  waterhousii 

delayed  development  and  monestry 

Glossophaga  soricirui 

continuous  or  bimodal  polyestry 

Leptonycteris  sanborni 

monestry  or  bimodal  polyestry 

Choeronycteris  mexicana 

monestry 

Carollia  castanea 

bimodal  polyestry 

Carollia  subrufa 

continuous  or  bimodal  polyestry 

Carollia  brevicauda 

bimodal  polyestry 

Carollia  perspicillata 

bimodal  polyestry 

Sturnira  1  ilium 

bimodal  polyestry 

Uroderma  bilobatum 

bimodal  polyestry 

Vampyrops  helleri 

bimodal  polyestry 

Vampyrodes  caraccioli 

bimodal  polyestry 

Vampyressa  pusilla 

bimodal  polyestry 

Vampyressci  nymphaea 

bimodal  polyestry 

Artibeus  cinereus 

bimodal  polyestry 

Artibeus  watsoni 

bimodal  polyestry 

Artibeus  phaeotis 

bimodal  polyestry 

Artibeus  jamaicensis 

bimodal  polyestry  and  delayed  development 

Artibeus  lituratus 

geographically  variable 

Desmodus  rotundus 

continuous  polyestry 

of  peak  flower  abundance.  In  October,  individuals  migrate  back  to  Mexico  for 
the  winter.  A  variation  of  this  pattern  is  found  in  Macrotus  californicus ,  where 
the  embryos  undergo  delayed  development  during  the  autumn  and  winter  months 
and  begin  developing  at  a  more  normal  rate  in  spring.  This  results  in  parturition 
and  weaning  periods  similar  to  those  of  Leptonycteris.  There  is  a  possibility  that 
some  individuals  of  Leptonycteris  sanborni  have  a  second  period  of  reproductive 
activity  resulting  in  the  production  of  offspring  in  Mexico  in  November.  If  so  this 
species  would  more  properly  belong  in  the  next  category,  that  of  bimodal 
polyestry. 

The  majority  of  species  of  phyllostomatids  for  which  there  is  ample  data  show  a 
reproductive  pattern  involving  an  extended  breeding  season  with  two  birth  peaks 
a  year.  In  these  species  (for  example,  some  members  of  the  genera  Glossophaga, 
Carollia,  Uroderma,  and  Artibeus),  the  young  from  the  first  birth  peak  are  weaned 
at  the  beginning  of  the  rainy  season,  and  those  from  the  second  pregnancy  of  the 
year  are  weaned  well  into  the  rainy  season.  These  two  peaks  are  followed  by  an 
inactive  period,  which  results  in  no  young  being  weaned  during  the  stressful  dry 
season. 

At  the  other  extreme  from  monestry  are  those  animals  that  are  completely 
polyestrous  and  produce  young  continuously  and  asynchronously  throughout  the 
year.  The  evidence  to  date  shows  only  the  vampire  bat  Desmodus  rotundus  to 
be  in  this  category.  These  animals  are  adapted  to  a  food  supply  (primarily  blood 
from  domestic  cattle)  that  is  available  throughout  the  year  over  much  of  their 
range.  However,  because  their  gestation  period  is  five  or  six  months  long,  the 
net  result  still  is  only  two  young  per  year. 


372 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  14  summarizes  the  type  of  reproductive  pattern  for  the  20  species  for 
which  there  is  a  reasonable  amount  of  data.  It  should  be  noted  that  many  of  these 
species  will  show  geographic  variation  in  the  timing  of  reproductive  events,  and, 
in  some  cases  ( Artibeus  lituratus,  for  example),  the  species  may  have  completely 
different  patterns  in  different  areas.  This  is  hardly  surprising  in  view  of  the  wide 
geographic  and  ecologic  range  of  many  of  the  species. 

All  of  these  patterns  may  be  thought  of  as  variations  on  a  single  theme — 
maximizing  the  production  of  offspring  with  available  environmental  energy 
resources.  Further  study  will  undoubtedly  add  a  wealth  of  data  on  the  fine  tuning 
of  the  various  mechanisms  involved  in  selecting  for  a  particular  reproductive 
strategy  for  a  given  species. 


Acknowledgments 

I  gratefully  thank  the  following  people:  Michael  A.  Bogan,  Estella  C.  Duell, 
Robert  D.  Fisher,  Theodore  H.  Fleming,  Alfred  L.  Gardner,  Barbara  A.  Harvey, 
Daniel  H.  Janzen,  Clyde  Jones,  Patricia  Mehlhop,  S.  Jerrine  Nichols,  and  William 
A.  Wimsatt. 


Literature  Cited 

Allen,  G.  M.  1939.  Bats.  Harvard  Univ.  Press,  Cambridge,  Massachusetts,  x  +  368  pp. 
Alvarez,  T.  1963.  The  Recent  mammals  of  Tamaulipas,  Mexico.  Univ.  Kansas  Publ., 
Mus.  Nat.  Hist.,  14:363-473. 

Alvarez,  T.,  and  J.  Ramirez- Pulido.  1972.  Notas  acerca  de  murcielagos  mexicanos. 
An.  Esc.  Nac.  Cien.  Biol.,  Mexico,  19:167-178. 

Anderson,  S.  1960.  Neotropical  bats  from  Western  Mexico.  Univ.  Kansas  Publ.,  Mus. 
Nat.  Hist.,  14:1-8. 

- .  1969.  Macrotus  waterhousiu  Mammalian  Species,  1 : 1-4. 

- .  1972.  Mammals  of  Chihuahua:  taxonomy  and  distribution.  Bull.  Amer.  Mus. 

Nat.  Hist.,  148:149-410. 

Anthony,  H.  E.  1918.  The  indigenous  land  mammals  of  Puerto  Rico,  living  and  extinct. 
Mem.  Amer.  Mus.  Nat.  Hist.,  n.s.,  2:331-435. 

Arata,  A.  A.,  and  J.  B.  Vaughn.  1970.  Analyses  of  the  relative  abundance  and  re¬ 
productive  activity  of  bats  in  southwestern  Colombia.  Caldasia,  10:517-528. 
Armstrong,  D.  M.  1969.  Noteworthy  records  of  bats  from  Costa  Rica.  J.  Mamm., 
50:808-810. 

Axtell,  R.  W.  1962.  An  easternmost  record  for  the  bat  Choeronycteris  mexicana  from 
Coahuila,  Mexico.  Southwestern  Nat.,  7:76. 

Baker,  R.  H.  1956.  Mammals  of  Coahuila,  Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat. 
Hist.,  9:125-335. 

Baker,  R.  H.,  and  J.  K.  Greer.  1962.  Mammals  of  the  Mexican  state  of  Durango.  Publ. 
Mus.,  Michigan  State  Univ.,  2:25-154. 

Baker,  R.  H.,  and  D.  Womochel.  1966.  Mammals  from  southern  Oaxaca.  Southwestern 
Nat.,  1 1:306. 

Baker,  R.  J.,  H.  H.  Genoways,  and  A.  Cadena.  1972.  The  phyllostomatid  bat,  Vampyres- 
sa  brocki,  in  Colombia.  Bull.  S.  California  Acad.  Sci.,  71:54. 

Baker,  R.  J.,  and  J.  K.  Jones,  Jr.  1975.  Additional  records  of  bats  from  Nicaragua,  with 
a  revised  checklist  of  Chiroptera.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  32:1-13. 
Baker,  R.  J.,  W.  J.  Bleier,  and  W.  R.  Atchley.  1975.  A  contact  zone  between  karyotypi- 
cally  characterized  taxa  of  Uroderma  bilobatum  (Mammalia:Chiroptera). 
Syst.  Zool.,  24:133-142. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


373 


Barbour,  R.  W.,  and  W.  H.  Davis.  1969.  Bats  of  America.  Univ.  Press  Kentucky, 
Lexington,  286  pp. 

Barlow,  J.  C.,  and  J.  R.  Tamsitt.  1968.  Twinning  in  American  leaf-nosed  bats  (Chirop- 
tera:  Phyllostomatidae).  Canadian  J.  Zool.,  46:290-292. 

Birney,  E.  C.,  J.  B.  Bowles,  and  R.  M.  Timm.  1974.  Mammalian  distributional  records 
in  Yucatan  and  Quintana  Roo,  with  comments  on  reproduction,  structure,  and 
status  of  peninsular  populations.  Occas.  Papers  Bell  Mus.  Nat.  Hist.,  13:1-25. 

Blake,  H.  A.  1885.  Note  on  the  parturition  of  a  West  Indian  bat.  Sci.  Proc.  Royal 
Dublin  Soc.,  4:449. 

Bloedel,  P.  1955.  Observations  on  the  life  histories  of  Panama  bats.  J.  Mamm., 
36:232-235. 

Bowles,  J.  B.  1973.  Notes  on  reproduction  in  four  species  of  bats  from  Yucatan,  Mexico. 
Trans.  Kansas  Acad.  Sci.,  75:271-272. 

Bradshaw,  G.  V.  R.  1961.  Le  cycle  de  reproduction  de  Macrotus  californicus  (Chirop- 
tera:  Phyllostomatidae).  Mammalia,  25:1 17-1 19. 

- .  1962.  Reproductive  cycle  of  the  California  leaf-nosed  bat,  Macrotus  californicus. 

Science,  136:645. 

Brosset,  A.,  and  G.  Dubost.  1967.  Chiropteres  de  la  Guyane  Francaise.  Mammalia, 
31:583-594. 

Buden,  D.  W.  1975.  A  taxonomic  and  zoogeographic  appraisal  of  the  big-eared  bat 
( Macrotus  waterhousii  Gray)  in  the  West  Indies.  J.  Mamm.,  56:758-769. 

- .  1976.  A  review  of  the  bats  of  the  endemic  West  Indian  genus  Erophylla.  Proc. 

Biol.  Soc.  Washington,  89:1-16. 

- .  1977.  First  records  of  bats  of  the  genus  Brachyphylla  from  the  Caicos  Islands, 

with  notes  on  geographic  variation.  J.  Mamm.,  58:221-225. 

Burns,  R.  J.  1970.  Twin  vampire  bats  born  in  captivity.  J.  Mamm.,  51:391-392. 

Burns,  J.,  and  R.  F.  Crespo.  1975.  Notes  on  local  movement  and  reproduction  of  vam¬ 
pire  bats  in  Colima,  Mexico.  Southwestern  Nat.,  19:446-449. 

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. 
Zool.,  Univ.  Michigan,  117:1-69. 

Campbell,  B.  1934.  Notes  on  bats  collected  in  Arizona  during  the  summer  of  1933. 
J.  Mamm.,  15:241-242. 

Carter,  D.  C.  1966.  A  new  species  of  Rhinophylla  (Mammalia,  Chiroptera,  Phyl¬ 
lostomatidae)  from  South  America.  Proc.  Biol.  Soc.  Washington,  79:235-238. 

- .  1968.  A  new  species  of  Anoura  (Mammalia:  Chiroptera:  Phyllostomatidae) 

from  South  America.  Proc.  Biol.  Soc.  Washington,  81:427-430. 

Carter,  D.  C.,  R.  H.  Pine,  and  W.  B.  Davis.  1966.  Notes  on  Middle  American  bats. 
Southwestern  Nat.,  11:488-499. 

Cockrum,  E.  L.  1955.  Reproduction  in  North  American  bats.  Trans.  Kansas  Acad. 
Sci.,  58:487-511. 

Cockrum,  E.  L.,  and  G.  V.  R.  Bradshaw.  1963.  Notes  on  mammals  from  Sonora, 
Mexico.  Amer.  Mus.  Novit.,  2138:1-9. 

Cockrum,  E.  L.,  and  E.  Ordway.  1959.  Bats  of  the  Chiricahua  Mountains,  Cochise 
County,  Arizona.  Amer.  Mus.  Novit.,  1938:1-35. 

Colinvaux,  P.  A.  1973.  Introduction  to  ecology.  John  Wiley  and  Sons,  Inc.,  New  York, 
ix  +  621  pp. 

Crespo,  J.  A.,  J.  M.  Vanella,  B.  D.  Blood,  and  J.  M.  De  Carlo.  1961.  Observaciones 
ecologicas  del  vampiro  Desmodus  rotundas  (Geoffroy)  en  el  norte  de  Cordoba. 
Rev.  Mus.  Argentino  Cien.  Nat.  “Bernardo  Rivadavia,"  6:131-160. 

Dalquest,  W.  W.  1953.  Mammals  of  the  Mexican  state  of  San  Luis  Potosf.  Louisiana 
State  Univ.  Studies,  Biol.  Sci.  Ser.,  1:1-229. 

- .  1955.  Natural  history  of  the  vampire  bats  of  eastern  Mexico.  Amer.  Midland 

Nat.,  53:79-87. 


374 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Davis,  W.  B.  1966.  The  mammals  of  Texas.  Bull.  Texas  Parks  and  Wildlife  Dept., 
41:1-267. 

- .  1968.  Review  of  the  genus  Uroderma  (Chiroptera).  J.  Mamm.,  49:676-698. 

- .  1969.  A  review  of  the  small  fruit  bats  (genus  Artibeus)  of  Middle  America. 

Southwestern  Nat.,  14:15-29. 

- .  1970.  A  review  of  the  small  fruit  bats  (genus  Artibeus)  of  Middle  America. 

Part  II.  Southwestern  Nat.,  14:389-402. 

- .  1975.  Individual  and  sexual  variation  in  Vampyressa  bidens.  J.  Mamm., 

56:262-265. 

Davis,  W.  B.,  and  D.  C.  Carter.  1964.  A  new  species  of  fruit-eating  bat  (genus  Artibeus) 
from  Central  America.  Proc.  Biol.  Soc.  Washington,  77:119-122. 

Davis,  W.  B„  D.  C.  Carter,  and  R.  H.  Pine.  1964.  Noteworthy  records  of  Mexican  and 
Central  American  bats.  J.  Mamm.,  45:375-387. 
de  la  Torre,  L.  1954.  Bats  from  southern  Tamaulipas,  Mexico.  J.  Mamm.,  35:1 13-1 16. 
DeVerteiul,  E„  and  F.  W.  Urich.  1936.  The  study  and  control  of  paralytic  rabies  trans¬ 
mitted  by  bats  in  Trinidad,  British  West  Indies.  Trans.  Royal  Soc.  Trop.  Med. 
Hyg.,  29:317-347. 

Easterla,  D.  A.  1972.  Status  of  Leptonycteris  nivalis  (Phyllostomatidae)  in  Big  Bend 
National  Park,  Texas.  Southwestern  Nat.,  17:287-292. 

Enders,  R.  K.  1935.  Mammalian  life  histories  from  Barro  Colorado  Island,  Panama. 

Bull.  Mus.  Comp.  Zool.  Harvard  College,  78:385-502. 

Felten,  H.  1956«.  Quiropteros  (Mammalia:Chiroptera)  en  El  Salvador.  Comm. 
Inst.  Trop.  Invest.  Cient.,  Univ.  El  Salvador,  5:153-170. 

- .  19566.  Fledermause  (Mammalia:  Chiroptera)  aus  El  Salvador.  Part  2.  Senck. 

Biol.,  37:69-86. 

- .  1956c.  Fledermause  (Mammalia:  Chiroptera)  aus  El  Salvador.  Part  3.  Senck. 

Biol.,  37:179-212. 

Fenton,  M.  B.  1969.  The  carrying  of  young  by  females  of  three  species  of  bats.  Cana¬ 
dian  J.  Zool.,  47:158-159. 

Findley,  J.  S.,  and  C.  Jones.  1965.  Northernmost  records  of  some  Neotropical  bat 
genera.  J.  Mamm.,  46:330-331. 

Fleming,  T.  H.  1971.  Artibeus  jamaicensis’.  delayed  embryonic  development  in  a  Neo¬ 
tropical  bat.  Science,  171:402-404. 

- .  1973.  The  reproductive  cycles  of  three  species  of  opossums  and  other  mammals 

in  the  Panama  Canal  Zone.  J.  Mamm.,  54:439-455. 

Fleming,  T.  H.,  E.  T.  Hooper,  and  D.  E.  Wilson.  1972.  Three  Central  American  bat 
communities:  structure,  reproductive  cycles,  and  movement  patterns.  Ecology, 
53:555-569. 

Forment,  W.  L.,  U.  Schmidt,  and  A.  M.  Greenhall.  1971.  Movement  and  population 
studies  of  the  vampire  bat  ( Desmodus  rotundas)  in  Mexico.  J.  Mamm.,  52:227- 
228. 

Gardner,  A.  L.  1976.  The  distributional  status  of  some  Peruvian  mammals.  Occas. 

Papers  Mus.  Zool.,  Louisiana  State  Univ.,  48: 1-18. 

Gardner,  A.  L.,  and  J.  P.  O'Neill.  1969.  The  taxonomic  status  of  Sturnira  bidens 
(Chiroptera:  Phyllostomatidae)  with  notes  on  its  karyotype  and  life  history. 
Occas.  Papers  Mus.  Zool.,  Louisiana  State  Univ.,  38:1-8. 

Gardner,  A.  L.,  R.  K.  LaVal,  and  D.  E.  Wilson.  1970.  The  distributional  status  of 
some  Costa  Rican  bats.  J.  Mamm.,  51:712-729. 

Genoways,  H.  H.,  and  R.  J.  Baker.  1972.  Stenoderma  rufum.  Mammalian  Species, 
18:1-4. 

Genoways,  H.  H.,  and  J.  K.  Jones,  Jr.  1975.  Additional  records  of  the  Stenodermine 
bat,  Sturnira  thomasi,  from  the  Lesser  Antillean  island  of  Guadeloupe.  J.  Mamm., 
56:924-925. 


BIOLOGY  OF  THE  PHYLLOSTOMAT1DAE 


375 


Goodwin,  G.  G.  1958.  Three  new  bats  from  Trinidad.  Amer.  Mus.  Novit.,  1877:1-6. 

Goodwin,  G.  G.,  and  A.  M.  Greenhall.  1961.  A  review  of  the  bats  of  Trinidad  and 
Tobago.  Bull.  Amer.  Mus.  Nat.  Hist.,  122:187-302. 

Goodwin,  R.  E.  1970.  The  ecology  of  Jamaican  bats.  J.  Mamm.,  51:571-579. 

Greenhall,  A.  M.  1965.  Notes  on  behavior  of  captive  vampire  bats.  Mammalia, 
29:441-451. 

- .  1968.  Notes  on  the  behavior  of  the  false  vampire  bat.  J.  Mamm.,  49:337-340. 

Grinnell,  H.  W.  1918.  A  synopsis  of  the  bats  of  California.  Univ.  California  Publ. 
Zool.,  17:223-404. 

Hahn,  W.  L.  1907.  A  review  of  the  bats  of  the  genus  Hemiderma.  Proc.  U.S.  Nat. 
Mus.,  32:103-118. 

Hall,  E.  R.,  and  W.  W.  Dalquest.  1963.  The  mammals  of  Veracruz.  Univ.  Kansas 
Publ.,  Mus.  Nat.  Hist.,  14:165-362. 

Hall,  E.  R.,  and  W.  B.  Jackson.  1953.  Seventeen  species  of  bats  recorded  from  Barro 
Colorado  Island,  Panama  Canal  Zone.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist., 
5:641-646. 

Hall,  E.  R.,  and  B.  Villa-R.  1949.  An  annotated  checklist  of  the  mammals  of  Michoa- 
can,  Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  1:431-472. 

Hamlett,  G.  W.  D.  1935.  Notes  on  the  embryology  of  a  phyllostomid  bat.  Amer.  J. 
Anat.,  56:327-353. 

Harrison,  D.  L.  1975.  Macrophyllum  macrophyllum.  Mammalian  Species,  62:1-3. 

Hatt,  R.  T.  1938.  Notes  concerning  mammals  collected  in  Yucatan.  J.  Mamm.,  19:333- 
337. 

Hayward,  B.  J.,  and  E.  L.  Cockrum.  1971.  The  natural  history  of  the  western  long- 
nosed  bat  Leptonycteris  sanbornl  WRI-SCI  (Western  New  Mexico  Univ.), 
1:75-123. 

Heithaus,  E.  R.,  T.  H.  Fleming,  and  P.  A.  Opler.  1975.  Foraging  patterns  and  resource 
utilization  in  seven  species  of  bats  in  a  seasonal  tropical  forest.  Ecology,  56:841- 
854. 

Hoffmeister,  D.  R.,  and  W.  W.  Goodpaster.  1954.  The  mammals  of  the  Huachuca 
Mountains,  southeastern  Arizona.  Illinois  Biol.  Monogr.,  24:1-152. 

Homan,  J.  A.,  and  J.  K.  Jones,  Jr.  1975.  Monophyllus  redmani.  Mammalian  Species, 
57:1-3. 

Huey,  L.  M.  1925.  Food  of  the  California  leaf-nosed  bat.  J.  Mamm.,  6:196-197. 

Jones,  J.  K.,  Jr.  1964a  Bats  new  to  the  fauna  of  Nicaragua.  Trans.  Kansas  Acad. 
Sci.,  67:506-508. 

- .  1964 b.  Bats  from  western  and  southern  Mexico.  Trans.  Kansas  Acad.  Sci., 

67:509-516. 

- .  1964c.  Additional  records  of  mammals  from  Durango,  Mexico.  Trans.  Kansas 

Acad.  Sci.,  66:750-753. 

- .  1966.  Bats  from  Guatemala.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist.,  16:439-472. 

Jones,  J.  K.,  Jr.,  and  D.  C.  Carter.  1976.  Annotated  checklist,  with  keys  to  subfamilies 
and  genera.  Pp.  7-38,  in  Biology  of  bats  of  the  New  World  family  Phyl- 
lostomatidae.  Part  I  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.),  Spec. 
Publ.  Mus.,  Texas  Tech  Univ.,  10:1-218. 

Jones,  J.  K.,  Jr.,  and  T.  E.  Lawlor.  1965.  Mammals  from  Isla  Cozumel,  Mexico,  with 
description  of  a  new  species  of  harvest  mouse.  Univ.  Kansas  Publ.,  Mus.  Nat. 
Hist.,  16:409-419. 

Jones,  J.  K.,  Jr.,  and  C.  J.  Phillips.  1976.  Bats  of  the  genus  Sturnira  in  the  Lesser 
Antilles.  Occas.  Papers  Mus.,  Texas  Tech  Univ.,  40:1-16. 

Jones,  J.  K.,  Jr.,  and  G.  L.  Phillips.  1964.  A  new  subspecies  of  the  fruit-eating  bat 
Sturnira  ludovici,  from  western  Mexico.  Univ.  Kansas  Publ.,  Mus.  Nat.  Hist., 
14:475-481. 


376 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Jones,  J.  K.,  Jr.,  and  A.  Schwartz.  1967.  Bredin-Archbold-Smithsonian  Biological  Survey 
of  Dominica.  6.  Synopsis  of  bats  of  the  Antillean  genus  Ardops.  Proc.  U.S.  Nat. 
Mus.,  124(3634):  1-13. 

Jones,  J.  K.,  Jr.,  J.  D.  Smith,  and  T.  Alvarez.  1965.  Notes  on  bats  from  the  Cape  region 
of  Baja  California.  Trans.  San  Diego  Soc.  Nat.  Hist.,  14:53-56. 

Jones,  J.  K.,  Jr.,  J.  D.  Smith,  and  R.  W.  Turner.  1971«.  Noteworthy  records  of  bats 
from  Nicaragua,  with  a  checklist  of  the  chiropteran  fauna  of  the  country.  Occas. 
Papers  Mus.  Nat.  Hist.,  Univ.  Kansas,  2:1-35. 

Jones,  J.  K.,  Jr.,  H.  H.  Genoways,  and  R.  J.  Baker.  19716.  Morphological  variation  in 
Stenoderma  rufum.  J.  Mamm.,  52:244-247. 

Jones,  J.  K.,  Jr.,  J.  R.  Choate,  and  A.  Cadena.  1972.  Mammals  from  the  Mexican  State 
of  Sinaloa.  II.  Chiroptera.  Occas.  Papers  Mus.  Nat.  Hist.,  Univ.  Kansas,  6:1-29. 
Jones,  J.  K.,  Jr.,  J.  D.  Smith,  and  H.  H.  Genoways.  1973.  Annotated  checklist  of 
mammals  of  the  Yucatan  Peninsula,  Mexico.  I.  Chiroptera.  Occas.  Papers  Mus., 
Texas  Tech  Univ.,  13:1-31. 

Jones,  T.  S.  1946.  Parturition  in  a  West  Indian  fruit  bat  (Phyllostomidae).  J.  Mamm., 
27:327-330. 

Kuhlhorn,  F.  1953.  Saugetierkundliche  studien  aus  Sud-Mattogrosso.  Saugetierk. 
Mitteil.,  1:115-122. 

Lackey,  J.  A.  1970.  Distributional  records  of  bats  from  Veracruz.  J.  Mamm.,  51:384- 
385. 

LaVal,  R.  K.  1969.  Records  of  bats  from  Honduras  and  El  Salvador.  J.  Mamm., 
50:819-822. 

- .  1972.  Distributional  records  and  band  recoveries  of  bats  from  Puebla,  Mexico. 

Southwestern  Nat.,  16:449-451. 

- .  1977.  Notes  on  some  Costa  Rican  bats.  Brenesia,  10/1 1:77-83. 

Linhart,  S.  B.  1971.  A  partial  bibliography  of  the  vampire  bats  ( Desmodus,  Diphylla, 
Diaemus).  U.S.  Bureau  Sport  Fisheries  and  Wildlife,  Denver,  iv  +  51  pp. 

Mares,  M.  A.,  and  D.  E.  Wilson.  1971.  Bat  reproduction  during  the  Costa  Rican  dry 
season.  BioScience,  21:471-477. 

Marinkelle,  C.  J.  1970.  Vampyrops  intermedius  sp.  n.  from  Colombia  (Chiroptera: 

Phyllostomatidae).  Rev.  Brasil.  Biol.,  30:49-53. 

Matson,  J.  O.,  and  D.  R.  Patten.  1975.  Notes  on  some  bats  from  the  state  of  Zacatecas, 
Mexico.  Contrib.  Sci.,  Nat.  Hist.  Mus.  Los  Angeles  Co.,  263:1-12. 

McNab,  B.  K.  1976.  Seasonal  fat  reserves  of  bats  in  two  tropical  environments. 
Ecology,  57:332-338. 

Merriam,  C.  H.  1898.  Mammals  of  Tres  Marias  Islands,  off  western  Mexico.  Proc. 
Biol.  Soc.  Washington,  12:13-19. 

Miller,  G.  S.,  Jr.  1904.  Notes  on  the  bats  collected  by  William  Palmer  in  Cuba.  Proc. 
U.S.  Nat.  Mus.,  27:337-348. 

Mumford,  R.  E„  and  D.  A.  Zimmerman.  1962.  Notes  on  Choeronycteris  mexicana. 
J.  Mamm.,  43:101-102. 

Mumford,  R.  E.,  L.  L.  Oakley,  and  D.  A.  Zimmerman.  1964.  June  bat  records  from 
Guadalupe  Canyon,  New  Mexico.  Southwestern  Nat.,  9:43-45. 

Murie,  A.  1935.  Mammals  from  Guatemala  and  British  Honduras.  Misc.  Publ.  Mus. 
Zool.,  Univ.  Michigan,  26:1-30. 

Nellis,  D.  W.  1971.  Additions  to  the  natural  history  of  Brachyphylla  (Chiroptera). 
Caribbean  J.  Sci.,  1 1:91. 

Novick,  A.  1960.  Successful  breeding  in  captive  Artibeus.  J.  Mamm.,  41:508-509. 
Osburn,  W.  1865.  Notes  on  the  Chiroptera  of  Jamaica.  Proc.  Zool.  Soc.  London, 
pp.  61-85. 

Pearse,  A.  S.,  and  R.  Kellogg.  1938.  Mammalia  from  Yucatan  caves.  Publ.  Carnegie 
Inst.  Washington,  491:301-304. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


377 


Peracchi,  A.  L.,  and  S.  T.  de  Albuquerque.  1971.  Lista  provisoria  dos  quiropteros  des 
estados  do  Rio  de  Janeiro  e  Guanatara,  Brazil  (Mammalia,  Chiroptera).  Rev. 
Brasil.  Biol.,  31:405-413. 

Phillips,  C.  J.,  and  J.  K.  Jones,  Jr.  1971.  A  new  subspecies  of  the  long-nosed  bat, 
Hylonycteris  underwood  i,  from  Mexico.  J.  Mamm.,  52:77-80. 

Pine,  R.  H.  1972.  The  bats  of  the  genus  Carollia.  Tech.  Monogr.  Agric.  Exp.  Sta., 
Texas  A&M  Univ.,  8:1-125. 

Pirlot,  P.  1963.  Algunas  consideraciones  sobre  la  ecologia  de  los  marruferos  del  oeste  de 
Venezuela.  Rev.  Universidad  Zulia,  Kasmera,  1:169-214. 

Pirlot,  P.,  and  J.  R.  Leon.  1965.  Chiropteres  de  1’est  du  Venezuela.  I.  Region  de  Cumana 
et  lie  de  Margarita.  Mammalia,  29:367-374. 

Rasweiler,  J.  J.  1972.  Reproduction  in  the  long-tongued  bat,  Glossophaga  soricina. 

I.  Preimplantation  development  and  history  of  the  oviduct.  J.  Reprod.  Fert., 
31:249-262. 

Rick,  A.  M.  1968.  Notes  on  bats  from  Tikal,  Guatemala.  J.  Mamm.,  49:516-520. 

Rouk,  C.  S.,  and  D.  C.  Carter.  1972.  A  new  species  of  Vampyrops  (Chiroptera: 

Phyllostomatidae)  from  South  America.  Occas.  Papers  Mus.,  Texas  Tech  Univ., 
1:1-7. 

Schaldach,  W.  J.,  Jr.  1965.  Notas  breves  sobre  algunos  mamiferos  del  sur  de  Mexico. 
Anal.  Inst.  Biol.,  35:129-137. 

- .  1966.  New  forms  of  mammals  from  southern  Oaxaca,  Mexico,  with  notes  on 

some  mammals  of  the  coastal  range.  Saugetierk.  Mitteil.,  14:286-297. 

Schmidly,  D.  J.,  and  C.  O.  Martin.  1973.  Notes  on  bats  from  the  Mexican  state  of 
Queretaro.  Bull.  S.  California  Acad.  Sci.,  72:90-92. 

Schmidt,  U.,  and  U.  Manske.  1973.  Die  Jugendentwicklung  der  Vampirfledermause 
(Desmodtts  rotundus).  Zeit.  Saugetierk.,  38:14-33. 

Schwartz,  A.,  and  J.  K.  Jones,  Jr.  1967.  Bredin-Archbold-Smithsonian  Biological 
Survey  of  Dominica.  7.  Review  of  bats  of  the  endemic  Antillean  genus  Mono 
phyllus.  Proc.  U.S.  Nat.  Mus.,  124(3635):  1-20. 

Smith,  J.  D.,  and  H.  H.  Genoways.  1974.  Bats  of  Margarita  Island,  Venezuela,  with 
zoogeographic  comments.  Bull.  S.  California  Acad.  Sci.,  73:64-79. 

Spenrath,  C.  A.,  and  R.  K.  LaVal.  1970.  Records  of  bats  from  Queretaro  and  San  Luis 
Potosi,  Mexico.  J.  Mamm.,  51:395-396. 

Starrett,  A.,  and  L.  de  la  Torre.  1964.  Notes  on  a  collection  of  bats  from  Central 
America,  with  the  third  record  for  Cyttarops  alecto  Thomas.  Zoologica,  49:53-63. 

Taddei,  V.  A.  1976.  The  reproduction  of  some  phyllostomidae  (Chiroptera)  from  the 
Northwestern  region  of  the  state  of  Sao  Paulo.  Bolm.  Zool.,  Univ.  Sao  Paulo, 
1:313-330. 

Tamsitt,  J.  R.  1966.  Altitudinal  distribution,  ecology,  and  general  life  history  of  bats  in 
the  Andes  of  Colombia.  Amer.  Phil.  Soc.  Yearbook,  pp.  372-373. 

- .  1970.  Comparative  biochemistry  and  ecology  of  bats  of  the  Puerto  Rican  region. 

Amer.  Phil.  Soc.  Yearbook,  pp.  342-343. 

Tamsitt,  J.  R.,  and  C.  Mejia.  1962.  The  reproductive  status  of  a  population  of  the  neo¬ 
tropical  bat,  Artibeus  jamaicensis,  at  Providencia.  Caribbean  J.  Sci.,  2:139-144. 

Tamsitt,  J.  R.,  and  D.  Valdivieso.  1961.  Notas  sobre  actividades  nocturnas  y  estados 
de  reproduccion  de  algunos  quiropteros  de  Costa  Rica.  Rev.  Biol.  Trop., 
9:219-225. 

- .  1963a  Reproductive  cycle  of  the  big  fruit-eating  bat,  Artibeus  lituratus  Olfers. 

Nature,  198:104. 

- .  19636.  Records  and  observations  on  Colombian  bats.  J.  Mamm.,  44:168-180. 

- .  1964.  Informations  sur  la  reproduction  des  cheiropteres  phyllostomides  de 

Colombie.  Mammalia,  28:397-402. 

- .  1965a  Reproduction  of  the  female  big  fruit-eating  bat,  Artibeus  lituratus 

pulmarum,  in  Colombia.  Caribbean  J.  Sci.,  5:157-166. 


378 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


- .  1965  6.  The  male  reproductive  cycle  of  the  bat  Artibeus  lituratus.  Amer.  Mid¬ 
land  Nat.,  73:150-160. 

- .  1966a.  Taxonomic  comments  on  Anoura  caudifer,  Artibeus  lituratus  and 

Molossus  molossus.  J.  Mamm.,  47:230-238. 

- .  19666.  Parturition  in  the  red  fig-eating  bat  Stenoderma  rufum.  J.  Mamm., 

47:352-353. 

- .  1970.  Observations  on  bats  and  their  ectoparasites.  Pp.  El  23-El  28,  in  A 

tropical  rain  forest  (H.  T.  Odum  and  R.  F.  Pigeon,  eds.),  U.S.  Atomic  Energy 
Commission,  Washington,  D.C.,  xiv  +  1652  pp. 

Tamsitt,  J.  R.,  D.  Valdivieso,  and  J.  Hernandez  C.  1965.  Additonal  records  of 
Choeroniscus  in  Colombia.  J.  Mamm.,  46:704. 

Thomas,  M.  E.  1972.  Preliminary  study  of  the  annual  breeding  patterns  and  population 
fluctuations  of  bats  in  three  ecologically  distinct  habitats  in  Southwestern  Colom¬ 
bia.  Unpublished  Ph.D.  dissertation,  Tulane  Univ.,  161  pp. 

Thomas,  M.  E.,  and  D.  N.  McMurray.  1974.  Observations  on  Sturnira  aratathomasi 
from  Colombia.  J.  Mamm.,  55:834-836. 

Tuttle,  M.  D.  1970.  Distribution  and  zoogeography  of  Peruvian  bats,  with  comments  on 
natural  history.  Univ.  Kansas  Sci.  Bull.,  49:45-86. 

Valdez,  R.,  and  R.  K.  LaVal.  1971.  Records  of  bats  from  Honduras  and  Nicaragua. 
J.  Mamm.,  52:247-250. 

Valdivieso,  D.,  E.  Conde,  and  J.  R.  Tamsitt.  1968.  Lactate  dehydrogenase  studies  in 
Puerto  Rican  bats.  Comp.  Biochem.  Physiol.,  27:133-138. 

Villa-R.,  B.  1966.  Los  murcielagos  de  Mexico.  Inst.  Biol.,  Univ.  Nac.  Autonoma  de 
Mexico,  xvi  +  491  pp. 

Villa-R.,  B.,  and  M.  Villa-C.  1969.  Algunos  murcielagos  del  norte  de  Argentina.  Misc. 
Publ.  Mus.  Nat.  Hist.,  Univ.  Kansas,  51:407-428. 

Walker,  E.  P.  1975.  Mammals  of  the  World.  Johns  Hopkins  Press,  Baltimore,  3rd  ed., 
rev.,  l:xlx  + 1-644. 

Watkins,  L.  C.,  J.  K.  Jones,  Jr.,  and  H.  H.  Genoways.  1972.  Bats  of  Jalisco,  Mexico. 
Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  1:1-44. 

Webb,  R.  G.,  R.  H.  Baker,  and  P.  L.  Dalby.  1967.  Vertebrados  de  la  Isla  del  Toro, 
Veracruz.  An.  Inst.  Biol.,  Univ.  Nac.  Autonoma  Mexico,  38:1-8. 

Wilson,  D.  E.  1973.  Reproduction  in  Neotropical  bats.  Period.  Biol.,  75:215-217. 

Wilson,  D.  E„  and  J.  S.  Findley.  1970.  Reproductive  cycle  of  a  Neotropical  insectivorous 
bat,  Myotis  nigricans.  Nature,  225: 1 155. 

Wimsatt,  W.  A.,  and  H.  Trapido.  1952.  Reproduction  and  the  female  reproductive  cycle 
in  the  tropical  American  vampire  bat,  Desmodus  rotundus  murinus.  Amer.  J. 
Anat.,  91:415-445. 


EMBRYOLOGY 


William  J.  Bleier 


Over  the  years,  there  have  been  numerous  reports  concerning  reproduction 
in  the  phyllostomatid  bats,  but  a  survey  of  the  literature  reveals  that  data  on 
the  embryology  of  the  Phyllostomatidae  are  limited  to  gross  morphological 
observations  of  reproductive  tissues,  embryos,  and  mammary  glands  of 
individuals  from  natural  populations.  These  reports  have  provided  useful 
information  concerning  times  of  pregnancy,  lactation,  and  spermatogenesis. 
Thus,  a  basic  knowledge  of  reproductive  cycles  for  a  number  of  the  phyl¬ 
lostomatid  bats  has  been  accumulated  (for  review,  see  Wilson,  this  volume). 

However,  there  have  been  few  microscopic  studies  of  reproduction  and 
embryological  development  in  the  Phyllostomatidae.  With  respect  to  the 
details  of  the  embryology  of  these  bats,  only  seven  species  representing 
five  genera  have  been  studied  microscopically.  Some  of  these  works  are 
based  on  tissues  collected  from  natural  populations;  others,  on  tissues  from 
laboratory  colonies. 

This  paper  reviews  the  data  now  available  on  the  embryology  of  the 
Phyllostomatidae.  In  order  to  facilitate  this  presentation,  developmental 
events  will  serve  as  major  subdivisions,  and,  within  these  subdivisions,  the 
data  available  on  the  various  species  will  be  presented.  The  subdivisions  to 
be  considered  are  ovulation,  fertilization,  preimplantation  embryonic 
development,  implantation,  postimplantation  embryonic  development,  and 
placentation. 


Ovulation 

Macrotus  californicus. — Studies  of  M.  californicus  indicate  that  ovulation 
is  from  the  right  ovary  only  (even  though  both  ovaries  develop  Graafian 
follicles),  and,  typically,  that  only  one  ovum  is  released.  It  is  not  known  if 
ovulation  is  spontaneous  in  Macrotus  (Bradshaw,  1961). 

Glossophaga  soricina. — Ovulation  in  G.  soricina  may  occur  from  either 
ovary,  and  there  is  a  tendency  for  it  to  alternate  between  the  two.  Ovulation  is 
spontaneous  and  usually  only  one  ovum  is  released  per  cycle.  Menstruation 
occurs  in  G.  soricina  and  ovulation  takes  place  at,  or  very  close  to,  the  time 
of  menstruation  (Hamlett,  1935;  Rasweiler,  1972). 

Carollia  perspicillata,  C.  brevicauda,  and  Desmodus  rotundus. — Ovulation 
in  these  three  species  is  basically  the  same  as  in  G.  soricina.  However,  it  is 
not  known  if  ovulation  is  spontaneous.  Menstruation  in  Carollia  and  Desmodus 
is  similar  to  that  of  G.  soricina. 

Artibeus  lituratus. — Ovulation  in  Artibeus  lituratus  may  occur  from  either 
ovary  (Tamsitt  and  Valdivieso,  1963,  1965). 


379 


380 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fertilization 

Because  there  is  no  evidence  for  sperm  storage  in  the  female  reproductive 
tracts  of  phyllostomatid  bats,  it  appears  that  fertilization  occurs  shortly  after 
copulation.  Hence,  the  phenomenon  of  delayed  fertilization  that  has  been 
observed  in  some  of  the  Vespertilionidae  (Wimsatt,  1942)  has  not  been  reported 
in  any  of  the  Phyllostomatidae. 

Preimplantation  Embryonic  Development 

Macrotus  californicus. — Studies  on  M.  californicus  have  revealed  the 
sequence  of  events  prior  to  implantation;  however,  the  timing  of  these  events 
has  not  been  determined  (Bleier,  1975a).  Development  to  a  blastocyst 
occurs  in  the  oviduct  and  was  predicted  to  require  10  to  20  days  (Bleier,  1975/?). 
Embryonic  development  follows  the  pattern  typical  for  other  therian  mammals. 
There  is  no  information  concerning  the  loss  of  the  zona  pellucida  in  Macrotus. 

Glossophaga  soricina. — In  studies  of  a  laboratory  colony,  Rasweiler 
(1972)  was  able  to  time  the  sequence  of  events  in  embryonic  growth  of  G. 
soricina.  The  two-celled  stage  of  development  is  attained  by  day  2  or  3  post¬ 
ovulation.  The  eight-celled  stage  is  reached  by  days  5  to  7;  the  32-celled  stage, 
by  day  8;  the  blastocyst  stage,  by  day  10.  Compared  to  development  in  other 
mammals,  cleavage  rate  in  Glossophaga  is  slow.  The  zona  pellucida  is  usually 
lost  on  day  1 2  or  1 3,  and,  prior  to  its  loss,  the  embryo  has  been  contained  within  the 
ampulla  of  the  oviduct.  Upon  loss  of  the  zona  pellucida,  the  embryo  is  located  in 
the  intramural  uterine  cornu,  which  is  the  site  of  implantation.  There  is  no  evidence 
of  differentiation  of  germ  layers  during  this  preimplantation  period. 

Carollia  perspicillata  and  C.  brevicauda. — Cleavage  in  C.  perspicillata 
and  C.  brevicauda  also  proceeds  slowly.  De  Bonilla  and  Rasweiler  (1974) 
reported  that  the  first  blastocyst  was  observed  on  day  10  postcoitum.  Again, 
development  to  the  blastocyst  stage  and  loss  of  the  zona  pellucida  occurs  in  the 
oviduct.  Earliest  loss  of  the  zona  pellucida  was  day  10. 

Artibeus  jamaicensis. — The  only  information  available  on  early  embryonic 
development  in  A.  jamaicensis  was  reported  by  Fleming  (1971),  who  found 
two  reproductive  cycles  per  year  in  Panamanian  populations  and  noted  that 
the  embryo  reaches  the  blastocyst  stage  before  entering  the  uterus.  An  unusual 
feature  is  that  during  one  of  the  cycles  (August  to  March)  there  is  a  2.5-month 
period  of  delayed  embryonic  development.  During  this  period  of  retarded 
development,  the  only  noticeable  morphological  change  is  an  increase  in  the 
size  of  the  blastocyst. 

Desmodus  rotundus. — Slow  cleavage  also  is  characteristic  of  D.  rotundus. 
Quintero  and  Rasweiler  (1974)  observed  a  two-celled  embryo  as  late  as  day 
7  postcoitum  in  an  individual  from  a  laboratory  colony.  A  blastocyst  was  not  ob¬ 
served  until  day  15.  Loss  of  the  zona  pellucida  occurred  in  the  oviduct,  and  the 
earliest  date  of  this  loss  was  day  15.  Wimsatt  (1954)  noted  that  endoderm  dif¬ 
ferentiation  in  the  blastocyst  begins  while  the  blastocyst  is  still  in  the  oviduct. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


381 


Implantation 

Macrotus  californicus. — Several  reports  are  available  concerning  implantation 
in  M.  californicus.  Bradshaw  (1962)  noted  that  implantation  occurs  during 
early  gestation.  Later  studies  by  Bodley  (1974)  and  Bleier  (1975a,  19756) 
have  provided  more  details  concerning  the  process  in  Macrotus.  Central 
implantation  is  initiated  shortly  after  the  arrival  of  the  blastocyst  into  the  uterus. 
Early  stages  are  characterized  by  a  deterioration  of  the  uterine  epithelium 
such  that  the  invading  trophoblast  comes  into  contact  with  the  basal  lamina  of 
the  uterine  epithelium.  Endoderm  differentiation  is  initiated  at  this  time. 
By  the  end  of  October,  implantation  has  progressed  to  the  point  that  the  entire 
uterine  epithelium  that  once  surrounded  the  embryo  has  now'  been  obliterated. 
The  trophoblast  is  largely  multilayered  at  this  time,  but  unilaminar  portions 
may  be  observed  in  the  abembryonic  regions.  Reichert’s  membrane  separates 
the  trophoblast  from  the  remaining  fetal  tissue  and  becomes  continuous 
throughout  the  embryonic  and  abembryonic  regions.  The  age  of  an  embryo  at 
this  stage  is  estimated  to  be  20  to  30  days  (Bleier,  19756).  By  mid  December, 
syncytiotrophoblast  has  differentiated;  there  is  considerable  proliferation 
of  the  syncytiotrophoblast  by  the  end  of  January.  At  this  time,  an  interstitial 
membrane  (presumptive  intrasyncytial  lamina)  is  conspicuous  between  the 
maternal  tissue  and  the  syncytiotrophoblast.  Reichert’s  membrane,  which 
reaches  its  greatest  thickness  in  late  January,  disappears  by  mid  February. 
Endoderm  completely  surrounds  the  yolk  sac  cavity  at  this  stage.  By  mid 
February  all  the  layers  that  comprise  the  definitive  placenta  are  present 
(Bodley,  1974;  Bleier,  19756). 

Glossophaga  soricina. — Implantation  in  G.  soricina  is  initially  central  and 
secondarily  interstitial  (Rasweiler,  1974).  Rasweiler  (1974)  divided  this 
process  of  implantation  into  eight  stages.  Stage  I  (12  to  14  days  postcoitum) 
blastocysts  resemble  ampullary  blastocysts;  however,  there  is  some  hypertrophy 
of  the  trophoblast  in  Stage  I  embryos.  The  uterine  epithelium  is  intact  but  at 
times  flattened.  The  blastocyst  is  oriented  such  that  the  inner  cell  mass  is 
toward  the  cephalic  side  of  the  blastocystic  cavity.  The  first  appearance  of 
endoderm  differentiation  is  at  this  stage.  Stage  II  blastocysts  (days  13  to  15) 
are  characterized  by  a  bilaminar  and  multilaminar  trophoblast  in  the  embryonic 
polar  region,  whereas  the  trophoblast  of  the  abembryonic  region  remains 
unilaminar.  Necrosis  of  the  maternal  epithelium  has  begun  in  the  bilaminar  and 
multilaminar  regions  and  the  trophoblast  has  penetrated  the  basal  lamina  of  the 
uterine  epithelium.  Stage  III  blastocysts  (days  14  to  16)  resemble  Stage  II  blas¬ 
tocysts,  but  the  uterine  epithelium  has  deteriorated  further.  In  some  areas,  the 
trophoblast  has  penetrated  to  the  maternal  basal  lamina.  Endoderm  is  clearly 
recognized  in  all  specimens  from  Stage  III.  Stage  IV  specimens  (days  15  to  17)  are 
characterized  by  complete  obliteration  of  the  uterine  luminal  epithelium  with 
encroachment  of  the  trophoblast  to  the  uterine  glands.  A  decidual  reaction 
first  appears  at  this  stage.  During  Stage  IV,  the  endoderm  and  inner  cell  mass 


T 


382  SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 

fill  almost  the  entire  space  of  the  blastocystic  cavity.  Solid  multilayered  masses 
of  endoderm  occur  on  the  ventral  side  of  the  inner  cell  mass,  and,  by  days 
16  to  17,  pockets  have  begun  to  develop  in  the  endoderm.  Endoderm  appears 
on  the  lateral  and  dorsal  surfaces  of  the  inner  cell  mass.  Stage  V  (days  16  to  21) 
is  recognized  by  the  presence  of  syncytiotrophoblast  in  the  region  of  the 
embryonic  pole.  Cytotrophoblast  at  this  stage  is  present  outside  of  the 
syncytiotrophoblast,  in  addition  to  its  position  inside  the  syncytiotrophoblast, 
and  in  some  regions  has  penetrated  the  glandular  epithelium.  The  fluid-filled 
pockets  in  the  endoderm  are  more  pronounced,  and  in  one  specimen  had 
coalesced  to  form  a  unilocular  condition.  By  Stage  VI  (days  20  to  22)  and 
Stage  VII  (days  23  to  25),  the  syncytiotrophoblast  has  proliferated  further 
and  has  begun  to  penetrate  the  decidua  basalis.  There  is  an  increase  in  vascular 
lacunae  and  a  decrease  in  maternal  endothelium  in  Stage  VII  individuals. 
A  lamina  that  is  probably  an  extension  of  the  abembryonic  portion  of  Reichert’s 
membrane  is  interposed  between  the  inner  cell  mass  and  the  endoderm  dorsal 
and  lateral  to  the  inner  cell  mass.  Coalescence  of  the  pockets  in  the  endoderm 
has  continued  so  that  most  embryos  are  unilocular.  In  Stage  VIII  (days  26  to 
30),  the  cytotrophoblast  has  penetrated  deep  into  the  syncytiotrophoblast. 
During  this  stage,  the  intrasyncytial  lamina  is  observed  and  significant  quantities 
of  maternal  blood  in  the  labyrinth  first  appear.  Amniogenesis  by  cavitation 
has  begun  at  this  stage.  By  day  32,  differentiation  of  ectoderm  has  been 
initiated,  and  thinning  of  the  roof  of  the  amnion  has  begun.  The  endoderm  and 
Reichert’s  membrane,  in  the  region  of  the  embryonic  pole,  have  disappeared. 
The  fate  of  Reichert’s  membrane  is  currently  unknown. 

Carollia  perspicillata  and  C.  brevicauda. — Little  is  known  about  implantation 
in  Carollia.  De  Bonilla  and  Rasweiler  (1974)  found  that  the  site  of  implantation 
in  C.  perspicillata  and  C.  brevicauda  is  similar  to  that  reported  for  G.  soricina; 
that  is,  implantation  occurs  in  the  segment  between  the  end  of  the  oviduct 
and  the  main  cavity  of  the  uterus. 

Artibeus  jamaicensis. — The  only  report  on  A.  jamaicensis  is  that  of  Fleming 
(1971).  Implantation  is  similar  to  that  observed  for  Glossophaga  soricina  and 
Desmodus  rotundus,  including  “(i)  precocious  development  of  the  blastocyst, 
which  by  the  time  it  reaches  the  uterus,  has  differentiated  into  a  trophoblast 
thickened  at  the  embryonic  pole  and  an  embryonic  cell  mass .  .  .  and  (ii) 
implantation  that  is  interstitial  and  cytolytic.” 

Desmodus  rotundus. — The  only  observations  of  implantation  in  D. 
rotundus  were  reported  by  Wimsatt  (1954):  implantation  is  “cytolytic  and 
completely  interstitial,”  occurring  antimesometrially  in  the  middle  of  the  uterine 
cornu  and  on  the  same  side  as  is  the  ovary  from  which  ovulation  occurred. 
During  early  implantation,  the  embryo  is  secured  to  the  uterus  only  in  the 
region  of  the  embryonic  cell  mass,  thereby  exposing  the  abembryonic  surface 
to  the  uterine  cavity.  The  trophoblast  near  the  embryonic  pole  is  multilaminar, 
whereas  the  trophoblast  associated  with  the  free  surface  (abembryonic)  is 
unilaminar.  Beneath  the  inner  cell  mass,  the  endoderm  has  hypertrophied; 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


383 


in  other  regions  it  remains  flattened.  Wimsatt  (1954)  also  observed  precocious 
formation  of  mesoderm,  but  Rasweiler  (1974)  speculated  that  this  may 
actually  be  endoderm. 

In  a  second,  older  specimen,  Wimsatt  (1954)  noted  that  implantation  was 
complete.  By  this  stage,  the  embryo  is  completely  embedded  in  the  endometrium, 
and  the  trophoblast  is  multilayered  in  the  embryonic  region  but  still  largely 
unilaminar  in  the  abembryonic  region.  In  both  specimens,  there  is  a  marked 
decidual  reaction  but  it  is  most  pronounced  in  the  older  specimen.  Amniogenesis 
is  accomplished  by  cavitation. 

Postimplantation  Embryonic  Development 

Macrotus  californicus. — Embryonic  growth  in  M.  californicus  to  the  end 
of  implantation  is  slow.  Fertilization  in  Macrotus  most  often  occurs  during 
October,  and,  by  the  end  of  implantation  (mid  February),  amniogenesis  by 
cavitation  has  begun.  Therefore,  the  embryo  requires  approximately  four 
months  to  reach  the  embryonic-disc  stage  (Bleier,  1975  a).  Growth  accelerates 
during  March,  and  embryos  at  the  limb-bud  stage  (crown-rump  length 
approximately  4.5  millimeters)  of  development  are  observed.  Embryonic 
growth  continues  at  a  rapid  rate,  and  most  parturitions  occur  during  June. 
Growth  and  differentiation  of  the  embryonic  tissues  and  organs,  following  the 
period  of  slow  development,  are  similar  to  the  pattern  that  has  been  described 
for  other  therian  mammals. 

Glossophaga  soricina. — Hamlett  (1935)  described  the  embryonic 
growth  in  G.  soricina  following  implantation.  His  description  included  a 
discussion  of  the  primitive  streak  and  mesoderm  formation.  Primary  mesoderm 
is  formed  early;  however,  Rasweiler  (1974)  provided  evidence  that  this 
“primary  mesoderm”  is  most  likely  endoderm.  Formation  of  secondary 
mesoderm  (that  derived  from  the  primitive  streak)  and  subsequent  primitive 
streak  activity  are  similar  to  that  of  any  typical  mammal.  By  the  six-somite 
stage  the  coelom  is  present  (but  absent  at  the  medullary-fold  stage)  and  the 
mesoderm  has  split  into  splanchnic  and  somatic  layers.  The  yolk  sac  remains 
large,  but  the  yolk  stalk  disappears  before  the  2.5-millimeter  stage.  There  is  no 
evidence  of  the  allantois  in  the  six-somite  specimen  (length  is  one  millimeter 
from  head  fold  to  end  of  primitive  streak),  but  by  the  time  the  embryo  reaches 
2.5  in  length,  the  allantois  has  attained  its  maximum  relative  size. 

Placentation 

Macrotus  californicus. — Bradshaw  (1961)  noted  that  the  definitive 
placenta  in  M.  californicus  is  hemochorial.  Recently,  Bodley  (1974)  used 
electron  microscopic  techniques  that  revealed  the  definitive  placenta  to  be 
hemodichorial.  Development  of  the  placenta  is  such  that  it  is  large  enough  to 
be  readily  visible  with  the  naked  eye  by  late  March.  At  this  time,  reduction  of 
the  cytotrophoblast  to  a  single  cell  layer  begins  and  syncytial  blocks 


384 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


(derivatives  of  the  syncytiotrophoblast)  replace  the  maternal  endothelium 
(Bodley,  1974;  Bleier,  1975  b).  Changes  from  March  to  June  involve  maturation 
of  the  placenta,  but  there  is  no  change  in  the  number  of  cell  layers.  The  layers 
of  the  hemodichorial  placenta  in  Macrotus  include  syncytial  blocks,  intrasyncytial 
lamina,  syncytiotrophoblast,  cytotrophoblast,  fetal  basal  lamina,  and  fetal  capillary 
endothelium  (Bodley,  1974). 

Glossophaga  soricina. — Hamlett  (1935)  and  Rasweiler  (1974)  classified 
the  placenta  in  G.  soricina  as  discoidal  and  hemochorial,  and  Rasweiler  (1974) 
indicated  that  formation  was  rapid.  There  is  an  interstitial  lamina  present, 
but  its  origin  is  uncertain — Rasweiler  (1974)  suggested  that  it  is  derived  from 
the  trophoblast.  The  trophoblast  differentiates  into  cytotrophoblast  and 
syncytiotrophoblast;  however,  the  cytotrophoblast  disappears  by  midgestation. 
In  addition,  the  walls  and  endothelium  of  the  maternal  blood  vessels  are  eroded 
(Hamlett,  1935)  so  that  there  are  three  cell  layers  that  separate  the  fetal  and 
maternal  blood  streams.  These  layers  are  fetal  endothelium,  loose  mesenchyme, 
and  syncytiotrophoblast. 

Carollia  perspicillata. — Little  is  known  concerning  the  placenta  in  C. 
perspicillata.  Wimsatt  (1958)  noted  that  the  placenta  is  discoidal  and 
endotheliochorial.  Also,  he  implied  that  the  cytotrophoblast  does  not  persist 
to  the  end  of  gestation.  There  is  a  conspicuous  interstitial  membrane  between 
the  syncytiotrophoblast  and  maternal  endothelium.  However,  this  observation 
was  made  by  using  light  microscopy.  Recent  studies  indicate  that  other 
phyllostomatid  bats  have  a  hemodichorial  type  placenta  and  that  the  “maternal 
endothelium”  is  actually  syncytiotrophoblast  (Bjorkman  and  Wimsatt,  1968; 
Rasweiler,  1974;  Bodley,  1974;  Bleier,  1975b).  Therefore,  it  would  not  be 
surprising  if  it  were  determined  that  the  “maternal  endothelium”  in  the  placenta 
were  syncytiotrophoblast.  If  this  were  true,  and  if  the  cytotrophoblast  is 
lost,  then  the  placenta  of  Carollia  would  be  a  hemochorial  type.  Further 
investigations  are  needed  to  confirm  the  type  of  placental  barrier  characteristic 
of  Carollia. 

Artibeus  jamaicensis. — Wislocki  and  Fawcett  (1941)  stated  that  the  placenta 
is  discoidal  and  hemochorial. 

Desmodus  rotundus. — Initial  reports  indicated  that  the  placenta  in  D.  rotundus 
is  discoidal  and  endotheliochorial  (Wimsatt,  1954,  1958).  However,  by  using 
electron  microscopic  methods,  Bjorkman  and  Wimsatt  (1968)  concluded 
that  the  definitive  placenta  is  hemodichorial,  but  in  earlier  stages  before  the 
loss  of  the  maternal  endothelium  it  is  endotheliochorial.  Thus,  the  definitive 
placenta  consists  of  the  following  layers:  intrasyncytial  lamina,  syncytiotropho¬ 
blast,  cytotrophoblast,  a  thick  basement  membrane,  mesenchyme,  and  fetal 
endothelium. 


Summary  and  Conclusions 

From  the  data  summarized  in  this  paper,  several  trends  can  be  seen  in  the 
embryology  of  the  Phyllostomatidae.  In  general,  ovulation  may  occur  from 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


385 


either  ovary,  except  in  Macrotus  californicus,  and  fertilization  follows 
immediately  after  ovulation  and  copulation.  Embryonic  development  to  the 
blastocyst  stage  appears  to  be  similar  to  that  reported  for  other  therian 
mammals;  however,  the  process  seems  to  be  considerably  slower  in  the 
phyllostomatid  bats  studied  thus  far.  Implantation  is  interstitial  except  in 
M.  californicus.  The  placenta  is  discoidal,  and  it  is  likely  that  the  placental 
barrier  is  either  hemodichorial  or  hemochorial. 

There  are  several  features  of  phyllostomid  embryology  that  should  stimulate 
further  investigations  of  the  species  reported  in  this  paper.  In  addition,  studies 
of  other  species  should  be  encouraged  for  they  might  reveal  embryological 
strategies  other  than  the  ones  presently  known.  Some  of  the  areas  deserving  the 
application  of  sophisticated  research  techniques  include  ovulation  from  only  the 
right  ovary  in  M.  californicus,  delayed  embryonic  development  in  M.  californicus 
and  Artibeus  jamaicensis,  the  length  of  gestation  in  Desmodus ,  and  menstruation 
and  interstitial  implantation  in  Glossophaga,  Carollia ,  and  Desmodus. 

Literature  Cited 

Bjorkman,  N.  H.,  and  W.  A.  Wimsatt.  1968.  The  allantoic  placenta  of  the  vampire 
bat  ( Desmodus  rotund  us  murinus):  a  reinterpretation  of  its  structure  based  on 
electron  microscopic  observations.  Anat.  Rec.,  162:83-98. 

Bleier,  W.  J.  1975a.  Early  embryology  and  implantation  in  the  California  leaf-nosed 
bat,  Macrotus  californicus.  Anat.  Rec.,  182:237-254. 

- .  1975ft.  Fine  structure  of  implantation  and  the  corpus  luteum  in  the  California 

leaf-nosed  bat,  Macrotus  californicus.  Unpublished  Ph.D.  dissertation, 
Texas  Tech  Univ.,  75  pp. 

Bodley,  H.  D.  1974.  Ultrastructural  development  of  the  chorioallantoic  placental 
barrier  in  the  bat  Macrotus  waterhousii.  Anat.  Rec.,  180:351-368. 

Bradshaw,  G.  VR.  1961.  A  life  history  study  of  the  California  leaf-nosed  bat,  Macrotus 
californicus.  Unpublished  Ph.D.  dissertation,  Univ.  Arizona,  89  pp. 

- .  1962.  Reproductive  cycle  of  the  California  leaf-nosed  bat,  Macrotus  californicus. 

Science,  136:645-646. 

De  Bonilla,  H„  and  J.  J.  Rasweiler,  IV.  1974.  Breeding  activity,  preimplantation 
development,  and  oviduct  histology  of  the  short-tailed  fruit  bat,  Carollia , 
in  captivity.  Anat.  Rec.,  179:385-404. 

Fleming,  T.  H.  1971.  Artibeus  jamaicensis:  delayed  embryonic  development  in 
a  Neotropical  bat.  Science,  171:402-404. 

Hamlett,  G.  W.  D.  1935.  Notes  on  the  embryology  of  a  phyllostomatid  bat.  Amer. 
J.  Anat.,  56:327-349. 

Quintero,  F.,  and  J.  J.  Rasweiler  IV.  1974.  Ovulation  and  early  embryonic 
development  in  the  captive  vampire  bat,  Desmodus  rotundus.  J.  Reprod. 
Fert.,  41:265-273. 

Rasweiler,  J.  J.,  IV.  1972.  Reproduction  in  the  long-tongued  bat,  Glossophaga 
soricina.  I.  Preimplantation  development  and  histology  of  the  oviduct.  J.  Reprod. 
Fert.,  31:249-262. 

- .  1974.  Reproduction  in  the  long-tongued  bat,  Glossophaga  soricina.  II.  Im¬ 
plantation  and  early  embryonic  development.  Amer.  J.  Anat.,  139:1-36. 

Tamsitt,  J.  R„  and  D.  Valdivieso.  1963.  Reproductive  cycle  of  the  big  fruit-eating 
bat,  Artibeus  lituratus  Olfers.  Nature,  198:104. 


386 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


- .  1965.  Reproduction  of  the  female  big  fruit-eating  bat,  Artibeus  lituratus 

palmarum,  in  Colombia.  Caribbean  J.  Sci.,  5:157-166. 

Wimsatt,  W.  A.  1942.  Survival  of  spermatozoa  in  the  female  reproductive  tract 
of  the  bat.  Anat.  Rec.,  83:299-307. 

- .  1954.  The  fetal  membranes  and  placentation  of  the  tropical  American  vampire 

bat  Desmodus  rotundus  murinus  with  notes  on  the  histochemistry  of  the  placenta. 
Acta  Anat.,  21:285-341. 

- .  1958.  The  allantoic  placental  barrier  in  chiroptera:  a  new  concept  of  its 

organization  and  histochemistry.  Acta  Anat.,  32:141-186. 

Wislocki,  G.  B.,  and  D.  W.  Fawcett.  1941.  The  placentation  of  the  Jamaican  bat 
(Artibeus jamaicensis  parvipes).  Anat.  Rec.,  81:307-317. 


ONTOGENY  AND  MATERNAL  CARE 


D.  G.  Kleiman  and  T.  M.  Davis 


Although  many  aspects  of  phyllostomatid  biology  have  received  increasing 
attention  in  recent  years,  there  is  still  a  dearth  of  information  on  the  growth  and 
behavioral  ontogeny  of  this  diverse  family  of  bats.  This  is  in  contrast  with 
studies  of  the  Vespertilionidae,  where  both  field  and  laboratory  investigations 
of  development  have  been  common,  although  by  no  means  numerous  (Jones, 
1967;  Pearson  et  ai,  1952;  Kleiman,  1969;  Orr,  1970;  Gould,  1971).  The 
lack  of  interest  in  chiropteran  ontogeny  is  discouraging  because  the  special 
adaptations  for  flight,  including  echolocation,  and  diverse  feeding  strategies 
should  provide  fertile  ground  for  developmental  studies,  as  Gould  (1970) 
has  pointed  out. 

In  this  chapter  we  will  attempt  to  review  some  aspects  of  ontogeny  in  the 
phyllostomatid  bats,  concentrating  on  growth  and  development  in  Carollia 
perspicillata,  which  we  have  studied  in  captivity.  Field  and  laboratory  observations 
of  other  species  will  be  included  where  they  are  available.  The  vampire  bat, 
Desmodus  rotundus,  is  the  only  other  phyllostomatid  for  which  detailed  in¬ 
formation  is  available  (Schmidt  and  Manske,  1973). 

The  colony  of  Carollia  perspicillata  was  originally  captured  in  Trinidad  in 
April  1972  and  maintained  at  Johns  Hopkins  University  for  six  months  by 
E.  Gould.  During  this  period,  several  births  occurred.  Sixteen  Carollia  were 
brought  to  the  National  Zoological  Park,  Washington,  D.C.,  in  October  1972. 
At  this  time,  one  female  had  a  small  infant;  a  second  female  gave  birth  three 
days  after  the  arrival  of  the  colony.  Both  young  were  reared.  Table  1  presents 
the  history  of  the  colony  between  January  1973  and  January  1974.  Three 
Glossophaga  soricina  (two  males,  one  female)  were  acquired  with  the  Carollia, 
of  which  one  adult  male  died  and  one  male  was  born.  Nine  Anoura  geoffroyi 
(four  males,  five  females)  also  were  received,  but  all  but  a  pair  died  within 
the  first  three  days.  No  breeding  of  Anoura  occurred. 

The  colony  was  housed  in  a  climate-controlled  room  measuring  approximately 
3  by  3  by  2.5  meters.  Temperatures  averaged  29 °C  (range  27  to  31  °C);  relative 
humidity,  70  per  cent  (range  50  to  80  per  cent).  A  light  cycle  of  12  hours  of 
light  to  12  hours  of  dark  was  used.  Two  wire  mesh  cages  with  wooden  frames 
and  burlap  covers  were  provided  for  roosts  in  an  elevated  position.  Several 
branches  were  placed  between  the  roosts  and  from  the  roosts  to  the  floor. 

Bats  were  fed  a  peach-nectar  mixture  developed  by  Rasweiler  and  De  Bonilla 
(1972)  for  nectarivorous  phyllostomatids,  although  there  is  evidence  that 
Carollia  also  feeds  on  insects  (Pine,  1972;  Ayala  and  D’ Alessandro,  1973). 
Water  was  available  ad  libitum,  as  were  ripe,  peeled  bananas  that  were 
suspended  from  branches.  Dishes  with  the  nectar  diet  were  placed  in  brackets 


387 


388 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Table  1. —  History  o/Carollia  perspicillata  colony  from  January  1973  to  January  1974. 


Males 

Females 

Total 

Number  of  original  adults 

6 

11 

17 

Number  of  births 

Number  of  deaths: 

17 

13 

30 

Adults 

0 

1* 

1 

Juveniles 

1 

5* 

6 

*One  mother  and  young  died  accidentally. 


attached  to  the  outside  of  the  roosts  so  that  bats  could  feed  while  in  flight  or 
while  hanging  on  the  roost. 

Bats  were  caught  with  butterfly  nets;  adults  initially  were  examined 
bimonthly  beginning  in  January  1973,  but  weekly  examinations  were  instituted 
in  April  1973.  Young  were  weighed  and  measured  every  two  to  four  days. 
Individuals  were  identified  by  a  number  punch  marked  on  the  wing  membrane 
(Bonaccorso  and  Smythe,  1972;  Kleiman  and  Davis,  1974).  Behavioral 
observations  and  retrieval  tests  were  conducted  at  irregular  intervals. 

Reproductive  Cycle 

After  the  two  births  in  October  1972,  there  were  three  birth  peaks  in 
Carollia:  February  1973,  June  and  July  1973,  and  November  and  December 
1973  (Table  2).  Known  interbirth  intervals  ranged  from  115  to  173  days. 
During  the  first  peak  of  parturition,  females  were  highly  synchronized — nine 
of  1 1  females  gave  birth  within  a  17-day  period.  The  births  were  more  scattered 


Table  2. — Dates  of  birth  and  interbirth  intervals  for  12  Carollia  perspicillata  females, 

between  January  1973  and  January  1974. 


No.  of 
females 

Birth  no.  1 

Birth  no.  2 

Birth  no.  3 

Interbirth 

interval 

(days) 

4 

12  Feb.  73 

1  July  73 

25  Oct.  73 

138, 

116 

6 

26  Jan.  73 

29  May  73 

18  Nov.  73 

123, 

173 

7 

28  Feb.  73* 

23  June  73 

6  Dec.  73* 

115, 

165 

10 

20  Feb.  73* 

21  June  73* 

121 

11 

14  Feb.  73* 

16 

5  Mar.  73 

21  July  73* 

10  Dec.  73 

138, 

142 

17 

16  Feb.  73* 

3  Aug.  73* 

168 

19 

12  Feb.  73 

24  June  73 

8  Nov.  73* 

132, 

137 

20 

18  Feb.  73* 

1  July  73 

15  Nov.  73* 

133, 

137 

25 

22  Feb.  73* 

12  July  73* 

7  Dec.  73* 

140, 

148 

26 

22  Feb.  73* 

20  June  73* 

10  Dec.  73 

118, 

173 

35 

14  Jan.  74* 

"■Indicates 

accurate  birth  date.  Other 

dates  are  estimated 

and  parturition  might 

have  occurred  a 

maxi- 

mum  of  three  days  earlier. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


389 


Pre-partum  Post-partum 

Weeks 

Fig.  1. — Average  weights  and  ranges  of  weights  in  pre  and  postpartum  Carollia 
perspicillata,  based  on  25  births  of  1 1  females. 


during  the  succeeding  two  parturition  periods.  The  shortest  interbirth  intervals 
ranged  from  115  to  123  days  (N  =  5).  Rasweiler  and  De  Bonilla  (1972) 
found  an  implanting  blastocyst  in  a  female  killed  21  days  postpartum, 
suggesting  that  estrus  may  occur  shortly  after  parturition.  If  an  immediate 
postpartum  heat  occasionally  occurs,  the  gestation  period  for  Carollia  perspicillata 
may  be  approximately  115  to  120  days.  The  single  Glossophaga  soricina  female 
gave  birth  in  March  1973  and  did  not  become  pregnant  again  for  a  full  year. 

A  total  of  30  Carollia  young  were  born  (see  Table  1),  of  which  24  survived 
through  weaning.  No  females  aborted  nor  were  any  infants  rejected  after  birth. 
The  majority  of  juvenile  deaths  occurred  at  weaning,  and  at  least  four  of  these 
might  have  resulted  from  feeding  on  spoiled  food  or  a  disfunction  in  the 
humidity  control,  which  caused  a  rapid  drop  in  humidity  in  the  flight  room. 
Adult  losses  were  limited  to  a  single  female  and  her  young,  which  died 
accidentally. 


390 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Females  gained  approximately  one-third  of  their  initial  weight  during 
pregnancy  (see  Fig.  1).  Average  weight  during  the  last  week  of  pregnancy  was 
22.9  grams  as  compared  with  17.3  grams  during  the  first  week  postpartum. 
During  the  final  weeks  of  pregnancy,  females  were  reluctant  to  fly  and 
maneuvered  less  efficiently  when  they  flew.  Fetuses  were  palpable  from 
about  five  to  six  weeks  before  birth,  and  were  in  a  transverse  position. 

The  nipples  of  pregnant  females  were  not  obvious  prior  to  birth,  but  within 
two  days  of  parturition  the  surrounding  fur  had  been  shed  and  the  mammary 
region  had  become  pink  in  color.  Thick  milk  could  be  expressed  from  the 
nipples  up  until  approximately  33  days  postpartum  (range  21  to  49  days). 
Thereafter,  the  milk  began  to  thin,  but  fluid  could  be  expressed  until 
approximately  56  days  after  birth  (range  42  to  72  days).  The  area  around 
the  nipples  began  to  assume  a  darker  pigmentation  and  the  fur  began  to  re¬ 
appear  from  48  days  postpartum  (range  37  to  66  days);  however,  the  mammary 
region  did  not  assume  prepartum  condition  until  72  days  postpartum  (range 
64  to  87  days).  From  these  observations,  it  would  appear  that  heavy  lactation 
continues  for  slightly  over  one  month  after  birth,  but  females  continue  to 
produce  milk  until  approximately  1.5  to  2  months  postpartum. 

Data  available  for  length  of  lactation  in  other  phyllostomatids  indicate  a 
lactation  period  of  one  to  two  months  (Jenness  and  Studier,  1976).  In  the 
vampire  bat,  Desmodus,  nursing  may  continue  for  nine  months  although 
weaning  is  initiated  at  three  (Schmidt  and  Manske,  1973).  In  Macrotus  and 
Leptonycteris ,  lactation  continues  for  one  month  and  four  to  eight  weeks, 
respectively  (see  Jenness  and  Studier,  1976).  A  single  Glossophaga  soricina 
female  in  our  colony  continued  lactating  for  approximately  two  months. 

Maternal  Care 

No  births  were  observed  in  Carollia  although  females  were  seen  eating 
placentas  and  licking  newborn  young.  The  umbilical  cord  was  rarely  severed 
at  the  base,  but  usually  dried  up  and  fell  off  within  a  day  following  birth. 

Parturition  has  been  described  for  Stenoderma  rufum  (Tamsitt  and 
Valdivieso,  1966b),  Artibeus  lituratus,  Glossophaga  soricina,  Vampyrops 
helleri  (Tamsitt  and  Valdivieso,  1965),  and  Choeronycteris  mexicana 
(Barbour  and  Davis,  1969).  In  all  species,  parturition  occurred  in  the  normal 
head-down  position;  this  seems  to  be  typical  of  phyllostomatids  but  rare  in 
vespertilionids  (Wimsatt,  1960),  except  for  Nyctalus  noctula  (Kleiman,  1969). 

In  the  species  observed  by  Tamsitt  and  Valdivieso  (1965,  1966b),  a  head 
presentation  was  found.  Placentophagia  has  not  been  reported  for  the  above- 
mentioned  species,  nor  for  Desmodus  (Schmidt  and  Manske,  1973). 

During  the  first  few  days,  young  Carollia  were  carried  parallel  to  the  mother’s 
body  and  held  under  the  wing.  Thereafter,  the  typical  carrying  position,  both 
at  rest  and  in  flight,  was  cross-wise  on  the  mother’s  ventral  surface,  just  posterior 
to  the  throat.  Carollia  infants  (up  to  14  days)  were  rarely  observed  hanging 
alone.  Young  attached  themselves  primarily  with  the  mouth  and  hind  feet; 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


391 


the  wings  were  tightly  closed  and  partially  covered  the  infant’s  body.  Claws  on 
the  thumbs  were  not  used  for  clinging  because  the  distal  portion  of  the  forearm 
was  pressed  tightly  against  and  covered  the  infant’s  head  and  ears.  Young  removed 
from  the  mother’s  nipple  occasionally  remained  in  this  carrying  posture  for 
several  seconds,  even  when  placed  on  their  back.  The  cross-wise  carrying 
posture  was  also  seen  in  our  individual  of  Glossophaga  soricina,  Desmodus 
(see  fig.  2  in  Schmidt  and  Manske,  1973),  and  might  be  present  in  Choeronycteris 
(see  fig.  8  in  Barbour  and  Davis,  1969).  It  appears  to  be  an  adaptation  for  carrying 
young  while  the  female  is  flying.  For  the  first  10  days,  captive  young  of  Artibeus 
were  reported  (Novick,  1 960)  to  hang  head  down  under  the  mother’s  wing  with  the 
hindfeet  around  the  mother’s  thigh. 

Carollia  mothers  preferred  to  hang  freely  from  a  horizontal  ceiling  when 
carrying  attached  young.  Thus,  it  was  impossible  for  infants  to  be  attached 
simultaneously  to  the  nipple  and  support  themselves  by  the  hind  feet  until 
they  were  about  half  the  size  of  the  mother.  Young  were  capable  of  hanging 
from  the  ceiling  by  the  age  of  18  days,  but  still  remained  attached  to  a  nipple. 
Similar  observations  were  made  of  a  young  Glossophaga.  In  Desmodus,  young 
do  not  support  themselves  until  at  least  two  weeks  of  age  (Schmidt  and  Manske, 
1973). 

From  our  observations,  it  appeared  that  resting  Carollia  females  supported 
the  bulk  of  their  infant’s  weight  for  at  least  14  days.  An  added  advantage  to 
the  cross-wise  carrying  position  assumed  by  the  young,  other  than  providing 
balance,  was  that  they  did  not  need  to  readjust  their  position  when  a  female 
flew.  Young  were  last  observed  attached  to  the  mother  approximately  23.5 
days  postpartum  (range  19  to  31;  N  =  15),  when  they  were  approximately 
57  per  cent  of  the  mother’s  weight. 

Because  we  were  unable  to  observe  the  bats  without  disturbing  them, 
especially  at  night,  we  do  not  know  whether  females  foraging  in  the  wild  carry 
their  young  or,  if  they  do,  for  how  long.  One  1 1  -day-old  young  was  seen 
hanging  alone  next  to  its  mother  approximately  45  minutes  after  the  lights 
went  out,  but  the  infant  attached  to  the  nipple  and  moved  back  into  a  cross-wise 
carrying  position  immediately  after  we  entered  the  room.  The  mother  flew 
as  soon  as  the  young  attached.  This  suggests  that  mothers  may  detach  from  the 
young  at  night,  but  we  had  no  evidence  that  young  were  ever  left  in  a  creche. 
Mothers  with  attached  young  were  more  reluctant  to  fly  when  disturbed  than 
were  unencumbered  bats  but  did  so,  nevertheless,  and  seemed  able  to 
maneuver  efficiently. 

Observations  of  development  in  a  single  young  Glossophaga  soricina  were 
similar  to  those  for  Carollia.  The  young  was  last  seen  attached  to  the  mother 
when  it  was  20  days  old. 

Both  from  our  Carollia  observations  and  some  field  reports,  it  appears 
as  though  some  species  of  phyllostomatid  bats  commonly  carry  their  young 
and,  unlike  vespertilionids  (see  Fenton,  1969;  Davis,  1970),  do  not  leave  them 
in  creches. 


392 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Felten  (in  Pine,  1972)  apparently  netted  a  Carollia  perspicillata  with  a 
half  grown  young,  and  Tamsitt  and  Valdivieso  (1963  a)  caught  lactating 
Artibeus  lituratus  and  Glossophaga  sorcina  carrying  young  in  the  vicinity  of 
fruit  trees  where  presumably  they  were  foraging.  One  A.  lituratus  female 
carried  a  young  53.8  per  cent  of  her  weight  (Tamsitt  and  Valdivieso,  1965). 
Mumford  and  Zimmerman  (1964)  reported  netting  lactating  Choeronycteris 
mexicana  with  attached  young  at  a  distance  of  approximately  200  yards  from 
the  main  daytime  roost.  Bradshaw  (1961)  captured  a  female  Macrotus 
californicus  in  a  roost  carrying  a  young  weighing  57  per  cent  of  her  weight; 
Cockrum  (in  Davis,  1970)  observed  female  Leptonycteris  sanborni  moving 
young  within  a  cave  as  well  as  carrying  advanced  young  to  a  previously 
abandoned  roost.  Schmidt  and  Manske  (1973)  indicated  that  Desmodus 
females  can  carry  young  up  to  eight  weeks  old.  A.  M.  Greenhall  (personal 
communication)  has  observed  Desmodus  females  with  attached  young  of 
unknown  age  feeding  on  cattle;  however,  these  bats  were  similar  in  size  to 
young  that  he  had  observed  crawling  around  in  roosts  without  the  mothers. 
These  young  were  not  newborn  and  might  have  been  approaching  weaning 
age. 

Observations  discussed  above  suggest  that  phyllostomatids  may  carry 
attached  young  of  an  advanced  age.  Whether  females  forage  with  the  young  or 
simply  move  them  from  roost  to  roost  remains  to  be  determined.  Certainly, 
except  for  Macrotus  waterhousii  (Goodwin,  1970),  Leptonycteris  sanborni 
(Hoffmeister,  1959),  and  Phyllostomus  hastatus  (J.  Bradbury,  personal 
communication),  one  does  not  find  reports  of  creches  of  infants  in  phyllostomatids, 
although  lactating  females  may  roost  colonially  and  segregate  themselves 
from  males.  Bradbury  (personal  communication)  suggested  that  female 
Carollia,  for  example,  may  move  their  babies  from  the  day  roost  to  a  night 
roost  prior  to  foraging,  which  may  partly  account  for  the  well-developed 
tendency  to  carry  young  in  captivity. 

Retrieval  of  young  Carollia  was  observed  under  several  experimental 
conditions.  Mothers  and  young  were  released  into  a  small  holding  cage  after 
being  weighed  and  measured;  typically,  they  reestablished  contact  within 
30  minutes  to  an  hour  (that  is,  before  being  released  into  the  flight  room). 
On  several  occasions,  young  were  deliberately  separated  and  hung  on  the 
outside  of  the  roost,  after  which  time  the  other  bats  were  released  into  the 
flight  room.  Several  different  bats  would  fly  past  hanging  infants,  pausing 
briefly  to  hover,  as  though  to  inspect  the  young.  Usually,  a  juvenile  was  inspected 
several  times  (both  by  its  mother  and  other  bats)  before  the  mother  would 
alight  above  her  offspring  and  crawl  down  to  it. 

Juveniles  that  were  too  young  to  fly  were  never  observed  attempting  to  regain 
contact  with  their  mother  by  climbing  higher  on  the  roost.  Normally,  they 
hung  motionless  until  the  mother  made  tactile  contact  with  them.  Audible 
vocalizations  (ultrasonic  calls  were  given  by  the  mother  and  young,  Gould, 
1975)  were  not  heard  nor  did  the  infant  reveal  much  sign  of  disturbance. 
Licking  of  the  young  by  the  mother  usually  accompanied  retrieval,  especially 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


393 


before  the  mother  flew  again.  The  latency  to  retrieve  was  highly  variable 
in  the  females,  ranging  from  two  to  30  minutes.  The  age  of  the  young  did  not 
seem  to  affect  this  latency  because  infants  between  one  and  three  days  old 
were  retrieved  within  two  to  22  minutes. 

Mothers  clearly  recognized  their  own  offspring;  we  never  caught  a  female 
with  an  alien  young  attached  to  her.  Moreover,  mothers  and  young  retained 
an  association  (roosted  near  each  other)  long  after  weaning.  One  Carollia 
mother  and  daughter  were  regularly  caught  together  until  the  daughter  was 
five  months  old,  about  a  week  prior  to  the  next  birth. 

Development  of  Young 

Carollia  are  born  in  an  advanced  state,  with  the  eyes  open  (Fig.  2). 
Neonates  are  fully  furred  on  the  dorsum,  and  the  more  sparsely  furred  venter 
and  muzzle  become  covered  within  two  to  three  days  after  birth.  The  dark 
brown  juvenile  pelage  is  complete  by  day  7  to  10. 

Of  the  neonatal  phyllostomatids  observed,  Macrotus,  Leptonycteris 
(Gould,  1975),  Carollia,  Glossophaga  (this  study  and  Klfma  and  Gaisler, 
1968),  Choeronycteris  (Mumford  and  Zimmerman,  1964),  and  Artibeus 
(Tamsitt  and  Valdivieso,  1966a)  are  born  well  furred.  Desmodus  (Schmidt 
and  Manske,  1973;  Gould,  1975),  Phyllostomus  discolor  (Klima  and  Gaisler, 
1968),  and  P.  hastatus  (Gould,  1975)  are  sparsely  furred  at  birth. 

Eyes  are  open  at  birth  in  Carollia  (this  study),  Artibeus  (Tamsitt  and 
Valdivieso,  1966a),  Desmodus  (Schmidt  and  Manske,  1973;  Gould,  1975), 
Macrotus  (Gould,  1975),  and  Phyllostomus  hastatus  (Gould,  1975).  Only 
Leptonycteris  and  Phyllostomus  discolor  have  been  reported  (Tamsitt  and 
Valdivieso,  1963  a)  to  have  the  eyes  closed  at  birth. 

Carollia  neonates  were  active  from  birth  and  when  handled  would 
squirm,  try  to  crawl  away,  and  often  vocalize.  This  contrasted  with  their 
behavior  in  the  flight  room  during  retrieval  tests  when  they  hung  motionless 
on  the  bat  roost.  The  increased  activity  might  have  been  caused  by  the 
temperature  of  the  room  in  which  weights  and  measurements  were  taken, 
which  was  cooler  than  was  the  flight  room.  Gould  (1975)  stated  that  the  young 
of  Desmodus,  Phyllostomus  hastatus,  and  Leptonycteris  sanborni  are  active 
during  reunions  with  the  mother,  whereas  those  of  Macrotus  californicus 
are  passive. 

C.  perspicillata  young  are  born  with  a  complete  set  of  22  deciduous  teeth, 
with  the  formula  di  2-2/2 -2,  dc  1 -1/1-1,  dpm  3-3/2-2  =  22.  A  comparison 
of  preserved  skulls  from  the  U.S.  National  Museum  with  living  neonates  suggests 
that  only  16  of  the  22  deciduous  teeth  are  functional.  The  four  lower  incisors, 
barely  penetrating  the  gingivum,  disappear  several  days  after  birth,  and  the  first 
upper  deciduous  premolars  are  not  even  visible  in  live  specimens.  Lower  deciduous 
premolars  are  simple,  highly  reduced  spicules,  undifferentiated  in  width  from  root 
to  crown.  The  second  and  third  upper  premolars,  although  more  prominent  than 
the  lower  ones,  are  tiny  pegs  that  taper  to  a  fine  point  at  the  crown.  The  second  milk 


394 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fig.  2. — Neonate  of  Carollia  perspicillata  on  the  day  of  birth.  Note  that  the 
eyes  are  open,  and  the  animal’s  dorsum  is  fully  furred.  The  venter  typically  has  only 
sparse  fur. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


395 


Fig.  3. — The  loss  of  deciduous  teeth  in  juvenile  Carollia  perspicillata.  Observations 
within  a  given  time  period  may  include  the  same  individual.  (Symbols  are  closed  circles, 
upper  outer  incisors;  open  circles,  upper  inner  incisors;  closed  triangles,  lower  canines; 
open  triangles,  upper  canines;  closed  squares,  lower  premolars;  and  open  squares, 
upper  premolars). 

premolar  is  weakly  recurved.  Lower  canines  are  slender,  mildly  recurved  spicules 
that  gradually  taper  to  a  point.  The  upper  canines  and  upper  outer  incisors  are  the 
largest,  most  strongly  recurved  stylettes;  also,  they  are  retained  longest.  Upper 
inner  incisors  are  bifid  at  the  distal  extremity. 

The  comparative  rate  of  loss  of  deciduous  teeth  is  represented  in  Fig.  3. 
Lower  deciduous  premolars  are  lost  during  the  first  two  weeks  postpartum. 
Lower  canines,  upper  premolars,  and  upper  inner  incisors  are  shed  next.  The 
upper  canines  and  upper  outer  incisors  are  retained  until  one  month  postpartum, 
the  last  milk  tooth  being  lost  at  34  days  postpartum.  The  permanent  dentition 
of  the  upper  jaw  emerged  first.  By  day  22  postpartum,  one-third  of  the 
permanent  teeth  had  emerged;  by  day  26,  two-thirds;  and  by  day  31,  all  were 
present. 

Deciduous  upper  and  lower  canines  and  upper  outer  incisors  are  the  teeth 
primarily  used  to  attach  to  a  nipple.  Two  observed  perforations  in  a  female’s 
nipple  were  a  clear  result  of  the  upper  canines,  the  distances  between  the 
perforations  and  the  canines  both  measuring  2.6  millimeters.  Carollia 
resembles  Tonatia,  Mimon,  Chrotopterus,  Choeronycteris,  and  Phyllostomus 
in  that  the  upper  outer  incisors  are  more  prominent  than  the  upper  inner  ones. 


396 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


In  Macrotus,  Glossophaga,  and  Leptonycteris,  both  outer  and  upper  inner 
incisors  are  functional  (Phillips,  1971). 

In  general,  the  deciduous  dentition  of  most  phyllostomatids  is  reduced 
and  less  complex  than  that  of  vespertilionids  (Phillips,  1971;  Miller,  1907). 
This  seems  to  correlate  with  the  tendency  to  carry  attached  young  rather  than 
deposit  them  in  creches,  thus  suggesting  that  increased  complexity  in  the 
deciduous  dentition  of  vespertilionids  may  function  to  grasp  the  returning 
mother  (or  any  female  in  species  that  nurse  promiscuously)  rather  than  to  main¬ 
tain  a  hold  on  the  nipple  when  already  attached. 

The  development  of  flight  in  C.  perspicillata  was  investigated  by  periodically 
dropping  infants  and  juveniles.  Prior  to  day  14,  all  young  drop  straight  to  the 
ground,  with  the  wings  extended.  As  infants  approached  14  days  of  age,  they 
occasionally  flapped  their  wings  once  or  twice  as  they  fell.  Between  days  14  and 
16,  young  bats  began  flapping  the  wings  when  dropped,  but  could  not  maintain 
altitude  or  turn.  They  also  were  unable  to  land  and  often  collided  with  obstacles  or 
eventually  dropped  to  the  floor.  By  day  1 8,  they  could  maintain  (and  gain)  altitude, 
take  off  from  a  roosting  position,  turn,  and  avoid  obstacles.  However,  their  landing 
ability  was  poor,  and  they  often  landed  with  the  wings  extended.  Between  days  20 
and  23,  the  ability  to  land  upside-down  with  the  wings  folded  perfected,  and, 
after  day  24,  flight  development  essentially  was  complete.  Juveniles,  however, 
could  be  distinguished  from  adults  by  their  flight  patterns  for  several  weeks  more 
because  they  flew  more  slowly  and  erratically.  Juveniles  were  first  captured  inde¬ 
pendent  of  the  mother  on  an  average  of  27.6  days  (range  23  to  31,  N=  16)  after 
birth. 

There  is  little  information  available  on  flight  development  in  other  young 
phyllostomatids.  In  Desmodus,  young  achieve  flight  capability  at  eight  to  ten 
weeks  of  age  (Schmidt  and  Manske,  1973);  Novick  (1960)  reported  that  a 
young  Artibeus  began  to  fly  at  approximately  28  days  of  age.  A  single  juvenile 
Glossophaga  soricina  was  first  found  separate  from  its  mother  and  flying  at 
age  25  to  28  days. 

Neonates  of  Carollia  perspicillata  average  5.0  grams  at  birth  (range  4.1  to 
5.9;  N=  13),  which  is  28.4  per  cent  of  the  postpartum  weight  of  females. 
Initial  growth  in  weight  is  rapid  (Fig.  4),  but  juveniles  do  not  achieve  adult 
weight  until  10  to  13  weeks  of  age.  Forearm  length  at  birth  is  24.4  millimeters 
(range  22.4  to  27.5  mm;  N=  10),  and  forearm  growth  essentially  is  complete 
at  six  weeks  (Fig.  4).  At  approximately  24  days  of  age,  when  the  young  first 
begin  to  fly,  forearm  length  is  93.4  per  cent  and  weight  63.0  per  cent  of  that 
for  adults  (N=  10). 

Neonatal  and  postpartum  weights  and  measurements  are  not  available  for 
most  phyllostomatid  bats.  Table  3  presents  some  accurate  and  estimated 
neonatal  to  mother  weight  and  measurement  ratios  for  both  phyllostomatid 
and  vespertilionid  bats,  based  on  known  and  derived  data.  Weights  and 
measurements  were  taken  from  full-term  fetuses  and  nonlactating  females. 
Young- to-mother  weight  ratios  are  poor  for  comparative  purposes  because 
weights  tend  to  fluctuate  seasonally,  captive  and  field  weights  frequently 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


397 


Days  Weeks 

Age 


Fig.  4. — Increase  in  average  weight  (bottom)  and  forearm  length  (top)  for  Carollia 
perspicillaux.  A,  average  day  when  young  were  last  observed  attached  to  the  mother; 
B,  average  day  when  the  mother’s  milk  began  to  thin;  C,  average  day  when  milk  no  longer 
could  be  expressed  from  the  mother's  nipples.  These  averages  are  based  on  measurements 
from  17  individuals  (8  females,  9  males)  of  known  age.  The  open  squares  indicate  the 
mean  weight  and  forearm  length  (and  range)  for  12  adult  males  for  comparison. 

differ,  and  species  may  have  one  to  three  young  per  litter.  However,  most 
phyllostomatids  exhibit  ratios  greater  than  0.25  (for  single  births).  Orr  (1970) 
noted  that  the  ratio  in  vespertilionids  depends  on  species  size,  larger  species 
tending  to  have  a  smaller  ratio.  Neonatal-to-mother  forearm  ratios  are  a 
better  comparative  measure.  Table  3  indicates  that  phyllostomatid  bats  may 
be  born  in  a  more  advanced  stage  than  vespertilionids  because  seven  of  eight 
species  of  phyllostomatids  have  a  ratio  usually  exceeding  0.41  whereas  this 
ratio  is  exceeded  in  only  three  of  1 3  vespertilionids. 

Discussion  and  Conclusions 

The  paucity  of  information  on  phyllostomatid  development  not  withstanding, 
available  data  suggest  that  ontogeny  and  maternal  care  in  phyllostomatids 
differs  in  several  characteristics  from  those  in  vespertilionids. 


Table  3. _ Average  neonate  to  adult  weight  and  forearm  length  ratios  in  selected  phyllostomatid  and  vespertilionid  bats.  Weights  are  given 


398 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


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BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


1.  Phyllostomatids  generally  are  born  in  a  more  precocial  condition, 
(furred,  eyes  open,  mobile,  and  size  large  relative  to  that  of  the  mother)  than 
vespertilionids.  As  Gould  (1975)  pointed  out,  there  is  no  clear  dividing  line 
between  altriciality  and  precociality,  but  within  the  two  families,  the  degree 
of  overlap  in  such  characteristics  as  mobility,  eye  opening,  and  pelage 
development  is  small. 

2.  In  phyllostomatids,  deciduous  teeth  are  reduced  in  size,  relatively  simple 
in  form,  and  functional  teeth  are  fewer  in  number.  The  deciduous  dentition 
might  be  related  to  permanent  dentition  and  different  feeding  strategies,  but 
it  might  also  correlate  with  maternal  care  patterns,  as  discussed  in  point  3 
below. 

3.  Phyllostomatid  young  usually  are  not  deposited  in  large  creches  by 
foraging  mothers.  Instead,  they  remain  attached  to  the  mother  in  the  roost 
during  the  day  and  might  be  carried  during  foraging.  Bradbury  (personal 
communication)  suggested  that  young  might  be  carried  to  a  nocturnal  roost 
before  the  female  begins  to  forage.  The  cross-wise  position  assumed  by  attached 
young  could  be  an  adaptation  of  phyllostomatids  to  frequent  carrying  by  the 
mother. 

The  occurrence  of  these  three  characteristics  in  many  species  of  phyllostomatid 
bats  is  intriguing,  especially  when  considering  how  such  adaptations  evolved. 
Carrying  young  during  foraging  or  transferring  young  to  individual  nocturnal 
roosts  before  foraging  could  serve  as  an  antipredator  strategy  for  bats  living  under 
conditions  where  other  bats  have  evolved  as  predators.  However,  transferral 
to  a  nocturnal  roost  might  be  an  adaptation  that  could  evolve  only  under  stable 
tropical  conditions  where  temperature  fluctuations  are  not  great.  By  contrast, 
creches  of  vespertilionids  might  function,  in  part,  to  retain  heat  in  the  altricial 
young.  Clearly,  behavioral  studies  in  the  field  are  needed  to  determine  how  onto- 
gency  and  maternal  care  in  the  Phyllostomatidae  relate  to  feeding  strategies,  social 
organization,  roosting  behavior,  and  possible  antipredator  mechanisms. 

Acknowledgments 

We  are  grateful  to  E.  Gould  for  providing  us  with  the  captive  colony  of 
Carollia  perspicillata,  Anoura  geoffroyi,  and  Glossophaga  soricina.  The 
study  specimens  examined  from  the  United  States  National  Museum  of 
Natural  History,  Smithsonian  Institution,  included  five  Carollia  perspicillata 
and  two  C.  brevicaudatum  (USNM  104551;  104552;  179612;  284511- 
284513;  65475).  We  thank  R.  H.  Pine  for  his  assistance.  The  literature  review 
for  this  article  was  completed  in  early  1975. 

Literature  Cited 

Ayala,  S.  C.,  and  A.  D’Alessandro.  1973.  Insect  feeding  of  some  Colombian  fruit¬ 
eating  bats.  J.  Mamm.,  54:266-267. 

Barbour,  R.  W.,  and  W.  H.  Davis.  1969.  Bats  of  America.  Univ.  Press  Kentucky, 
Lexington,  286  pp. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


401 


Bogan,  M.  A.  1972.  Observations  on  parturition  and  development  in  the  hoary  bat, 
Lasiurus  cinereus.  J.  Mamm.,  53:61 1-613. 

Bonaccorso,  F.  J.,  and  N.  Smythe.  1972.  Punch-marking  bats:  an  alternative  to 
banding.  J.  Mamm.,  53:389-390. 

Bradshaw,  G.  V.  R.  1961.  A  life  history  study  of  the  California  leaf-nosed  bat, 
Mcicrotus  californicus.  Unpublished  Ph.D.  dissertation,  Univ.  Arizona, 
Tucson,  89  pp. 

Burns,  R.  J.  1970.  Twin  vampires  born  in  captivity.  J.  Mamm.,  51:391-392. 

Crespo,  R.  F.,  R.  J.  Burns,  and  S.  B.  Linhart.  1970.  Loadlifting  capacity  of  the  vampire 
bat.  J.  Mamm.,  51:627-628. 

Davis,  R.  1969.  Growth  and  development  of  young  pallid  bats,  Antrozous  pallidus. 
J.  Mamm.,  50:729-736. 

- .  1970.  Carrying  of  young  by  flying  female  North  American  bats.  Amer. 

Midland  Nat.,  83:186-196. 

Davis,  W.  H.,  R.  W.  Barbour,  and  M.  D.  Hassell.  1968.  Colonial  behavior  of  Eptesiats 
fuscus.  J.  Mamm.,  49:44-50. 

Fenton,  M.  B.  1969.  The  carrying  of  young  by  females  of  three  species  of  bats.  Canadian 
J.  Zool.,  47:158-159. 

Goodwin,  G.  G.,  and  A.  M.  Greenhall.  1961.  A  review  of  the  bats  of  Trinidad  and 
Tabago.  Bull.  Amer.  Mus.  Nat.  Hist.,  122:195-301. 

Goodwin,  R.  E.  1970.  The  ecology  of  Jamaican  bats.  J.  Mamm.,  51:571-579. 

Gould,  E.  1970.  Echolocation  and  communication  in  bats.  Pp.  144-161,  in 
About  bats  (B.  H.  Slaughter  and  D.  W.  Walton,  eds.),  Southern  Methodist 
Univ.  Press,  Dallas,  vii  +  1-339. 

- .  1971.  Studies  of  maternal-infant  communication  and  development  of 

vocalizations  in  the  bats  Myotis  and  Eptesicus.  Comm.  Behav.  Biol.,  5:263-313. 

- .  1975.  Neonatal  vocalizations  in  bats  of  eight  genera.  J.  Mamm.,  56:15-29. 

Hoffmeister,  D.  F.  1959.  Distributional  records  of  certain  mammals  from 

Southern  Arizona.  Southwestern  Nat.,  4: 14-19. 

Jenness,  R„  and  E.  H.  Studier.  1976.  Lactation  and  milk.  Pp.  201-218,  in 
Biology  of  bats  of  the  New  World  family  Phyllostomatidae.  Part  1  (R.  J.  Baker, 
J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.),  Spec.  Publ.  Mus.,  Texas  Tech 
Univ.,  10:1-218. 

Jennings,  W.  L.  1958.  The  ecological  distribution  of  bats  in  Florida.  Unpublished 
Ph.D.  dissertation,  Univ.  Florida,  Gainesville,  126  pp. 

Jones,  C.  1967.  Growth,  development,  and  wing-loading  in  the  evening  bat 
Nycticeius  humeralis.  J.  Mamm.,  48:1-19. 

Kleiman,  D.  G.  1969.  Maternal  care,  growth  rate,  and  development  in  the  noctule 
( Nyctalus  noctula),  pipistrelle  ( Pipistrellus  pipistrellus),  and  serotine 
{Eptesicus  serotinus)  bats.  J.  Zool.,  157:187-211. 

Kleiman,  D.  G.,  and  T.  M.  Davis.  1974.  Punch-mark  renewal  in  bats  of  the  genus 
Carollia.  Bat  Research  News,  15:29-30. 

Klima,  M.,  and  J.  Gaisler.  1968.  Study  on  growth  of  juvenile  pelage  in  bats,  III.  Phyl¬ 
lostomatidae.  Zoolog.  Listy,  17:1-18. 

Kunz,  T.  H.  1973.  Population  studies  of  the  cave  bat  ( Myotis  velifer ):  reproduction, 
growth,  and  development.  Univ.  Kansas  Occas.  Papers  Mus.  Nat.  Hist., 
15:1-43. 

Lane,  H.  K.  1946.  Notes  on  Pipistrellus  subflavus  subflavus  (F.  Cuvier)  during  the 
season  of  parturition.  Proc.  Pennsylvania  Acad.  Sci.,  20:57-61. 

Medway,  L.  1972.  Reproductive  cycles  of  the  flat-headed  bats,  Tylonycteris  pachypus 
and  T.  robustula  (Chiroptera:  Vespertilioninae)  in  a  humid  equatorial  environ¬ 
ment.  Zool.  J.  Linn.  Soc.,  51:33-61. 


402 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Miller,  G.  S.,  Jr.  1907.  The  families  and  genera  of  bats.  Bull.  U.  S.  Nat.  Mus., 
57:1-282. 

Mumford,  R.  E.,  and  D.  A.  Zimmerman.  1964.  Notes  on  Choeronycteris  mexicana.  J. 
Mamm.,  43:101-102. 

Novick,  A.  1960.  Successful  breeding  in  captive  Artibeus.  J.  Mamm.,  41:508-509. 

O'Farrell,  M.  J.,  and  E.  H.  Studier.  1973.  Reproduction,  growth,  and  development 
in  Myotis  thyscinodes  and  M.  lucifugus  (Chiroptera:  Vespertilionidae).  Ecology, 
54:18-30. 

Orr,  R.  T.  1954.  Natural  history  of  the  pallid  bat,  Antrozous  pall  id  us  ( LeConte).  Proc. 
California  Acad.  Sci.,  28:165-246. 

- .  1970.  Development:  prenatal  and  postnatal.  Pp.  217-231,  in  Biology  of 

Bats  (W.  A.  Wimsatt,  ed.).  Academic  Press,  New  York,  1:406  pp. 

Pearson,  O.  P.,  M.  A.  Koford,  and  A.  K.  Pearson.  1952.  Reproduction  of  the 
lump-nosed  bat  ( Corynorhinus  rafinesquii)  in  California.  J.  Mamm.,  33:273-320. 

Phillips,  C.  J.  1971.  The  dentition  of  the  glossophagine  bats:  development,  morphological 
characteristics,  variation,  pathology  and  evolution.  Misc.  Publ.  Mus.  Nat. 
Hist.,  Univ.  Kansas,  54:1-138. 

Pine,  R.  H.  1972.  The  bats  of  the  genus  Carollia.  Tech.  Monogr.,  Texas  Agric. 

Exp.  Sta.,  Texas  A&M  Univ.,  8:1-125. 

Rasweiler,  J.  J.,  and  H.  De  Bonilla.  1972.  Maintaining  nectarivorous  phyllostomatid 
bats  in  the  laboratory.  Lab.  Anim.  Sci.,  22:658-663. 

Schmidt,  U.,  and  U.  Manske.  1973.  Die  Jugendentwicklung  der  Vampirfledermause 
(Desmodus  rotundus).  Z.  Saugetierk.,  38:14-33. 

Tamsitt,  J.  R.,  and  D.  Valdivieso.  1963a.  Records  and  observations  on  Colombian 
bats.  J.  Mamm.,  44:168-180. 

- .  1963  6.  Reproductive  cycle  of  the  big  fruit-eating  bat,  Artibeus  lituratus, 

Olfers.  Nature,  198:194. 

- .  1965.  Reproduction  of  the  female  big  fruit-eating  bat,  Artibeus  lituratus 

pal  mar  um,  in  Colombia.  Caribbean  J.  Sci.,  5:157-166. 

- .  1966a.  Taxonomic  comments  on  Anoura  caudifer,  Artibeus  lituratus,  and 

Molossus  molossus.  J.  Mamm.,  47:230-238. 

- .  19666.  Parturition  in  the  red  fig-eating  bat,  Stenoderma  rufum.  J.  Mamm., 

47:352-353. 

Wimsatt,  W.  1960.  An  analysis  of  parturition  in  Chiroptera,  including  new  observations 
on  Myotis  l.  lucifugus.  J.  Mamm.,  41:183-200. 


GENERAL  PHYSIOLOGY 


John  M.  Burns 


At  first  exposure  to  this  volume,  as  well  as  its  previous  companions,  one  is 
amazed  at  the  amount  of  information  that  has  accumulated  concerning  the  biology 
of  New  World  leaf-nosed  bats.  Upon  closer  inspection,  however,  it  is  apparent 
that  the  vast  majority  of  this  information  deals  with  taxonomy,  distribution, 
natural  history,  and  various  aspects  of  morphology.  Physiological  study  of  these 
biologically  important  mammals  has  been  a  neglected  area,  at  least  as  judged  by 
the  published  literature. 

Two  physiological  systems  that  have  been  examined  to  a  substantial  degree  and 
warrant  separate  consideration  are  sensory  physiology  (primarily  echolocation) 
and  thermoregulation.  Gould  (1977)  and  McManus  (1977)  have  provided  excel¬ 
lent  reviews  of  these  respective  topics  and  I  shall  not  atterqpt  to  duplicate  here 
the  information  presented  in  these  two  papers. 

Endocrine  studies,  in  particular,  are  lacking  for  phyllostomatids.  The  reason 
for  this  probably  can  be  attributed  to  the  fact  that  bats  are  small  and  therefore 
have  small  blood  volumes.  Until  the  last  decade,  measurements  of  hormone 
concentrations  were  dependent  mostly  on  bioassays  that  required  blood  to  be 
pooled  from  several  bats.  Determination  of  hormone  concentration  is  no  longer 
a  major  problem  because  such  techniques  as  radioimmunoassay  (RIA)  and 
fluorescent  immunoenzyme  assay  require  only  50  to  100  microliters  of  plasma. 
Echolocation  and  thermoregulation  studies,  on  the  other  hand,  have  allowed 
investigators  to  work  with  entire  animals  without  the  need  for  expensive  equip¬ 
ment. 

Many  interesting  questions  can  be  raised  as  to  the  role  of  chiropteran  endocrine 
systems  in  such  physiological  endeavors  as  water  balance,  bone  and  calcium 
metabolism,  and  digestion.  For  the  moment,  we  can  only  surmise  that  such 
endocrine  regulation  is  similar  to  that  known  for  other  mammals. 

Reproductive  Physiology 

In  view  of  the  great  deal  of  emphasis  placed  on  reproductive  physiology  of 
animals  over  the  past  several  decades,  one  would  be  inclined  to  suppose  that  there 
is  a  wealth  of  such  information  for  phyllostomatid  bats.  However,  the  vast 
majority  of  literature  on  reproduction  in  leaf-nosed  bats  deals  with  studies  of 
comparative  anatomy,  morphology,  natural  history,  and  fecundity  rather  than 
with  the  physiological  processes  of  reproduction.  As  Wilson  (this  volume) 
pointed  out,  reproductive  strategies  of  phyllostomatids  are  varied.  These  include 
such  schemes  as  monestry,  polyestry,  and  a  system  that  Bradshaw  (1962)  termed 
delayed  development  for  Macrotus  californicus\  a  similar  system  was  reported 
(Fleming,  1971)  for  Artibeus  jamaicensis.  A  unique  gestation  pattern  was  re¬ 
ported  in  Macrotus  californicus  for  thyroid  hormone  (Bums  et  al.,  1972), 


403 


404 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


estrogens  (Burns  and  Wallace,  1975),  and  for  progesterone  (Burns  and  Easley, 
1977).  In  each  of  these  reports,  biphasic  patterns  were  described  in  which  one 
peak  coincided  with  the  fertilization  and  implantation  period  of  October  and 
November,  followed  by  a  second  peak  in  May  and  June  that  corresponded  to 
fetal  maturation  and  parturition.  The  hormonal  data  (summarized  in  Table  1), 
as  well  as  the  histological  studies  of  Bleier  (1975a,  1975  b),  suggest  that  the 
reproductive  scheme  in  M.  californicus  is  quite  different  from  delayed  im¬ 
plantation. 

Krutzsch  et  al.  (1976)  reported  changes  in  plasma  testosterone  and  testicular 
ascorbic  acid  in  reproductively  active  male  M.  californicus ;  testosterone  and 
testicular  ascorbic  acid  reached  a  peak  concentration  of  2.7  ng/ml  and  38  ug/ml, 
respectively,  in  late  summer,  and  spermatazoa  were  present  in  the  epididymides 
from  August  to  early  December.  The  testes  began  to  atrophy  by  late  September, 
and  the  levels  of  testosterone  and  testicular  ascorbic  acid  declined  by  December 
but  were  detectable  the  year  around  (minimum  concentrations  observed  for 
testosterone  were  0.25  ng/ml;  ascorbic  acid,  1  ug/ml.). 

The  seminal  vesicles  and  prostate  glands  were  at  maximum  size  in  September 
(15  mm  diameter,  19  mg  weight)  and  slowly  digressed  beginning  in  late  autumn. 

In  my  studies  at  Texas  Tech  University,  I  also  found  that  M.  californicus  is 
an  adaptable  animal  for  laboratory  study.  After  individuals  are  fed  by  hand  for 
2  to  3  days,  they  are  tamed  quite  rapidly.  When  bats  were  housed  in  large  cages, 
which  allow  for  adequate  freedom  of  flight,  attempts  to  establish  breeding  colonies 
proved  successful  (unpublished). 


Thyroid 

Reports  on  thyroid  physiology  are  scarce  and  usually  play  a  minor  role  in  larger 
studies  related  to  thermoregulation  or  reproduction.  Sadler  and  Tyler  (1960a) 
examined  thyroid  function  in  a  nonhibernating  bat,  Macrotus  californicus ,  by 
means  of  131 1  uptake.  Animals  were  tested  over  a  temperature  range  of  24°  to 
37  °C,  and  it  was  found  that  chronic  exposure  to  these  temperatures  did  not 
influence  thyroid  activity.  This  is  quite  different  from  responses  of  hibernating 
species  of  vespertilionids,  which  show  drastic  changes  in  the  rate  of  thyroid 
uptake  of  radioactive  iodine  when  subjected  to  a  similar  temperature  regime  as 
described  above  for  M.  californicus  (Sadler  and  Tyler,  1960 b). 

Burns  et  al.  (1972)  reported  a  drastic  decrease  in  plasma  thyroxine  for  M. 
californicus  during  the  second  trimester  of  pregnancy  (see  Table  1).  It  was  found 
later,  however,  that  triiodothyronine  (T3)  levels  were  elevated  throughout  the 
gestation  period  to  the  extent  that  total  thyroid  hormone  concentration  in  the 
blood  during  pregnancy  remained  essentially  unchanged  (unpublished). 

Adrenal  Glands 

Studies  that  attempt  to  describe  the  role  of  either  adrenocortical  or  medullary 
hormones  in  regulating  a  host  of  physiological  processes  in  phyllostomatid  bats 
are  unknown.  Such  a  work  would  represent  a  “first”  for  comparative  physiology. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


405 


Table  1 . — Changes  in  plasma  concentrations  of  various  hormones  during  pregnancy  /nMacrotus 

californicus. 


Hormone 

Aug. 

Sept. 

Oct. 

Nov. 

Dec. 

Jan. 

Feb. 

Mar. 

Apr. 

May 

June 

Thyroxine1 

3.5 

2.0 

1.0 

1.0 

2.0 

5.0 

5.0 

Progesterone2 

1 

1  1 

16 

22 

5 

8 

7 

13 

31 

15 

Estrone3 

0 

10 

12 

15 

12 

12 

25 

Estradiol- 17  6 

0 

40 

55 

60 

35 

40 

75 

■Thyroxine  concentrations  were  measured  by  a  column  chromatography-colorimeter  technique  and  are 
expressed  as  microgram  per  cent.  The  June  sample  is  from  lactating  females.  From  Burns  et  at.  (1972). 

2Progesterone  concentrations  were  determined  by  radioimmunoassay  and  are  expressed  as  nanograms 
per  milliliter.  The  June  sample  represents  preparturition  samples.  From  Burns  and  Easley  (1977). 

3Estrogens  determined  by  radioimmunoassay  and  expressed  as  picograms  per  milliliter.  The  values  for 
August  mean  that  estrogen  levels  were  not  detectable  with  this  assay.  From  Burns  and  Wallace  (1975). 


There  are  a  few  reports  that  describe  the  basic  morphology  of  chiropteran  adrenal 
glands  (for  example,  see  Christian,  1963)  but  no  attempt  has  been  made  to 
elucidate  the  role  of  the  adrenal  glands  in  a  physiological  sense. 

Parathyroid  Glands 

I  was  unable  to  find  a  published  report  of  any  investigation  that  dealt  with  the 
function  or  particular  characteristics  of  the  parathyroid  glands  in  any  member 
of  the  Phyllostomatidae. 


Renal  Physiology 

The  great  success  of  Chiroptera  in  general,  and  phyllostomatids  in  particular, 
suggests  that  some  species  have  evolved  elaborate  and  highly  efficient  renal 
mechanisms  for  conserving  water.  Most  studies  of  renal  function  pertain  to 
evaporative  water  loss,  however,  and  are  orientated  more  toward  thermoregulation 
than  anything  else  (see  McManus,  1977).  There  is  a  paucity  of  information 
concerning  renal  physiology  and  evidently  an  absolute  absence  of  data  dealing 
with  the  endocrine  regulation  of  renal  function  in  leaf-nosed  bats. 

McFarland  and  Wimsatt  (1965,  1969)  reported  on  the  unusual  ability  of  the 
kidneys  in  the  vampire  bat  Desmodus  rotundus  to  concentrate  urine.  At  first, 
one  might  question  the  physiological  demand  that  would  result  in  development  of 
a  versatile  renal  system  in  an  animal  with  a  diet  that  is  approximately  98  per  cent 
water.  McFarland  and  Wimsatt  (1969)  proposed  that  the  majority  of  the  water 
content  ingested  with  a  blood  meal  must  be  eliminated  rapidly  for  purposes  of 
flight.  This  would  result  in  a  meal  residue  composed  almost  entirely  of  protein, 
which  represents  a  nitrogen  load  that  must  be  excreted  with  a  minimum  of  urinary 
water  loss.  McFarland  and  Wimsatt  (1969)  also  reported  that  the  vampire  bat 
concomittantly  forms  urine  at  a  high  rate  (4  ml/kg/minute)  and  a  low  osmolality 
(475  mOs)  during  feeding.  Five  to  six  hours  after  feeding,  the  rate  of  urine  pro¬ 
duction  falls  to  approximately  0.2ml/kg/minute,  with  a  surprising  high  urine 
concentration  (4656  mOs).  Wimsatt  and  Guerriere  (1962)  also  reported  on  the 
relationship  of  volume  of  blood  consumed  by  D.  rotundus  to  amount  of  urine 


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excreted.  For  example,  if  the  blood  meal  is  35  milliliters,  the  urine  volume  excreted 
shortly  after  feeding  is  approximately  26  milliliters.  Also  of  interest  is  the 
observation  (Wimsatt  and  Guerriere,  1962)  that  isolated  D.  rotundus  have  a 
somewhat  higher  average  daily  consumption  of  blood  than  do  bats  held  captive  in 
groups  (21.2  as  compared  to  15.5  milliliters).  The  physiological  significance  of 
these  observations  is  not  known. 

Whereas  Desmodus  rotundus  demonstrates  a  remarkable  ability  to  concentrate 
urine,  the  nectarivorous  Leptonycteris  sanborni  has  little  physiological  capability 
in  this  regard.  Carpenter  (1969)  showed  that  even  when  individuals  of  L.  sanborni 
collected  from  desert  habitats  were  placed  on  a  high  protein  diet,  the  maximum 
urine  concentration  was  only  342  mOs.  This  value  is  even  less  concentrated  than 
that  reported  by  Schmidt-Nielsen  and  O’Dell  (1961)  for  semiaquatic  mammals 
such  as  beaver,  Castor  canadensis.  Normally,  L.  sanborni  feeds  on  nectar  from  a 
variety  of  desert  plants  that  are  high  in  water  and  carbohydrates.  Howell  (1974) 
showed  that  this  species  obtains  proteins  and  amino  acids  by  consuming  pollen 
of  the  saguaro  cactus  as  a  dietary  supplement.  The  pollen’s  nitrogenous  degradation 
products  are  concentrated  in  the  urine  and  then  actively  ingested  by  the  bat.  This 
behavior  results  in  a  positive  nitrogen  balance,  a  condition  otherwise  impossible 
on  a  pollen-free  diet. 


Respiratory  Physiology 

Inasmuch  as  bats  lack  the  more  efficient  flow-through  air  sac  arrangement 
characteristic  of  birds,  they  must  devote  a  substantially  greater  portion  of  their 
body  to  respiratory  surface  tissue.  For  example,  the  common  crow,  Corvus 
brachyrhynchos,  has  a  respiratory  surface  area  in  its  lungs  of  approximately 

O. 6  square  centimeter  per  gram  of  body  weight  (McCauley,  1971),  whereas  small 
bats,  such  as  those  in  the  vespertilionid  genus  Myotis,  must  devote  100  square 
centimeters  per  gram  of  body  weight  so  as  to  meet  the  metabolic  demand  of 
flight.  It  does  not  appear,  however,  that  this  poses  an  anatomical  disadvantage 
for  bats  because  flight  is  an  efficient  method  of  travel  for  chiropterans.  For 
example,  Thomas  (1975)  calculated  that  Phyllostomus  hastatus  requires 
only  one-sixth  the  energy  needed  by  a  terrestrial  mammal  of  the  same  size  to 
cover  a  given  distance.  He  also  calculated  the  metabolic  rate,  in  watts,  for  flying 

P.  hastatus  (0.93  kg)  as  130.4  w/kg-1.  Thomas  also  stated  that  such  metabolic 
rates  are  essentially  the  same  as  the  predicted  values  for  flying  birds  of  similar 
body  size,  but  that  they  are  two  and  a  half  to  three  times  greater  than  the  highest 
metabolic  rates  of  which  exercising  terrestrial  mammals  of  similar  size  appear 
capable. 

Thomas  and  Suthers  (1972)  provided  some  interesting  data  concerning  the 
differences  in  respiration  at  rest  and  during  flight  for  Phyllostomus  hastatus, 
which  are  summarized  in  Table  2. 

They  also  reported  that  the  heart  rate  of  preflight  P.  hastatus  was  8.7  beats 
per  second  as  compared  to  13  beats  per  second  (780  beats  per  minute)  in  the 
first  few  seconds  of  flight.  Lastly,  Thomas  and  Suthers  recorded  the  hematocrit 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


407 


Table  2. — Comparison  of  difference  in  respiration  for  resting  and  flying  Phyllostomus  hastatus 
(from  Thomas  and  Slithers,  1972).  Weight  is  given  in  grams  and  metabolic  rate  in  terms  of 
milliliters  of  oxygen  per  gram  of  body  weight  per  hour. 


Weight 

Metabolic  rate 
ml02  (gh)-1 

Ventillation  rate 
(breaths/second) 

Before  flight 

101 

6.78  ±  0.85 

2.8 

87 

6.12  ±  1.15 

During  flight 

101 

27.53  ±  0.79 

10.6 

87 

24.68  ±  1.87 

of  P.  hastatus  as  60  per  cent.  This  is  considerably  greater  than  the  percentage 
of  red  blood  cells  found  in  a  given  volume  of  blood  from  any  avian  species  listed 
by  Sturkie  (1965);  the  higher  erythrocyte  number  probably  reflects  one  of  the 
general  physiological  adaptations  for  flight  in  bats. 

Electrophoretic  properties  of  some  phyllostomatid  hemoglobins  have  been 
described.  Valdivieso  et  al.  (1969)  found  a  single,  common  hemoglobin  band  for 
Monophyllus  redmani,  Artibeus  jamaicensis ,  Stenoderma  rufum,  and  Erophylla 
bombifrons.  A  similar,  more  comprehensive  electrophoretic  survey  was  reported 
by  Mitchell  (1966).  Additional  hematological  data  for  leaf-nosed  bats  were 
reported  by  Valdivieso  and  Tamsitt  (1971),  who  concluded  that  hematocrit 
values  for  frugivorous  species  are  lower  than  those  found  in  insectivorous  bats. 

Concluding  Remarks 

This  contribution  to  the  biology  of  New  World  leaf-nosed  bats  is  an  indication 
of  what  little  is  known  concerning  their  physiology  rather  than  a  survey  and 
review  of  a  substantial  body  of  knowledge.  It  also  represents  perhaps  a  subtle  plea 
to  comparative  physiologists  to  turn  their  attention  to  phyllostomatids.  Techniques 
now  are  available  for  measuring  biological  molecules  in  blood  samples  of  small 
volume.  Hopefully,  future  investigators  will  take  advantage  of  this  technology. 

Literature  Cited 

Bradshaw,  G.  V.  R.  1962.  Reproductive  cycle  of  the  California  leaf-nosed  bat,  Macrotus 
cal  ifornicus.  Science,  136:645. 

Bleier,  W.  J.  \915a.  Early  embryology  and  implantation  in  the  California  leaf-nosed 
bat,  Macrotus  californicus.  Anat.  Rec.,  182:237-254. 

- .  19756.  Fine  structure  of  implantation  and  the  corpus  luteum  in  the  California 

leaf-nosed  bat,  Macrotus  californicus.  Unpublished  Ph.D.  dissertation,  Texas 
Tech  Univ.,  Lubbock,  Texas,  vii+75  pp. 

Burns,  J.  M„  and  R.  G.  Easley.  1977.  Hormonal  control  of  delayed  development  in 
the  California  leaf-nosed  bat,  Macrotus  californicus.  III.  Changes  in  plasma 
progesterone  during  pregnancy.  Gen.  Comp.  Endocrinol.,  32:163-166. 

Burns,  J.  M.,  and  W.  C.  Wallace.  1975.  Hormonal  control  of  delayed  development 
in  Macrotus  waterhousii.  II.  Radioimmunoassay  of  plasma  estrone  and  estradiol 
17B  during  pregnancy.  Gen.  Comp.  Endocrinol.,  25:529-533. 


408 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Burns,  J.  M.,  R.  J.  Baker,  and  W.  J.  Bleier.  1972.  Hormonal  control  of  delayed 
development  in  Macrotus  waterhousii.  I.  Changes  in  plasma  thyroxine  during 
pregnancy  and  lactation.  Gen.  Comp.  Endocrinol.,  18:54-58. 

Carpenter,  R.  E.  1969.  Structure  and  function  of  the  kidney  and  the  water  balance  of 
desert  bats.  Physiol.  Zool.,  42:288-302. 

Christian,  J.  J.  1963.  Endocrine  adaptive  mechanisms  and  the  physiologic  regulation  of 
population  growth.  Pp.  189-353,  in  Physiological  mammalogy,  (R.  G.  Van 
Gelder  and  W.  Mayer,  eds.).  Academic  Press,  New  York,  l:xii+  1-381. 

Fleming,  T.  H.  1971.  Artibeus  jamaicensis :  delayed  embryonic  development  in  a 
neotropical  bat.  Science,  171:402-404. 

Gould,  E.  1977.  Echolocation  and  communication.  Pp.  247-279,  in  Biology  of  bats 
of  the  New  World  family  Phyllostomatidae.  Part  II  (R.  J.  Baker,  J.  K.  Jones,  Jr., 
and  D.  C.  Carter,  eds.),  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  13:1-364. 

Howell,  D.  J.  1974.  Bats  and  pollen:  physiological  aspects  of  the  syndrome  of  chi- 
ropterophily.  Comp.  Biochem.  Physiol.,  48A:263-276. 

Krutzsch,  P.  H.,  R.  W.  Watson,  and  C.  P.  Lox.  1976.  Reproductive  biology  of  the  male 
leaf-nosed  bat,  Macrotus  waterhousii,  in  the  southwestern  United  States.  Anat. 
Rec.,  184:611-636. 

McCauley,  W.  J.  1971.  Vertebrate  physiology.  W.  B.  Saunders  Co.,  Philadelphia, 
xiv  +  422  pp. 

McFarland,  W.  N.,  and  W.  A.  Wimsatt.  1965.  Urine  flow  and  composition  in  the 
vampire  bat.  Amer.  Zool.,  5:662. 

- .  1969.  Renal  function  and  its  relation  to  the  ecology  of  the  vampire  bat,  Desmodus 

rotundas.  Comp.  Biochem.  Physiol.,  28:985-1006. 

McManus,  J.  J.  1977.  Thermoregulation.  Pp.  281-292,  in  Biology  of  bats  of  the  New 
World  family  Phyllostomatidae.  Part  II  (R.  J.  Baker,  J.  K.  Jones,  Jr.,  and  D. 
C.  Carter,  eds.),  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  13:1-364. 

Mitchell,  H.  A.  1966.  Multiple  haemoglobins  in  bats.  Nature,  210:1067-1068. 

Sadler,  W.  W.,  and  W.  S.  Tyler.  1960a.  Thyroidal  activity  in  hibernating  Chiroptera. 
I.  Uptake  of  131I.  Acta.  Endocrinol.,  34:586-596. 

- .  19606.  Thyroidal  activity  in  hibernating  Chiroptera.  II.  Synthesis  of  radio- 

iodinated  amino  acids.  Acta  Endocrinol.,  34:597-604. 

Schmidt-Nielsen,  B.,  and  R.  O’Dell.  1961.  Structure  and  concentrating  mechanism  of 
the  mammalian  kidney.  Amer.  J.  Physiol.,  Zool.,  200:1 1 19-1 124. 

Sturkie,  P.  D.  1965.  Avian  physiology.  Cornell  University  Press,  Ithaca,  New  York, 
2nd  ed.,  xxvii  +  766  pp. 

Thomas,  S.  P.  1975.  Metabolism  during  flight  in  two  species  of  bats,  Phyllostomus 
liastatus  and  Pteropus  gouldii.  J.  Exp.  Biol.,  63:273-293. 

Thomas,  S.  P.,  and  R.  A.  Suthers.  1972.  The  physiology  and  energetics  of  bat  flight.  J. 
Exper.  Biol.,  57:317-335. 

Valdivieso,  D.,  and  J.  R.  Tamsitt.  1971.  Hematological  data  from  tropical  American 
bats.  Canadian  J.  Zool.,  49:31-36. 

Valdivieso,  D.,  J.  R.  Tamsitt,  and  E.  Conde-del  Pino.  1969.  Electrophoretic  properties 
of  neotropical  bat  hemoglobin.  Comp.  Biochem.  Physiol.,  30:1 17-122. 

Wimsatt,  W.  A.,  and  A.  Guerriere.  1962.  Observations  on  the  feeding  capacities  and 
excretory  functions  of  captive  vampire  bats.  J.  Mamm.,  43:17-27. 


POPULATION  AND  COMMUNITY  ECOLOGY 


Stephen  R.  Humphrey  and  Frank  J.  Bonaccorso 


Bats  are  the  numerically  dominant  group  of  mammals  in  the  Neotropics. 
They  comprise  52  per  cent  of  the  mammalian  species  in  Costa  Rica  (Robinson, 
1971)  and  46  per  cent  of  those  in  Panama  (Handley,  1966).  The  family 
Phyllostomatidae  accounts  for  55  per  cent  of  all  Costa  Rican  bat  species  and 
59  per  cent  of  the  species  in  Panama.  In  terms  of  number  of  individuals,  the 
density  of  some  phyllostomatid  species  far  exceeds  that  of  any  other  kind  of 
mammal  in  Central  America  (F.  J.  Bonaccorso  and  D.  Morrison,  unpublished). 
Additionally,  phyllostomatids  exhibit  great  diversity  in  the  types  of  food  used, 
with  specializations  for  eating  fruit,  nectar  and  pollen,  insects,  small  land 
vertebrates,  and  blood  of  birds  and  mammals.  The  importance  of  this  family 
in  diversity  and  relative  density  suggests  an  equivalent  functional  importance 
in  tropical  ecosystems. 

A  recurrent  theme  in  tropical  ecology  and  in  this  volume  is  the  seasonal 
variation  of  tropical  climate.  The  dominant  feature  of  tropical  climate  is  an 
annual  cycle  of  wet  and  dry  seasons  (see  Rumney,  1968).  It  is  not  uncommon  to 
find  tropical  dry  or  wet  forests  (forest  types  refer  to  the  classification  of  Holdridge, 
1967)  that  receive  200  to  400  millimeters  of  rain  per  month  in  the  wet  season 
and  no  measurable  rain  in  some  dry  season  months.  Tropical  wet  and  rain 
forests  have  less  distinct  dry  seasons  but  predictably  have  reduced  rainfall  in 
certain  months.  The  influence  of  wet-dry  seasonality  on  the  foraging  and 
reproduction  of  tropical  bats  was  discussed  by  Baker  and  Baker  (1936), 
Mutere  (1968,  1970),  Liat  (1970),  Mares  and  Wilson  (1971),  Fleming  et  al. 
(1972),  and  Heithaus  et  al.  (1975). 

Foraging  Strategy 

Optimal  foraging  strategy  requires  that  animals  maximize  food  intake 
(benefits)  while  minimizing  expenditure  of  time  and  energy  (costs)  of  acquiring 
food.  The  distribution  of  food  resources  in  time  and  space,  the  type  of  food 
eaten,  and  competition  for  food  all  weigh  heavily  in  shaping  foraging  strategy 
(Schoener,  1969).  Additionally,  transitional  stages  in  the  evolution  of  species 
or  individual  life  histories  may  coincide  with  less  than  optimal  time-energy 
budgets  when  animals  use  excessive  energy  to  exploit  new  resources.  For 
example,  some  phyllostomatids  that  change  their  diets  seasonally  may  incur 
such  increased  foraging  costs. 

In  this  section,  we  discuss  factors  influencing  the  foraging  strategies  of 
phyllostomatids.  We  suggest  that  Neotropical  bats  feeding  on  vertebrates  and 
blood  can  rely  on  stable  and  abundant  food  resources  throughout  the  year. 
On  the  other  hand,  fruits,  flowers,  and  insects  are  extremely  seasonal  in  abundance. 


409 


410 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Some  phyllostomatids  specializing  on  these  food  types  may  encounter  local 
shortages  at  predictable  times  of  the  year.  In  order  to  survive  such  food  shortages, 
foraging  strategies  of  tropical  bats  include  migration,  dietary  changes,  dis¬ 
continuation  of  reproduction,  and  successfully  competing  with  other  species 
for  limited  food  resources.  In  addition,  phyllostomatids  do  have  seasonal 
fat  cycles  (McNab,  1976)  and  undergo  at  least  diel  torpor  (McNab,  1969); 
these  strategies  also  might  help  in  the  accommodation  of  food  shortages. 

Fruit 

Fruit  availability  in  tropical  forests  varies  in  complex  ways.  Some  tree 
species  produce  fruit  synchronously  each  year  at  a  characteristic  season.  Some 
fruit  rhythmically  but  not  every  year.  Others  fruit  with  no  discernable  pattern 
from  once  every  few  years  to  several  times  a  year  (Richards,  1973;  Foster, 
1973;  Frankie  et  al.,  1975). 

Thorough  studies  of  fruiting  patterns  in  tropical  dry,  moist,  and  wet  forest 
plant  communities  have  been  conducted  in  Panama  by  Foster  (1973)  and  in 
Costa  Rica  by  Frankie  et  al.  (1975).  These  studies  show  that  edible  fruit  is 
available  throughout  the  year,  regardless  of  life  zone,  but  that  sharp  seasonal 
fluctuations  occur  in  the  number  of  species  fruiting  and  the  total  fruit  biomass. 
In  dry  forest,  a  single  peak  in  the  number  of  species  with  mature  fruits  occurs 
during  the  wet  season.  Both  moist  and  wet  forests  have  two  peaks  in  the 
number  of  species  fruiting,  one  each  in  the  dry  and  wet  seasons. 

Heithaus  et  al.  (1975)  studied  the  foraging  patterns  and  resource  use  of 
six  fruit-eating  phyllostomatids  near  Canas,  Costa  Rica.  The  tropical  lowland 
dry  forest  of  Canas  has  a  wet  season  from  May  to  early  November  and  a  dry 
season  from  mid-November  through  April.  Virtually  no  rain  falls  in  the  dry 
season,  and  the  forests  are  semideciduous,  with  about  half  the  tree  species  losing 
their  leaves  (Daubenmire,  1972).  In  each  month,  between  five  and  10  species 
of  plants  produce  fruit  eaten  by  bats.  A  single  strong  peak  in  the  number  of 
plant  species  with  “bat  fruits”  occurs  from  May  through  August.  During  the 
early  dry  season,  when  the  fewest  kinds  of  fruits  are  ripe,  a  peak  in  the  number 
of  species  of  blooming  “bat  flowers”  occurs.  At  that  time,  all  “fruit  bats”  at 
Canas  switch  in  part  to  a  pollen  and  nectar  diet.  Three  species  ( Carollia 
perspicillata,  Sturnira  lilium,  and  Artibeus  jamaicensis )  that  eat  fruit, 
nectar,  and  pollen  reproduce  twice  each  year — once  coinciding  with  the  dry 
season  and  once  with  the  fruit  abundance  in  the  wet  season.  S.  lilium  undergoes 
a  marked  change  from  nectarivory  in  the  dry  season  to  frugivory  in  the  wet 
season.  Thus,  female  S.  lilium  on  a  diet  that  is  either  primarily  nectar  and 
pollen  or  primarily  fruit  are  able  to  nurse  young. 

Our  own  unpublished  data  from  a  moist  forest  site,  Barro  Colorado  Island, 
Panama,  reveal  that  from  six  to  19  species  of  bat  fruits  are  available  each  month. 
Again,  two  peaks  in  fruit  abundance  occur,  one  in  the  wet  season  and  one  in 
the  dry  season  (Fig.  1),  and  Artibeus  jamaicensis  correspondingly  reproduces 
twice  each  year.  At  this  site,  few  bat  flowers  are  available,  and  A.  jamaicensis 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


411 


> 

30 

03 

m  -q 
C  m 
Co  *3 

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i_  m 
>  Z 

Is 

n  m 

m 

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o 


Fig.  1. — Seasonal  reproduction  of  female  Artibeus  jamaicensis  and  of  trees  supplying 
this  species  with  food,  on  Barro  Colorado  Island,  Panama. 


relies  on  a  diet  of  fruit  from  canopy  trees  throughout  the  year.  The  period  of  an 
adult  female  mammal’s  year  that  is  most  expensive  energetically  —  lactation 
(Miguela,  1969;  Studier  et  al.,  1973) — is  even  more  costly  in  the  first 
reproductive  peak  of  A.  jamaicensis ,  which  occurs  during  the  late  dry  season. 
Then  most  bats  are  simultaneously  lactating  and  pregnant  with  embryos  to  be 
born  in  the  wet  season.  Selective  pressure  for  this  postpartum  estrus  probably 
arose  from  the  combination  of  the  four-month  gestation  of  A.  jamaicensis 
and  the  occurrence  of  the  second  fruiting  peak  four  months  into  the  wet  season. 
This  reproductive  adaptation  places  the  end  of  the  second  lactation  period 
during  the  year’s  second  fruiting  peak.  Therefore,  coincidence  of  the  first 
lactation  with  the  year’s  larger  and  longer  fruiting  peak  is  a  doubly  vital 
phase  of  seasonal  timing.  The  wet  season  fruiting  peak  is  followed  by  two  months 
of  fruit  scarcity;  it  is  accompanied  by  another  postpartum  estrus  of  A.  jamaicensis, 
but  development  of  the  embryo  is  delayed  until  the  end  of  the  wet  season  (Fleming, 
1971). 

The  fruiting  patterns  of  individual  plant  species  often  are  less  important  to 
bats  than  are  the  fruiting  patterns  of  inclusive  genera.  Usually,  all  members  of 
a  genus  will  have  similar  fruits,  either  edible  or  not.  For  example,  the  1 8  species 
of  Miconia  eaten  by  birds  in  Trinidad’s  Arima  Valley  fruit  for  periods  of  one 
to  four  months,  with  fruiting  intervals  spaced  through  the  year  so  that  from  one 
to  seven  species  always  are  bearing  fruit  simultaneously  (Snow,  1965).  On 
Barro  Colorado  Island,  Panama,  and  at  Cahas,  Costa  Rica,  several  species 


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of  the  shrub  genus  Piper  (Fig.  2)  are  sympatric.  Each  fruits  cyclicly,  with 
species  cycles  offset  so  that  fruit  of  the  genus  is  available  all  year  (Heithaus 
et  al .,  1975;  our  data).  Pipers  are  the  most  important  food  species  for  bats  of 
the  genus  Carollia  in  Central  America  (Howell  and  Burch,  1974;  Heithaus 
et  al.,  1975;  our  data).  The  plant  genus  Ficus  (figs)  has  many  species  that 
consecutively  serve  as  dietary  staples  for  Artibeus  and  other  stenodermines. 
The  same  is  true  for  Cecropia  trees  and  Phyllostomus  discolor,  in  our  experience. 
The  year-round  availability  typical  of  such  dietary  staples  may  result  from 
long  coevolution  in  response  to  mutualistic  seed  dispersal  (Snow,  1965). 

Some  important  bat  fruits  are  available  only  for  several  months  and  do  not 
have  congeners  fruiting  at  other  times  of  the  year.  For  example,  Spondias 
mombin  is  ripe  only  from  September  to  December  on  Barro  Colorado  Island 
(Smythe,  1970).  (Croat,  1974,  reported  that  this  species  begins  to  fruit  in 
July,  but  this  is  true  only  along  watercourses  and  drainage  ditches.)  Its  only 
congener,  5.  radlkoferi,  also  fruits  within  this  period.  Both  species  of  Spondias 
are  important  food  items  for  bats  when  few  other  fruits  are  available  during  the 
heaviest  rains  of  the  wet  season. 

The  low  densities  of  tree  species  in  heterogeneous  Neotropical  forests  may 
force  large  foraging  distances  upon  herbivorous  bats.  Large-sized  specialists  on 
fruit  should  have  greater  foraging  distances  than  smaller  generalists.  For 
example,  Fleming  et  al.  (1972)  gave  mean  recapture  distances  of  347  meters 
for  Artibeus  jamaicensis  and  167  meters  for  Carollia  perspicillata.  Heithaus 
et  al.  (1975)  reported  that  small  species  feed  on  resources  of  high  abundance, 
whereas  large  species  use  resources  that  are  more  patchy  in  time  and  space. 

Nectar  and  Pollen 

Community  patterns  in  the  timing  of  flowering  for  Neotropical  plants, 
like  fruiting  patterns,  are  quite  complex  and  vary  by  life  zone.  In  dry  forests, 
most  species  flower  during  the  dry  season  (Allen,  1956,  summarized  by  Janzen, 
1967;  Fournier  and  Salas,  1966;  Daubenmire,  1972;  Frankie  et  al.,  1975). 
In  the  dry  forest  in  Costa  Rica  (Heithaus  et  al.,  1975),  the  number  of  species 
of  bat  flowers  in  bloom  varied  from  a  low  of  two  in  July  and  August  (wet  season) 
to  a  maximum  of  seven  in  January  and  March  (dry  season).  During  the  dry 
season,  while  bat  flowers  were  abundant,  seven  species  of  phyllostomatids 
were  regularly  covered  with  pollen  from  flower  visits.  At  this  time,  the  flowering 
periods  of  plants  were  displaced  —  adaptations  effectively  avoiding  competition 
for  the  services  of  bat  pollinators.  However,  as  the  wet  season  began  and  flowers 
decreased,  only  Glossophaga  soricina  continued  to  visit  flowers  regularly  for 
food.  Phyllostomus  discolor  apparently  responded  to  the  scarcity  of  flowers 

Fig.  2. — Flowers  and  fruits  used  as  food  by  phyllostomatids:  A,  Cecropia  eximia 
(Moraceae)  fruit;  B,  Piper  aeqaale  (Piperaeae)  fruit;  C,  Astrocaryum  standleyaniun 
(Palmaceae)  fruit;  D,  Ochroma  lagopus  (Bombacaceae)  flowers  closed  during  daytime; 
E,  Pseudobombax  septenatum  (Bombacaceae)  flower;  F,  Markea  sp.  (Solanaceae)  flower. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


413 


414 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


by  migrating.  The  other  five  species  switched  to  fruit  diets.  This  shows  how 
different  species  react  with  different  strategies  to  a  scarcity  in  food  resources 
that  are  shared  during  times  of  abundance. 

In  the  moist  forest  on  Barro  Colorado  Island,  only  five  kinds  of  flowers  appear 
to  be  fed  on  by  bats  (see  Figs.  2d  and  2e\  our  data).  These  flowers  are  available  only 
from  late  December  to  late  March,  during  the  dry  season.  Thus,  for  nine  months 
each  year,  little  nectar  or  pollen  is  available  to  bats.  Here,  nectar  and  pollen  are 
important  food  sources  only  for  G.  soricina  and  P.  discolor.  The  latter  does  not 
migrate  as  it  does  in  the  dry  forest  studied  by  Heithaus  et  al.  (1975);  instead,  it 
switches  to  a  diet  of  fruit  and  insects.  G.  soricina ,  which  typically  is  more  dependent 
on  floral  resources,  is  rare  on  the  island  but  apparently  also  switches  to  fruit. 

We  know  of  no  dry-season  flower  feeders  that  switch  wholly  to  insects  in 
the  wet  season.  Instead,  they  switch  to  fruit  or  fruit  and  insects  together  (for 
example,  G.  soricina ,  Fleming  et  al.,  1972,  and  P.  discolor  at  our  Panamanian 
site).  It  might  be  most  realistic  to  view  such  species  as  herbivores  with  omnivorous 
tendencies,  in  which  case  it  is  proper  to  wonder  if  a  plant-adapted  gastrointestinal 
tract  (Rouk  and  Glass,  1970)  could  function  effectively  on  a  wholly  insectivorous 
diet. 

Taxa,  such  as  Leptonycteris  and  Choeronycteris,  that  are  exceptional  in  not 
switching  from  plant  food,  migrate  to  stay  permanently  in  “dry  season”  environ¬ 
ments  by  moving  to  subtropical  and  warm  temperate  thorny  vegetation  zones 
where  suitable  flowers  occur  in  summer.  In  view  of  the  many  potential  competitors 
among  insectivorous  and  frugivorous  bats,  the  selective  pressure  for  migration 
is  true  nectar-pollen  specialists  should  not  be  underrated. 

Insects 

Wet-dry  seasonality  strongly  affects  the  distribution  and  abundance  of 
Neotropical  insects.  The  dry  season  presents  many  insects  with  food  shortages 
and  water  balance  problems.  Most  tropical  insects  survive  the  dry  season  as 
adults  (Janzen  and  Schoener,  1968)  rather  than  in  diapause  (as  in  winter 
survival  of  temperate  taxa).  However,  the  precise  impact  of  tropical  seasons 
on  the  food  of  insectivorous  bats  is  difficult  to  assess,  because  few  studies  deal 
with  the  particular  insects  of  interest.  These  are  nocturnal  species  either  in 
flight,  for  bats  that  catch  flying  prey,  or  active  on  leaves,  tree  trunks,  and  the 
ground,  for  bats  that  feed  by  gleaning. 

In  a  study  of  mosquito  seasonality  based  on  adults  flying  into  a  livestock-baited 
trap,  Bates  (1945)  showed  that  nocturnal  species  peak  in  abundance  immediately 
after  the  onset  of  the  wet  season.  Some  species  exhibited  a  secondary  peak 
near  the  end  of  the  wet  season,  and  all  species  were  least  common  in  the  dry 
season.  In  addition  to  this  annual  periodicity,  one  species  underwent  population 
irruptions,  with  a  hundredfold  difference  in  minimum  and  maximum  numbers 
over  a  two-year  period. 

Light-trap  samples  in  moist  forest  in  Panama  (Smythe,  1974)  document 
remarkable  seasonal  changes,  with  up  to  eight  times  as  much  insect  biomass 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


415 


Fig.  3. — Seasonality  of  tropical  insect  biomass  (after  Smythe,  1974).  This  pattern 
occurs  in  Central  America  where  distinct  dry  seasons  occur.  The  timing  varies 
geographically  by  one  to  three  months. 


in  the  wet  season  as  in  the  dry  season.  Large  taxa  ( >  5  millimeters  long)  were 
responsible  for  this  change,  with  Isoptera,  Diptera,  and  Lepidoptera  having 
particularly  dramatic  population  increases  early  in  the  wet  season.  By  contrast, 
small  taxa  (<5  millimeters  long)  were  of  constant  abundance  throughout  the 
year.  Combined  data  (Fig.  3)  show  biomass  increasing  shortly  after  the  wet 
season  begins,  peaking  about  a  month  later  when  reproduction,  growth,  and 
metamorphosis  is  complete,  and  remaining  high  for  the  next  three  or  four 
months.  Biomass  declines  late  in  the  wet  season  at  the  time  when  the  heaviest 
rains  occur,  and  it  remains  low  through  the  dry  season. 

Insectivorous  phyllostomatids  may  exhibit  at  least  three  responses  to  the 
seasonality  of  their  food.  One  would  be  to  bear  young  at  the  beginning  of  the 
wet  season;  the  limited  data  available  (see  Wilson,  this  volume)  suggest  that  this 
often  may  be  the  case.  Another  would  be  to  switch  to  other  types  of  food. 
A  partial  shift  occurs  in  Micronycteris  hirsuta,  which  gleans  insects  as  its  primary 
food  but  supplements  this  diet  with  fruit  during  the  dry  season  (Wilson,  1971). 
A  third  response  would  be  to  change  foraging  habitat.  In  the  dry  forest  near 
Canas,  Costa  Rica,  Janzen  (1973)  noted  that  night-time  numbers  of  beetle 
and  true  bug  species  decreased  much  less  during  the  dry  season  in  riparian 
forest  than  in  nearby  pasture  land  and  upland  deciduous  forest.  Thus,  riparian 
forest  may  serve  as  a  dry  season  refuge  for  food  of  insectivorous  bats,  assuming 
that  the  preferred  insect  taxa  behave  similarly. 


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Few  precise  data  exist  on  the  food  habits  of  insect-eating  phyllostomatids. 
Wilson  reported  that  large  roaches,  Orthoptera,  and  scarabaeid  beetles  are 
the  most  important  items  in  the  diet  of  Micronycteris  hirsuta  in  Panama.  These 
insects  spend  much  of  their  time  walking  and  feeding  on  vegetation  or  detritus. 
Wilson  concluded  that  M.  hirsuta  captures  most  of  its  prey  by  gleaning.  This 
pattern  appears  to  apply  to  other  members  of  the  genus  as  well  (Gardner,  1977). 
Macrotus  californicus  in  the  southwestern  United  States  also  feeds  heavily  on 
large  insects,  including  larval  Lepidoptera  that  probably  are  gleaned  from 
foliage  (Ross,  1967).  A  gleaning  mode  of  foraging  was  confirmed  for  Macrotus 
waterhousii  by  watching  one  (S.  R.  Humphrey,  aided  by  an  ultrasonic  sensor 
and  a  streetlight)  for  which  the  feeding  flight  was  confined  to  the  interior  of 
an  almost  spherical  tree  crown.  F.  J.  Bonaccarso  observed  a  captive  pair  of 
Tonatia  bidens  take  large  cicadas,  katydids,  grasshoppers,  and  beetles  by 
picking  them  off  the  walls  of  their  cage. 

Phyllostomatids  known  to  be  mainly  insectivorous  have  adaptations 
characteristic  of  bats  that  glean  prey  from  vegetation  or  the  ground.  Such 
adaptations  include  large  eyes,  large  ears,  a  long  robust  rostrum,  long  vibrissae, 
and  a  low  wing  aspect  ratio  that  promotes  vertical  flight  and  hovering.  Many 
insectivorous  phyllostomatids  with  these  features  also  have  long  nose  leafs. 
As  suggested  by  Wilson  (1971),  many  of  the  same  food-gathering  skills 
probably  are  involved  in  securing  fruit  and  resting  insects.  By  contrast,  none 
of  the  Neotropical  insectivores  of  other  families  (Emballonuridae,  Mormoopidae, 
Furipteridae,  Thyropteridae,  Vespertilionidae,  and  Molossidae,  possibly 
excepting  Natalidae)  appear  to  have  this  gleaning  morphology,  although 
several  Nearctic  vespertilionids  do  ( Antrozous ,  Euderma,  Plecotus,  Idionycteris, 
and  several  species  of  Myotis).  We  suspect  an  evolutionary  character  displacement 
at  the  family  level,  in  which  phyllostomatids  were  decisively  preeminent  as 
insect  gleaners.  A  gleaning  morphology  well  could  have  provided  suitable 
preadaptations  for  specializing  on  vertebrate  prey,  as  in  Chrotopterus  auritus 
and  Vampyrum  spectrum. 


Vertebrates 

The  literature  on  foods  of  vertebrate-eating  phyllostomatids  is  not  detailed, 
but  at  least  lizards,  birds,  mice,  and  bats  are  taken  (Goodwin  and  Greenhall, 
1961;  Gardner,  1977).  We  offer  information  on  seasonal  abundance  of  birds 
by  way  of  example.  Peaks  of  bird  breeding  should  coincide  with  high  population 
levels.  Tropical  birds  may  breed  continuously,  regularly  in  concert  with  wet-dry 
seasons,  or  irregularly.  In  continuous  breeders,  individuals  or  pairs  breed 
according  to  their  own  activity  cycles,  with  all  reproductive  stages  present  in 
the  population  at  any  time.  Most  regular  breeders  in  regions  with  a  weak  dry 
season  breed  in  the  drier  months.  In  regions  with  a  pronounced  dry  season, 
most  breed  in  the  wet  season,  but  few  specialists  breed  in  the  dry  season.  Regions 
with  two  annual  wet-dry  cycles  have  some  species  breeding  once  a  year  and 
others  breeding  twice  (Immelmann,  1971).  Superimposed  on  this  complex 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


417 


pattern  is  the  arrival  of  numerous  migrants  during  austral  and  boreal  winters. 
Thus,  at  least  in  a  general  sense,  it  appears  that  night-roosting  birds  should 
be  in  ample  supply  at  all  times  of  the  Neotropical  year. 

Blood 

Aside  from  man  and  domestic  animals,  food  for  blood-eating  phyllostomatids 
should  be  a  seasonally  stable  resource,  as  the  birds  and  mammals  parasitized 
are  large  and  have  long  life  spans.  For  the  same  reason,  these  hosts  are  likely 
to  occur  at  low  densities  and  therefore  to  be  difficult  for  vampires  to  locate. 
Conversely,  humans  and  domesticants  are  high-density  hosts  that  are  predictably 
accessible  in  time  and  space.  One  account  (Benzoni,  in  Turner,  1975)  suggests 
that  humans  were  a  major  host  for  vampires  along  the  east  coast  of  Costa  Rica 
in  the  sixteenth  century.  Now,  greater  use  of  houses  that  limit  access  makes 
human  bites  unusual,  but  vampires  commonly  and  regularly  feed  on  domestic 
birds  and  mammals.  In  accord  with  these  observations,  our  impression  from 
mist-netting  is  that  sanguivorous  bats  are  rare  except  where  domestic  animals 
are  abundant;  see  Fig.  6  for  some  illustrative  data. 

Roosting  Strategy 

Like  other  bats,  phyllostomatids  spend  the  daylight  hours  at  rest.  A  good 
roost  should  provide  some  protection  from  adverse  weather,  predators,  and 
nonresident  parasites  such  as  diurnal  mosquitoes  and  biting  flies.  Beyond  this, 
a  roost  should  afford  microclimatic  conditions  that  are  not  stressful  and  that 
favor  effective  use  of  available  energy.  At  the  very  least  (though  hardly  a 
problem),  a  roost  should  prevent  prolonged  exposure  to  direct  sunlight, 
because  phyllostomatids  die  at  body  temperatures  of  37  to  42 °C  (McManus, 
1977).  More  importantly,  microclimate  should  be  optimal  for  growth  during 
periods  of  gestation  and  lactation. 


Roost  Type 

Phyllostomatids  use  an  amazing  variety  of  natural  and  man-made  structures. 
These  include  caves,  culverts,  buildings,  bridges,  cisterns,  steam  banks,  cliff 
crevices,  tree  foliage,  tree  hollows  (even  hollow  tree  trunks  lying  on  the  forest 
floor),  rabbit  burrows,  and  old  termite  nests.  The  tent-making  bat  ( Uroderma 
bilobatum )  makes  shelters  by  clipping  palm  fronds  so  that  the  frond  tips  fold  down. 
Most  species  seem  unrestricted  to  particular  sorts  of  roosts.  For  example,  Des- 
modus  rotundus  occurs  in  tree  hollows,  caves,  and  culverts.  Commonly  several 
species  will  share  a  roost,  in  bodily  contact  with  each  other  (Goodwin  and  Green- 
hall,  1961). 


Roost  Microclimate 

The  microclimates  of  phyllostomatid  roosts  are  known  from  a  single  study. 
McNab  (1969)  recorded  air  temperature  and  relative  humidity  in  roosts  of 


418 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


12  species  at  the  time  of  capture.  Temperature  ranged  from  13  to  29  °C  and 
humidity  from  70  to  98  per  cent,  so  microclimate  is  characteristically  mild  and 
moist.  Studies  of  diel,  seasonal,  and  regional  variation  of  these  microclimates 
have  not  been  reported. 

A  few  studies  characterize  tropical  forest  microclimate  and  indicate  conditions 
that  might  be  encountered  by  a  foliage-roosting  bat.  Dry  season  data  at  Barro 
Colorado  Island,  Panama,  show  a  weekly  temperature  range  of  25.8  to  27.4 °C 
at  the  ground,  24.9  to  33.0°  in  the  subcanopy,  and  27.0  to  37.5°  in  the  canopy 
(Allee,  1926).  Relative  humidity  and  light  intensity  were  likewise  stratified, 
and  conditions  were  more  extreme  in  sunflecks  than  in  the  shade.  Similar  daytime 
temperature  profiles  occur  in  other  tropical  forests  (Hales,  1949;  Baynton 
et  al.,  1965).  Allen  et  al.  (1972)  showed  that  such  stratification  is  stable  all 
day,  breaking  down  in  the  evening,  and  that  it  is  caused  by  the  ameliorating 
effect  of  vegetation  on  air  turbulence  rather  than  any  constancy  of  incident 
conditions.  Thus,  near-lethal  temperatures  occur  in  the  canopy,  but  a  bat  can 
easily  avoid  them  by  seeking  shaded  sites  in  lower  foliage. 

Studies  of  phyllostomatid  response  to  cooling,  such  as  would  be  encountered 
at  high  latitudes  or  altitudes,  are  inconsistent,  apparently  because  of  differing 
experimental  procedures.  When  bats  were  exposed  to  rapidly  dropping 
temperatures  for  two  hours  (McNab,  1969),  four  hours  (Carpenter  and  Graham, 
1967),  or  exposed  to  cold  for  several  days  with  food  provided  ad  libitum 
(Arata  and  Jones,  1967;  Arata,  1972),  they  responded  endothermically, 
surviving  by  increasing  metabolic  rate.  Exceptions  were  small  stenodermines 
and  the  three  vampire  genera,  which  died  quickly  as  temperature  dropped. 
Animals  with  food  available  fed  many  times  a  day  when  cold.  Studier  and 
Wilson  (1970)  used  fed  animals  but  did  not  provide  food  during  their  experiments, 
lowering  temperatures  stepwise  from  34  to  2.5 °C  over  periods  of  seven  to  10 
hours,  and  allowing  body  temperatures  to  stabilize  at  each  step.  Most  individuals 
were  wholly  ectothermic  or  else  partially  so,  maintaining  body  temperatures 
5  to  15°C  above  ambient  temperatures  while  both  ambient  and  body  temperatures 
decreased.  Below  8°C  most  bats  went  into  torpor  and  died  after  failing  to  arouse. 
One  lactating  female  Carollia  perspicillata  remained  endothermic  at  ambient 
temperatures  as  low  as  5.7 °C. 

Obviously  a  bat  in  a  roost  with  a  temperature  that  is  too  low  can  leave  for 
an  alternate  site,  but  if  it  remains  it  cannot  feed  and  would  be  exposed  to  roost 
temperatures  for  approximately  eight  to  12  hours.  Realistic  thermoregulation 
studies  should  employ  microtemperatures  that  are  stable  or  that  increase 
during  the  day.  The  limited  data  on  thermal  response  lead  us  to  hypothesize 
that  at  low  roost  temperatures  (1)  reproducing  female  phyllostomatids 
thermoregulate,  incurring  the  consequent  metabolic  costs,  and  (2)  nonreproducing 
females  thermoregulate  weakly  or  not  at  all.  In  the  latter  case,  presumably  the 
practice  would  not  be  fatal  at  roost  microclimates  encountered  at  low  altitude 
in  the  Neotropics.  At  higher  latitudes  or  altitudes,  ectothermy  could  be  fatal 
and  perhaps  phyllostomatids  in  such  circumstances  attempt  to  thermoregulate. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


419 


Roosts  as  a  Limiting  Factor 

Most  phyllostomatids  roost  alone  or  in  small  colonies  and  are  not  strongly 
specialized  to  be  highly  colonial  in  order  to  exploit  particular  roost  types  (Dwyer, 
1971),  as  is  common  in  families  characteristic  of  temperate  zones  (Humphrey, 
1975).  Known  highly  gregarious  exceptions  are  Phyllonycteris,  Erophylla, 
Desmodus,  Brachyphylla,  and  Phyllostomus  (Dalquest  and  Walton,  1970). 
Satisfactory  roosts  are  available  in  abundance  in  the  Neotropics.  For  these 
reasons,  and  because  of  the  probable  importance  of  food  as  a  limiting  factor 
(McNab,  1971),  it  would  be  expected  that  roosts  seldom  limit  phyllostomatid 
abundance  and  community  structure. 

Distributional  limits  of  herbivorous  and  nonmigratory  carnivorous 
phyllostomatids  should  be  determined  by  food.  However,  roosts  may  be  limiting 
factors  at  the  distributional  limits  of  many  carnivorous  phyllostomatids, 
including  sanguivores  and  migratory  vertebrate-eaters  and  insectivores.  Dwyer 
(1971)  predicted  that  such  bats  —  that  is,  tropical  species  adapted  to  tropical 
roost  microclimates  —  will  be  limited  at  higher  latitudes  and  altitudes  by 
absence  of  suitable  food  and  by  the  increased  cost  of  thermoregulation  in  caves 
with  cool  microclimates.  Including  bats  of  all  feeding  types,  Dwyer  judged 
that  food  would  be  a  more  critical  factor  than  roosts.  In  fact,  McNab  (1973) 
calculated  that  the  cost  of  thermoregulation  in  cool  roosts  prevents  Desmodus 
rotundus  (which,  as  in  any  animal,  can  consume  only  so  much  food  nightly) 
from  occupying  higher  latitudes,  even  though  its  preferred  food  is  abundant. 
That  no  insectivorous  phyllostomatids  are  known  to  migrate  to  temperate 
zones  may  reflect  both  their  thermoregulatory  disadvantage  in  cool  roosts  and 
a  probable  competitive  advantage  on  the  part  of  vespertilionid  insect-gleaners 
that  hibernate  in  winter.  The  migratory  nectar-pollen  feeders,  Leptonycteris 
and  Choeronycteris,  move  only  as  far  north  as  the  hottest  and  driest  areas  of 
Texas  and  Arizona,  although  one  preferred  food  (Agave)  occurs  much  farther 
northward.  That  these  bats  appear  to  be  highly  colonial  at  the  northern  limits 
of  their  range  (Easterla,  1972)  may  reflect  clustering  thermoregulatory 
behavior. 


Demography 

Every  animal  can  be  said  to  have  a  demographic  strategy  —  a  combination 
of  performances  that  adds  individuals  to  the  population  in  concert  with  factors 
that  subtract  individuals,  with  a  pattern  of  magnitude,  balance,  and  timing  that 
differs  for  each  species.  A  demographic  strategy  is  a  set  of  responses  to  an 
environment;  to  some  degree  a  species  may  vary  its  strategy  among  environments 
(for  example,  in  response  to  different  seasonal  regimens  of  climate  and  food  in 
tropical  lowland  dry  as  opposed  to  wet  forest).  On  the  other  hand,  the  evolved 
nature  of  some  demographic  phenomena  (for  example,  biotic  potential,  the 
dispersal  effect  of  pioneering,  and  the  ability  to  avoid  predation)  results  in 
reasonably  fixed  numerical  expressions.  The  totality  of  demographic  events 
produces  a  growth  rate  that  must  be  positive  or  zero  over  any  substantial  time 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


interval  if  a  population  is  to  survive.  Demographic  lability  enables  a  species 
to  survive  in  a  variety  of  environments,  but  demographic  limitations  make 
success  in  other  environments  unlikely.  Values  of  demographic  parameters 
show  specific  things  an  animal  does  to  succeed  and  simultaneously  reveal 
performances  that  must  be  modified  to  enable  use  of  another  location. 

Ways  that  demographic  factors  interrelate  and  operate  are  presented  in  a 
flow  diagram  (Fig.  4).  If  population  size  and  the  natural  rate  of  increase  (r) 
are  known,  growth  trends  can  be  predicted  and  used  to  evaluate  the  progress 
of  the  population.  Three  basic  demographic  parameters  integrate  to  determine 
population  growth  rate  and  size:  natality  rate,  survival  rate,  and  dispersal  rate. 
Factors  affecting  the  values  vary  quantitatively  as  a  function  of  population  age 
structure  because  the  importance  of  each  intrinsic  factor  (for  example,  emigration) 
changes  with  age.  The  extrinsic  factors  and  potential  regulatory  pathways  are 
speculative,  as  these  seldom  or  never  have  been  demonstrated  to  operate 
among  phyllostomatids  or  other  bats.  However,  studies  directed  toward  these 
factors  and  pathways  should  reveal  the  implications  of  demographic  adaptations 
of  phyllostomatids.  Although  no  demographic  strategy  is  documented  thoroughly 
for  these  bats,  piecemeal  data  on  intrinsic  factors  ( sensu  Fig.  4)  are  reported 
in  recent  literature. 


Number  of  Births  per  Year 

Most  phyllostomatids  studied  to  date  are  polyestrous  (Fleming  et  al., 
1972;  Wilson,  this  volume)  with  the  maximum  possible  number  of  estrous 
cycles  being  two  (possibly  three  may  occur  in  tropical  vespertilionids,  as  shown 
by  Wilson  and  Findley,  1971).  Two  peaks  in  parturition  clearly  are  indicated 
by  the  bimodal  pattern  of  pregnancy  of  most  species,  and  a  maximum  number  of 
two  is  dictated  by  the  long  gestation  period  of  Desmodus  rotundus.  Of  much 
more  interest  than  the  maximum  number  of  births  possible  annually  however, 
is  the  average  number  actually  occurring.  To  our  knowledge  such  data  are 
unavailable.  In  cases  of  bimodal  polyestry,  individual  females  could  produce 
offspring  at  none,  one,  or  both  peaks  during  a  year.  Frequent  observation  of 
females  simultaneously  lactating  and  pregnant  shows  the  latter  case  to  be 
common.  Macrotus  californicus  gives  birth  only  once  per  year  (Bradshaw, 
1962).  Members  of  the  genera  Leptonycteris  and  Choeronycteris  that  annually 
migrate  from  tropical  to  warm  temperate  regions  are  parturient  during  the 
temperate  zone  summer;  available  data  do  not  preclude  the  possibility  of 
a  second  birth  during  the  tropical  dry  season.  Data  on  Leptonycteris  sanborni 
(Cockrum  and  Ordway,  1959;  Howell,  1972)  suggest  two  peaks  in  parturition 
for  each  female  or  a  single  peak  that  occurs  either  in  the  temperate  summer  or 
the  tropical  dry  season. 


Number  of  Offspring 

All  phyllostomatids  presently  are  thought  to  have  a  single  young  at  a  time. 
Carter  (1970)  termed  the  family  “characteristically  monotocous,”  and 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


421 


Fig.  4. — Operational  pathways  of  demographic  factors. 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fleming  et  al.  (1972)  stated  that  in  seasonally  polyestrous  phyllostomatids 
“one  young  is  produced  in  each  pregnancy.”  Records  of  twinning  prove  to  be 
the  exception,  not  the  rule.  Barlow  and  Tamsitt  (1968)  reported  three  sets  of 
unborn  twins  in  195  pregnancies  in  A.  jamaicensis,  one  of  615  in  Glossophaga 
soricina,  and  one  of  10  in  Erophylla  bombifrons.  Burns  (1970)  noted  one  case  in 
Desmodus  rotundus.  In  Artibeus  lituratus,  the  single  young  results  from 
reducing  the  number  of  ova  shed;  in  a  sample  of  49  females,  the  average  number 
of  corpora  lutea  or  mature  follicles  was  1.37,  but  only  one  ovum  was  released 
at  estrus  (Tamsitt  and  Valdivieso,  1965). 

Production  of  a  single  young  in  phyllostomatids  is  consistent  with  the 
pattern  generally  expected  (Spencer  and  Steinhoff,  1968;  MacArthur,  1972) 
in  that  animals  in  tropical  latitudes  have  more  numerous  but  smaller  litters  than 
those  in  temperate  latitudes.  By  contrast,  Nearctic  vespertilionids  and  molossids 
are  monestrous,  with  mean  number  of  young  ranging  from  1  to  3.4  (Humphrey, 
1975).  The  possibility  of  larger  litters  in  phyllostomatids  should  not  be  ignored 
entirely,  however,  as  some  tropical  bats  of  other  families  regularly  produce 
twins.  Examples  are  the  tropical  vespertilionids  Rhogeesa  parvula  (Cockrum, 
1955)  and  R.  tumida  (Goodwin  and  Greenhall,  1961). 

Birth  Rate 

One  important  intrinsic  factor  can  be  measured  by  answering  the  question, 
“for  each  species,  what  percentage  of  females  gives  birth  during  each  birth 
pulse?”  The  pregnancy  rate  may  approximate  the  birth  rate  in  species  with 
synchronous  parturition  if  late  abortions  and  stillbirths  are  few,  as  is  true  of 
some  temperate-zone  vespertilionids.  Single,  fortuitous  captures  of  pregnant 
phyllostomatids  (Mares  and  Wilson,  1971;  Fleming  et  al.,  1972)  demonstrate 
asynchrony  or  partial  synchrony  of  breeding.  Because  births  are  not  simultaneous 
within  seasonal  birthpulses,  these  pregnancy  data  do  not  indicate  which 
individuals  are  breeding  and  which  are  not.  However,  in  our  experience, 
properly  timed  samples  accounting  for  pregnancies  and  early  lactations  can 
generate  estimates  of  the  proportion  reproducing.  To  our  knowledge,  no 
phyllostomatid  birth  rate  data  have  been  published  (the  values  for  Desmodus 
rotundus  in  Turner,  1975,  do  not  permit  calculation  of  annual  or  seasonal 
rates). 


Age  at  Sexual  Maturity 

In  Macrotus  californicus,  females  breed  during  their  first  autumn  and  give 
birth  at  the  age  of  one  year  (Bradshaw,  1962).  Males  do  not  breed  until  their 
second  autumn,  which  is  not  disadvantageous  so  long  as  enough  males  live 
that  long,  and  it  may  be  an  advantage  in  ensuring  that  only  successful  male 
genotypes  are  perpetuated. 

To  our  knowledge,  this  important  factor  has  not  been  documented  for  any 
other  phyllostomatid,  notwithstanding  the  unsubstantiated  suggestion  that 
females  of  Artibeus  jamaicensis  become  pregnant  in  the  dry  season  following 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


423 


their  birth  (Fleming  et  al.,  1972).  Data  on  age  at  sexual  maturity  come  only 
from  recapture  of  marked  females  of  known  age  taken  from  one  to  several 
breeding  seasons  after  their  birth.  For  example,  one  Desmodus  rotundus 
marked  as  an  infant  was  pregnant  when  recaptured  18  months  later  (Turner, 
1975).  Careful  study  may  reveal  that  this  parameter  varies  in  response  to 
unusual  hardship  dictated  by  climatic  or  habitat  variation  (see  Christian,  1971). 

Survival  Rate 

The  only  record  of  phyllostomatid  longevity  exceeding  two  or  three  years 
is  a  recapture  of  an  A.  jamaicensis  seven  years  after  banding  (Wilson  and 
Tyson,  1970).  Longevity  information  is  useful  to  indicate  the  maximum  age 
attainable  by  a  species  but  is  of  little  importance  as  a  numerical  expression  of 
demographic  strategy.  The  parameter  of  interest  is  the  mean  life  expectancy 
(mean  life  span),  the  value  designated  e0  in  a  life  table.  Mean  life  expectancy 
reflects  the  actual  performance  of  a  cohort  of  animals  in  nature,  and  it 
integrates  properly  with  the  other  rate  values.  For  example,  it  allows  easy 
calculation  of  the  number  of  offspring  produced  during  the  average  lifetime  of 
a  female. 

The  only  proven  technique  for  documenting  bat  survival  is  frequent  re¬ 
capture  of  marked  individuals  of  known  age  and  sex.  Failure  to  determine 
age  at  the  time  of  marking  (for  example,  banding  cohorts  of  bats  during 
temperate  winters)  has  resulted  in  voluminous  but  not  especially  useful  bat 
survival  data  in  the  literature.  Such  data  by  definition  produce  a  constant 
rate  of  survival  throughout  life  (or  nearly  so,  depending  on  details  of  sampling 
protocol),  when  actually  mammals  characteristically  have  lower  survival  in 
immature  and  elderly  stages  than  during  adulthood  (Caughley,  1966).  The 
most  satisfactory  period  to  mark  cohorts  of  immature  bats  is  just  prior  to 
weaning.  Obtaining  accurate  data  then  depends  on  recapturing  all  of  the  living 
cohort  members  at  least  once  annually  until  the  last  individual  has  died. 
Mortality  between  birth  and  weaning  should  be  documented  to  prevent 
overestimation  of  survival.  Because  marking  extremely  young  bats  would 
cause  many  deaths,  the  best  available  technique  is  to  determine  the  number  of 
young  born  in  a  roost,  remove  all  carcasses  from  the  roost  area,  and  count  the 
number  of  young  dying  before  they  begin  to  fly.  Examples  of  such  data  are 
preweaning  survival  of  a  molossid  (Herreid,  1967),  Tadarida  brasiliensis, 
and  postweaning  survival  of  a  vespertilionid  (Humphrey  and  Cope,  1976), 
Myotis  lucifugus.  These  studies  must  be  done  at  roosts  and  require  many  years 
for  long-lived  species. 

Aging  bats  according  to  tooth  cementum  annuli  is  an  exciting  prospect, 
as  it  would  provide  an  “instantaneous”  method  of  constructing  survival  curves. 
A  disadvantage  is  that  the  animals  must  be  killed  to  acquire  the  data.  This 
technique  has  been  applied  to  Desmodus  rotundus,  yielding  mean  age  values 
of  3.0  years  for  females  and  1.5  years  for  males  in  Mexico  (Linhart,  1973) 
and  4.13  years  for  females  and  3.01  years  for  males  in  Argentina  (Lord  et  al., 


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SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


1976).  The  latter  author  suggested  that  apparent  differences  in  sex-specific 
survival  are  artifacts  of  the  social  structure  of  the  sampled  populations. 
Unfortunately,  in  neither  study  were  annuli  counts  checked  against  known-age 
control  animals  to  confirm  that  a  line  represents  one  year’s  growth.  Why  growth 
should  be  periodic  when  food  supply  is  constant  is  unclear. 

Dispersal  Rate 

Pioneering  is  a  vital  phenomenon  for  finding  available  habitats  and  com¬ 
pensating  for  local  extinctions.  For  demographic  purposes,  dispersal  rate  is  the 
net  loss  or  gain  of  animals  by  one-way  movement  in  proportion  to  the  population 
in  a  given  area.  No  measurements  of  phyllostomatid  dispersal  rates  occur  in 
the  literature.  Studies  of  dispersal  rate  of  mobile  animals  must  include  large 
geographical  areas,  and  mammalian  dispersal  is  seldom  quantified.  For  examples 
and  discussion  of  procedural  difficulties,  see  Barkalow  et  al.  (1970)  and 
Humphrey  and  Cope  (1976).  Site  attachment  index  values  and  associated 
movement  data  on  two  species  of  temperate  vespertilionids  (Humphrey  and 
Cope,  1970,  1976;  Humphrey,  1975)  indicate  little  or  no  dispersal  of  recruited 
females  in  undisturbed  populations;  whether  such  a  pattern  applies  to  phyl- 
lostomatids  in  the  tropics  in  unknown. 

Migration  has  no  effect  on  the  dispersal  rate  if  a  migrating  individual  indeed 
returns.  If  the  migrator  stays  away,  then  it  becomes  a  dispersor,  and  if  it  dies 
while  migrating  the  effect  is  on  the  survival  rate.  These  distinctions  help  prevent 
confusion  about  the  demographic  implications  of  migration.  Migration  has 
not  been  demonstrated  clearly  for  any  phyllostomatid,  but  many  sorts  of 
collateral  evidence  suggest  that  Leptonycteris  and  Choeronycteris  are  migratory 
in  the  northern  part  of  their  range  (Hayward  and  Cockrum,  1971).  Further, 
our  unpublished  data  suggest  that  some  species  in  Belize  and  Panama  are 
migratory  or  at  least  nomadic  (see  beyond). 

Community  Diversity 

Field  biologists  recognize  great  differences  in  the  various  bat  communities 
that  they  sample.  Although  patterns  of  diversity  occur  in  and  among  these 
taxonomic  communities,  so  many  characteristics  of  species  and  habitat  factors 
are  involved  that  these  patterns  are  difficult  to  perceive  and  express. 

Species  Number 

The  simplest  measure  of  diversity  is  the  number  of  species  present  (in  the 
literature  termed  variously  faunal  size,  species  density,  species  diversity,  and 
species  richness).  Often  this  is  the  only  useful  measure  of  diversity  available 
from  specimens  taken  for  taxonomic  purposes.  Bat  communities  (and  numbers 
of  phyllostomatid  species)  in  the  Americas  are  largest  in  tropical  lowland  rain 
forest.  Moving  away  from  that  life  zone  in  moisture,  altitude,  or  latitude,  the 
number  of  species  diminishes  (Fig.  5).  Beyond  this  common  observation, 
analysis  of  species  number  reveals  little  about  the  nature  of  bat  communities. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


425 


COOL  TEMPERATE 


Desert 

Desert  Scrub 

Steppe 

Moist  Forest 

Wet  Forest 

Rain  Forest 

0.89(6,2) 

0.05(71) 

1.32(75) 

99/0 

83/0 

100/0 

Desert 

2.2808,1) 

99/0 

Rain  Forest 

Desert 

Rain  Forest 

Desert 

2.04(12,2) 

93/6 

1.8502,3) 

86/0 

Rain  Forest 

Desert 

Rain  Forest 

WARM  TEMPERATE 

Desert 

Deserf  Scrub 
1.33(?1) 
76/0 

Thorn  Steppe 

165(10,1) 

91/0 

Dry  Forest 

Moist  Forest 
128(8,2) 
95/0 

Wet 

Forest 

Rain  Forest 

SUBTROPICAL 

Desert 

Desert  Scrub 

Thorn 

Woodland 

Dry  Forest 

Moist  Forest 

Wet 

Forest 

Rain  Forest 

SUBALPINE 


MONTANE 


LOWLAND 


TROPICAL 

Desert 

Desert 

— 

— 

0  80(3,1) 
0/100 

2  26(16,1) 
2/84 

2  60(1^1) 
5/89 

Rain  Forest 

Rain  Forest 

LOWER 

MONTANE 

PRE- 

MONTANE 

LOWLAND 

Desert 

Desert  Scrub 

Thorn 

Woodland 

Very  Dry 
Forest 

Dry  Forest 

169(18,5) 

1/95 

Moist  Forest 

2.13(18,8) 

6/85 

Wet  Forest 

214(20,6) 

5/94 

Rain  Forest 

2.36(29,1) 

1/99 

Fig.  5. — Characteristics  of  bat  community  structure  according  to  life  zone  (after 
Holdridge,  1967).  Numbers  are  :  top  line,  average  diversity  value,  (average  number  of 
species,  number  of  samples);  bottom  line,  proportion  of  the  diversity  contributed  by 
vespertilionids/phyllostomatids.  Because  no  data  are  available  from  boreal  or  subpolar 
latitudes  or  alpine  altitudes,  corresponding  life  zones  are  omitted.  All  samples  were 
mist-netted  in  tropical  wet  seasons,  temperate  summers,  or  year-round.  Although  the 
best  available,  these  samples  are  not  ideal  for  diversity  analysis.  Samples  vary  in  habitat 
(for  example,  mature  forest,  riparian  forest,  slash-and-burn  agriculture)  and  adequacy  of 
netting  vertical  strata  and  full  nights.  All  tropical  samples  inadequately  represent  high-flying 
molossids  and  taxa  more  difficult  to  net  than  phyllostomatids  (for  example,  emballonurids, 
mormoopids,  and  vespertilionids).  Summaries  and  references  to  sample  data  are  available  from 
the  senior  author  on  request. 


Species  Diversity 

More  can  be  learned  by  finding  a  concise  way  to  compare  communities 
and  the  abundance  of  species  within  and  among  communities.  Such  comparison 
is  afforded  by  a  species  diversity  index.  Details  of  rationale  and  application 
of  this  analytical  tool  to  bat  communities  are  presented  by  Humphrey  (1975). 
Briefly,  the  standard  index  is  that  of  Shannon  and  Weaver  (1949),  H'  = 
—  2 PilogePi ,  where  p{  is  the  number  of  individuals  in  the  i,h  species  divided  by 
sample  size.  The  contribution  of  species  n  to  its  community’s  diversity  is 
H'n=  ~  Pnl°SePn- 

Parallel  to  the  pattern  of  species  number,  species  diversity  (Fig.  5)  is 
highest  in  tropical  lowland  rain  forest  and  decreases  along  gradients  of  moisture, 
altitude,  and  latitude.  The  most  diverse  single  sample  (//'  =  2.65)  was  taken  in 
garden  and  forest  habitats  at  San  Pablo,  Peru  (Tuttle,  1970);  no  doubt  this  is 
an  overestimate,  as  data  from  two  habitats  are  pooled.  Average  diversity  of 


426 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


warm  temperate  montane  dry  forest  also  is  overestimated,  because  all  three 
sites  were  chosen  for  exceptional  diversity  in  topography  and  vegetation. 

Although  fairly  diverse  communities  continue  into  middle  latitudes  and 
zones  of  intermediate  moisture,  a  pronounced  shift  in  the  importance  of 
phyllostomatids  occurs  between  subtropical  and  warm  temperate  zones.  In 
warm  temperate  zones,  phyllostomatids  are  replaced  by  vespertilionids. 
Presence  of  two  species  of  nectar-feeding  phyllostomatids  in  warm  temperate 
montane  thorrujsteppe  ( Choeronycteris  mexicana  and  Leptonycteris  sanborni 
in  Arizona  samples)  results  from  migration  to  take  advantage  of  seasonally 
available  Agave  and  cactus  flowers.  Bat  communities  are  least  diverse  in  zones 
of  extreme  dryness  or  high  altitude  or  latitude.  These  correlations  with  the 
Holdridgean  axes  of  precipitation,  humidity,  and  temperature  suggest  that 
phyllostomatids,  as  a  family,  are  best  adapted  to  regions  where  1)  annual 
precipitation  exceeds  1000  mm,  or  2)  the  ratio  of  potential  evapotranspiration 
to  precipitation  is  less  than  two,  and  3)  a  mean  annual  biotemperature  about 
17°C  is  available  for  at  least  one  season  of  the  year  (as  by  migration).  The 
ultimate  factors  responsible  for  this  pattern  will  become  clear  as  the  functions 
of  the  morphological,  behavioral,  demographic,  and  physiological  adaptations 
of  these  bats  are  better  understood.  We  infer  that  the  pattern  represents 
phyllostomatid  response  either  directly  to  climate  or  to  biological  factors  such 
as  vegetation  or  food. 

The  general  lack  of  anomalies  in  diversity  trends  of  phyllostomatid- 
dominated  faunas  is  striking.  One  exception  is  in  tropical  lower  montane  dry 
forest,  where  small  sample  size  (10)  of  the  single  sample  may  account  for 
low  diversity.  By  contrast,  no  clear  life-zone  pattern  appears  in  diversity  of 
warm  and  cool  temperate  bat  faunas.  As  shown  by  Humphrey  (1975),  the 
presence  of  suitable  roosts  enables  strongly  roost-adapted  vespertilionids  and 
molossids  to  become  exceptionally  abundant  there.  A  super-abundant  species 
affects  the  diversity  value  because  H'n  1<  H'n 2,  lowering  H' .  Thus  for  roost-adapted 
taxa,  perhaps  including  the  tropical  mormoopids,  we  expect  such  factors  as 
karst  topography  and  forest  management  practices  to  be  of  primary  importance. 

Some  indication  of  the  importance  of  certain  species  to  their  bat  communities 
is  given  in  Figs.  6  and  7.  Consistently  important  species  in  lowland  forest  are 
the  feeding  generalists  Carollia  perspicillata  and  small  species  of  Artibeus, 
which  eat  a  wide  variety  of  fruits.  When  fruit  is  scarce,  C.  perspicillata  also 
will  consume  nectar,  pollen,  and  insects.  Other  generalists  such  as  species  of 
the  genus  Sturnira,  however,  are  consistently  minor  community  members. 
Specialists  on  large  fruit,  Artibeus  jamaicensis  and  species  of  Vampyrops,  do 
best  in  wet  climates  and  decrease  in  importance  in  drier  forests.  High  H' „ 
of  A.  jamaicensis  in  dry  forest  is  an  artifact  in  that  all  samples  there  were  in 
fruit  plantations  or  riparian  gallery  forest  that  included  many  fig  trees.  High 
importance  of  both  A.  jamaicensis  and  small  species  of  Artibeus  in  forest  of 
intermediate  moisture  accords  with  our  unpublished  data  that  these  bats  eat 
different  species  of  fruit,  partitioning  food  on  the  basis  of  particle  size. 
Glossophaga  soricina,  a  species  that  specializes  on  nectar  and  pollen  but 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


427 


TROPICAL  LOWLAND  FOREST 

Fig.  6. — Contribution  to  bat  community  diversity  by  species  of  phyllostomatids  along 
a  moisture  gradient  in  tropical  lowland  life  zones.  Data  are  from  samples  used  in  Fig.  5. 


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TROPICAL  WET  FOREST 

Fig.  7. — Contribution  to  bat  community  diversity  by  species  of  phyllostomatids 
along  an  altitudinal  gradient  in  tropical  wet  forest  life  zones.  In  effect,  this  graph  adds  a 
third  dimension,  elevation,  to  Fig.  6,  with  the  origin  at  the  point  “tropical  lowland  wet 
forest.” 

switches  to  fruit  in  the  wet  season  when  flowers  are  scarce,  is  of  complementary 
importance  to  those  bats  that  eat  large  fruits,  increasing  its  contribution  in 
dry  communities.  Trachops  cirrhosus,  a  specialist  on  insects  and  small  vertebrates, 
and  Phyllostomus  hastatus,  a  large  bat  that  eats  fruit,  insects,  and  some 
vertebrates,  are  predictably  unimportant;  we  do  not  understand  the  higher 
contribution  of  the  latter  in  wet  forest.  Desmodus  rotundus,  a  specialist  on 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


429 


mammal  blood,  is  a  minor  constituent  of  wet  zones  but  becomes  increasingly 
important  in  drier  forest,  probably  a  function  of  increasing  livestock  density. 

Altitudinal  data  (Fig.  7)  reinforce  the  general  conclusions  derived  from 
lowland  data.  Carollia  perspicillata  is  an  important  community  member  at 
all  elevations  sampled.  G.  soricina,  already  seen  to  be  a  minor  member  in  wet 
lowland  forest,  is  equally  unimportant  in  higher  forest.  The  large  fruit 
specialists  A.  jamaicensis  and  Vampyrops,  which  respond  similarly  to  moisture 
change  in  lowland  forest,  show  opposite  trends  at  high  altitudes  in  wet  forest  — 
prominence  of  Vampyrops  and  disappearance  of  A.  jamaicensis.  The  frugivorous 
generalists  of  the  genus  Sturnira  and  small  species  of  Artibeus  show  a  pattern 
similar  to  that  of  Vampyrops ,  except  that  small  Artibeus  decrease  in  importance 
at  the  highest  elevation.  Perhaps  this  pattern  represents  a  competition-based 
displacement  or  poor  response  by  small  kinds  of  Artibeus  to  the  greater  daily 
variation  of  climate  in  high  altitude  forests.  The  prominence  of  Sturnira  may 
indicate  some  special  adaptations  to  highlands  in  view  of  its  unimportance 
in  all  lowland  forests  sampled. 

Careful  work  that  samples  vertical  strata  and  accounts  for  differences  of 
habitat  and  season  will  reveal  much  more  about  tropical  bat  communities 
and  phyllostomatids.  For  example,  ground-level  nets  at  a  site  at  Belem, 
Brazil,  yielded  18  species  with  a  diversity  of  1.93;  simultaneous  netting  with 
subcanopy  and  canopy  nets  placed  above  the  ground  nets  added  seven  species 
of  phyllostomatids  and  increased  diversity  to  2.40  (Handley,  1967).  In  all-night 
netting,  LaVal  (1970,  personal  communication)  alternately  sampled  riparian 
forest  and  banana  groves  at  Finca  La  Pacifica,  near  Canas,  Costa  Rica.  A 
decrease  in  diversity  in  the  plantation  (1.63  as  opposed  to  1.84  in  the  riparian 
forest)  was  accompanied  by  marked  shifts  in  species  composition;  most  striking 
was  the  omnivorous  Phyllostomus  discolor,  rare  in  the  forest  but  by  far  the 
most  abundant  species  in  banana  groves,  where  presumably  it  ate  banana  nectar 
and  pollen.  In  an  area  of  slash-and-bum  agriculture  near  Frijoles,  Canal  Zone, 
D.  E.  Wilson  (personal  communication)  sampled  during  the  wet  season  (shortly 
after  the  main  pulse  of  births  for  the  year)  and  in  the  following  dry  season. 
Wet  season  diversity  was  1.93  with  14  species,  but  dry  season  values  dropped 
to  1.74  and  seven,  respectively;  all  species  lost  in  the  dry  season  were 
phyllostomatids.  Such  data  suggest  exciting  patterns  of  phyllostomatid 
specialization  in  foraging  strata  and  habitat  and  the  possibility  of  seasonal 
migration  or  nomadism  among  habitats. 

Ecosystem  Functions 

The  dynamics  of  energy  flow  are  receiving  increased  attention  by  ecologists. 
Producers  and  decomposers  are  recognized  as  the  important  organisms  in 
contributing  to  net  productivity.  Consumers  account  for  little  of  the  energy 
flowing  through  their  ecosystems  (Fleharty  and  Choate,  1973;  Fittkau  and 
Klinge,  1973).  However,  they  do  play  key  roles  in  directing  ecosystem 
dynamics.  Long-term  growth,  succession,  and  stability  of  plant  associations 
depend  partly  on  ecosystem  functions  performed  by  herbivores.  Examples  are 


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seed  dispersal  by  birds,  primates,  rodents,  and  bats;  pollination  by  bees,  moths, 
and  bats;  and  successional  retardation  by  voles,  prairie  dogs,  and  some  ungulates. 

Phyllostomatid  bats  fall  into  several  consumer  trophic  levels.  These  range 
from  the  second  level  for  fruit  and  nectar-pollen  consumers,  the  third  for 
insectivores  and  sanguivores  (most  of  the  time),  and  the  fourth  or  fifth  for 
carnivorous  taxa  such  as  Vampyrum  spectrum  and  Trachops  cirrhosus, 
which  often  eat  other  carnivores.  The  interactions  of  the  phyllostomatids  with 
other  organisms  and  the  ecosystem  functions  performed  by  phyllostomatids 
will  be  discussed  in  the  following  sections. 

Seed  Dispersal 

Phyllostomatid  bats,  birds,  primates,  and  rodents  are  the  most  important 
agents  of  seed  dispersal  in  the  Neotropical  region  (van  der  Pijl,  1972).  Few 
data  exist  for  shrubs,  vines,  or  epiphytes,  but  phyllostomatids  act  as  dispersal 
agents  for  up  to  24  per  cent  of  the  forest  tree  species  at  some  sites.  At  Finca  la 
Selva  near  Puerto  Viejo  in  Costa  Rica,  Gary  Hartshorn  made  the  following 
unpublished  observations  in  a  tropical  wet  forest  (4000  millimeters  of 
rain  per  year;  no  dry  season):  in  an  area  containing  20  species  of  fruit-eating 
bats,  24  per  cent  of  the  273  tree  species  counted  bore  bat-dispersed  seeds. 
F.  J.  Bonaccorso  (unpublished  data)  recorded  similar  information  for  a  tropical 
moist  forest  (rainfall,  2750  millimeters  per  year;  four  to  five  month  dry  season) 
on  Barro  Colorado  Island,  Panama:  7.7  per  cent  of  the  approximately  350 
tree  species  identified  carried  seeds  dispersed  by  bats;  16  species  of  fruit-eating 
bats  were  present.  In  both  of  these  works,  a  fruit-eating  bat  was  defined  as  one 
with  20  per  cent  or  more  of  its  diet  consisting  of  fruit;  trees,  as  being  greater  than 
10  centimeters  dbh  or  5  meters  tall.  These  observations  suggest  that 
phyllostomatids  become  increasingly  important  as  dispersal  agents  in  wetter 
forests,  as  evidenced  by  the  percentage  of  trees  dispersed  by  bats  and  increased 
number  of  frugivorous  bat  species.  Where  long  dry  seasons  occur,  persistent 
winds  disperse  the  seeds  of  many  trees.  At  sites  without  a  strong  dry  season 
and  associated  winds,  animals  (and  water)  play  a  major  role  in  dispersing  plant 
species.  Some  dry  forest  sites  may  support  higher  than  expected  numbers  of 
frugivorous  species  because  of  the  abundant  quantities  of  bat  flowers  during  the 
dry  season.  At  that  time,  these  usually  frugivorous  bats  switch  to  diets  of  nectar 
and  pollen  (Fleming  et  al. ,  1972;  Heithaus  et  al.,  1975). 

The  bat-fruit  syndrome. — The  relationship  between  bats  and  fruiting  plants 
is  mutualistic.  The  plants  expend  energy  on  production  of  edible,  nutritious 
fruits  as  well  as  on  olfactory  and  visual  stimuli  that  attract  bats.  In  eating  fruits, 
bats  usually  transport  seeds  away  from  the  parental  crown  and  discard  them 
at  potential  germination  sites. 

Fruit-eating  bats  and  bat  fruits  have  undergone  considerable  coevolution, 
and  the  resulting  set  of  adaptations  are  characterized  as  the  “bat-fruit  syndrome” 
(Table  1).  Exceptions  to  this  syndrome  occur,  but  when  several  or  all  characters 
of  the  syndrome  occur  in  a  fruit,  it  is  likely  to  be  dispersed  by  bats. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


431 


Table  1. —  The  bat-fruit  syndrome  (after  Pijl,  1972). 


Fruit  characteristics  Bat  characteristics 


1 .  Strong,  musty  odor 

2.  Dull  color,  often  green  or  brown,  fruits 
visually  inconspicuous 

3.  Exposed  position  outside  dense  foliage  on 
periphery  of  branches  or  on  pendulous  branches 

4.  Attachment  to  tree  through  maturity 

5.  Hard  skin  or  pulp  to  deter  other  frugivores 
(many  bat  fruits  are  soft  externally) 

6.  Requires  animal  agent  to  disperse  seed 


Good  sense  of  smell 
Large  eyes  for  orientation, 
probably  color-blind 
Approach  fruit  from  air 

Harvest  fruit  from  tree,  not  ground 
Strong  dentition  for  tearing  fruit 

Carry  fruits  from  fruiting 
trees  to  night  roosts 


Fruiting  plants  are  under  selective  pressure  to  attract  seed  dispersal  agents 
and  repel  or  temporally  avoid  seed  predators,  such  as  squirrels  and  peccaries 
that  frequently  eat  fruit  and  seeds.  Also,  some  animals  may  eat  fruits  yet 
neither  destroy  the  seeds  nor  disperse  them,  but  simply  discard  seeds  below 
the  parental  crown.  Ceboid  monkeys  often  discard  large  seeds  in  this  manner. 
Mature  fruits  on  the  tree  are  available  to  bats,  birds,  and  arboreal  animals, 
but  not  to  terrestrial  rodents,  deer,  and  peccaries.  Pendulant  fruits  of  tree 
species  such  as  Cecropia  (Fig.  2),  exclude  arboreal  rodents,  but  not  birds  or 
bats  that  pluck  fruits  in  flight,  nor  monkeys  that  hang  from  prehensile  tails 
and  arms,  nor  procyonids  or  primates  that  pull  branches  with  their  feet  (see 
Kaufmann,  1962).  The  strong  odor  of  fruits  attracts  olfactory-orienting  mammals, 
whereas  dull  coloration  camouflages  fruits  from  visually  orienting  birds. 
Fruits  having  hard  edible  parts  or  edible  parts  covered  by  a  tough  husk  exclude 
most  birds  but  do  not  hinder  fruit-eating  bats  with  their  strong  teeth. 

Seed  survival  and  mortality. — Once  a  plant  releases  its  fruit  to  a  dispersal 
agent,  the  chances  of  seed  mortality  are  high.  In  response,  many  plants  produce 
vast  numbers  of  seeds,  a  few  of  which  survive  losses  to  seed  predators,  parasites, 
mechanical  damage,  and  inhospitable  germination  sites.  For  seeds  to  be 

dispersed  successfully  by  bats,  the  following  conditions  are  requisites:  1) 

fruit  harvest  must  occur  at  maturity;  2)  seed  displacement  must  be  beyond  the 
crown  of  the  parent  plant  (in  some  species);  3)  seed  deposition  must  be  at  a 
site  suitable  for  germination;  and  4)  seeds  must  not  be  severely  damaged. 

In  our  experience,  frugivorous  phyllostomatids  select  ripe  fruit.  Several 
factors  promote  this  pattern.  Unripe  fruits  have  little  odor  and  would  not 
attract  bats.  Also  unripe  fruits  are  hard  and  difficult  to  chew,  and  some  are 
distasteful  or  toxic  when  immature.  For  example,  some  immature  fruits  of 

Passiflora  contain  deadly  cyanide  compounds  but  when  ripe  are  eaten  by 

mammals  (Saenz  and  Nassar,  1972).  Unripe  figs  contain  latex,  which  is 
gummy,  and  have  a  bad  taste  (to  humans,  at  least).  The  great  difference  in 
sugar  content  of  green  and  ripe  fruit  (Snow,  1971)  suggests  that  there  is  little 


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selective  advantage  in  exploiting  unripe  fruit.  These  mechanisms  effectively 
protect  seeds  until  they  are  mature  and  viable. 

Janzen  (1971  cr,  1971  b)  and  Wilson  and  Janzen  (1972)  demonstrated  that 
seed  predators  cause  heavy  mortality  of  seeds  falling  under  the  parental 
crown  because  the  foraging  or  egg-laying  strategy  is  to  locate  areas  of  high 
seed  density.  Seed  survival  frequently  is  related  to  the  seed’s  ability  to 
escape  predators  in  space.  Figs  that  are  not  taken  away  by  bats  or  other 
dispersors,  for  example,  are  susceptible  to  heavy  seed  predation  by  lygaeid 
bugs  (Slater,  1972). 

As  a  rule,  the  Phyllostomatidae  and  the  paleotropical  Pteropodidae  carry 
fruits  from  resource  trees  to  night  feeding  roosts  (Greenhall,  1956,  1965; 
Nellis,  1971;  Osmaston,  1965;  Jones,  1972),  which  may  change  every  few 
days  (D.  Morrison,  personal  communication).  Because  hundreds,  or  even 
thousands,  of  bats  may  come  to  large  fruiting  trees  in  a  single  night,  the  use 
of  feeding  roosts  may  alleviate  crowding  and  aggression  at  resource  trees. 
Additionally,  this  behavior  may  reduce  the  attractiveness  of  bat  feeding 
aggregations  to  predators  such  as  owls  and  opossums. 

Each  plant  species  has  particular  requirements  with  respect  to  soil,  nutrients, 
drainage,  and  lighting  conditions  conducive  to  subsequent  growth  and 
development.  Once  taken  to  a  feeding  roost,  seeds  are  either  discarded  as  the 
fruit  is  eaten  (somatochory)  or  ingested  with  fruit  pulp  (endochory). 
Somatochores  (Fig.  2C)  have  seeds  too  large  to  swallow,  and  their  dispersal 
by  phyllostomatids  is  limited  by  the  nature  and  location  of  night  feeding  roosts. 
Of  these,  caves  and  buildings  are  particularly  bad  places  for  seeds  to  germinate. 

Endochores  (Fig.  2 A,  B)  have  small,  numerous  seeds  scattered  through 
the  edible  pulp.  Endochores  are  eliminated  with  the  feces  and  have  the 
potential  to  land  any  place  a  bat  moves  during  a  night.  Alimentary  passage 
time  is  usually  less  than  three  hours  for  seeds  (Arata,  1972;  S.  Farkas, 
personal  communication)  and  typically  may  be  half  an  hour  (Klite,  1965).  Quick 
seed  passage  time  keeps  bats  at  low  flight  weights  and  also  ensures  that 
many  seeds  will  be  eliminated  before  the  bat  returns  to  the  day  roost,  which 
usually  is  a  poor  germination  site. 

Seed  damage  may  result  from  mastication  or  digestion.  Bats  rarely  damage 
small  seeds,  and  excreted  seeds  have  a  high  germination  rate  (S.  Gaulin, 
personal  communication).  We  know  of  only  one  species  of  large-seeded  fruit 
species,  Anacardium  excelsum,  regularly  damaged  by  bats.  A.  excelsum 
seeds  commonly  are  eaten  by  Carollia  perspicillata,  which  acts  as  a  seed 
predator. 


Pollination  and  the  Bat-Flower  Syndrome 

Based  on  floral  form,  Vogal  (1969)  estimated  that  bats  play  some  part 
in  pollination  of  at  least  500  Neotropical  plant  species  of  96  genera.  It  appears 
that  phyllostomatids  increase  in  importance  as  pollinating  agents  from  mesic 
to  xeric  habitats.  This  pattern  is  the  opposite  of  that  of  phyllostomatids  acting 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


433 


Table  2. —  The  hat-flower  syndrome  ( after  Faegri  and  Pijl,  1971). 


Flower  characteristics  Bat  characteristics 


1.  Nocturnal  anthesis 

2.  Strong,  musty  odor 

3.  Dull  color,  often  whitish,  creamy,  or  purple 

4.  Exposed  position  outside  dense  foliage  on 
periphery  of  branches  or  on  pendulous  branches 

5.  Large  flowers 

6.  Copious  nectar  and  pollen  production 

7.  Flower  tube-like  with  anthers  protruding, 
or  brush-shaped  flower 


Nocturnal  foraging 
Good  sense  of  smell 
Large  eyes  for  orientation, 
probably  color-blind 
Approach  flower  from  air 

Large  body  size  compared  to 
other  pollinators 

High  metabolic  rate  and  large  body 
size 

Elongate  snout  and  protrusible  tongue 
for  probing  deep  into  flowers 


as  seed  dispersal  agents.  Glossophaga  soricina  in  Panama  uses  flower  resources 
for  only  five  months  of  the  year  (Fleming  et  al.,  1972),  whereas  in  the  dry 
forest  of  Costa  Rica,  G.  soricina  uses  flowers  all  year  long  (Heithaus  et  al ., 
1975). 

Flowers  pollinated  by  bats  are  distinguished  by  drab  colors,  musty  odors, 
tube  or  brush  shapes,  position  free  of  foliage,  large  size,  nocturnal  anthesis, 
and  copious  nectar  and  pollen  production.  These  floral  characters  (Fig.  2) 
and  corresponding  adaptations  found  in  nectarivorous  bats  are  summarized 
in  Table  2.  Caution  must  be  taken  because  bats  are  highly  opportunistic  in 
foraging  habits  and  may  sometimes  take  advantage  of  flowers  not  precisely 
fitting  the  “bat-flower  syndrome.”  Furthermore,  bats  may  ingest  nectar  or 
pollen  (or  both)  of  a  particular  plant  species  and  yet  not  provide  pollination 
services.  Floral  parasites  are  common  in  nature  (for  example,  flower-piercing 
hummingbirds  and  bees).  Ratcliffe  (1931)  reported  that  flying  foxes  of 
Australia  eat  entire  flowers,  but  we  have  found  no  reports  of  phyllostomatids 
regularly  eating  flowers.  Baker  et  al.  (1971)  suggested  that  Leptonycteris 
sanborni  occasionally  eats  anthers. 

The  association  between  these  bats  and  flowering  plants  is  mutualistic. 
Plants  divert  energy  into  production  of  odors  and  floral  parts  that  attract 
bats  as  well  as  nectar  and  pollen  that  feed  bats.  In  moving  from  flower  to  flower 
for  food,  bats  transport  some  pollen,  which  results  in  fertilization. 

It  has  long  been  obvious  that  flower  bats  obtain  carbohydrate  in  the  form  of 
sugars  from  floral  nectaries.  Recently  it  has  been  demonstrated  that  pollen  is 
an  important  source  of  protein  to  these  bats  (Howell,  1974).  The  cellular 
contents  of  pollen  grains  begin  to  extrude  through  the  micropores  when  pollen 
begins  to  germinate  in  the  gut.  Protein  then  is  extracted  by  hydrochloric  acid 
produced  in  the  stomach,  and  protein  is  leached  further  by  urea  from  urine 
ingested  by  the  bat  (at  least  in  Leptonycteris  sanborni).  Howell  also  found  that 
the  protein  content  of  pollen  eaten  by  L.  sanborni  was  44  per  cent  for  saguaro 
and  23  per  cent  for  paniculate  agave,  much  higher  than  in  pollen  of  closely 


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related  plants  for  which  pollen  is  not  eaten  or  dispersed  by  bats.  Inasmuch 
as  herbivores  must  have  a  rich  source  of  plant  protein  in  order  to  maintain  a 
high  rate  of  metabolism,  the  concentration  of  protein  in  pollen  may  be  an 
important  avenue  of  coevolution. 

If  flower  bats  eat  pollen,  then  how  do  they  function  as  pollinators?  While 
probing  the  flower  corolla  for  nectar,  bats  become  dusted  with  pollen  from 
noseleaf  to  uropatagium  (Baker,  1970).  Phyllostomatids  eat  pollen  only  as  they 
groom  their  flight  membranes  and  fur  after  a  foraging  bout  (Howell  and  Hodgkin, 
1976;  Heithaus  et  al.,  1975).  Such  behavior  would  provide  for  floral  pollination 
during  visits  to  successive  flowers  while  foraging  and  still  permit  the  bat  later 
to  eat  the  excess  pollen  covering  its  body. 

Some  bats  adapted  to  the  exploitation  of  flowers  are  known  to  feed  on  insects 
and  fruits.  Whether  insects  are  taken  in  the  process  of  nectar-feeding  or  hunted 
separately  is  unknown. 


Impact  of  Predation  by  Bats 

Little  is  known  of  the  precise  diets  of  insect  or  vertebrate-eating  phyllostomatids, 
so  the  impact  of  their  predation  on  prey  populations  is  undocumented.  The 
prominence  of  biotic  limiting  factors  in  the  Neotropics  suggests  that  investigation 
of  this  impact  is  worthwhile. 


Competition 

Competition  for  sunlight  or  food  is  thought  to  be  the  dominant  limiting 
factor  for  Neotropical  organisms  (see  Janzen,  1967,  and  MacArthur,  1969). 
The  likelihood  of  interspecific  competition  in  tropical  bats  has  been  discussed 
(Tamsitt,  1967;  McNab,  1971;  Dwyer,  1971;  Fleming  et  al.,  1972;  Howell 
and  Burch,  1974;  Heithaus  et  al.,  1975),  though  documentation  of  such 
competition  awaits  further  study.  Intraspecific  competition  for  both  food  and 
roost  space  are  most  likely  to  occur  in  the  most  colonial  phyllostomatids, 
mentioned  above. 

Enough  is  known  about  the  three  main  categories  of  phyllostomatid  food  — 
fruit,  nectar  and  pollen,  and  insects  —  to  discuss  them  briefly.  Potential 
competitors  in  all  three  categories  include  insects,  birds,  arboreal  mammals, 
and  other  bats,  plus  insectivorous  spiders.  Observations  cited  above  of  times 
when  food  may  be  in  short  supply  suggest  when  competition  could  be  acute. 

Fruit. — Because  fruits  are  available  on  a  “first  come,  first  served”  basis, 
fruit  searching  success  may  be  an  important  component  of  potential  competition. 
Birds  feed  heavily  on  ripe  fruits  of  species  eaten  by  bats.  Two  of  the  most 
important  bat  fruits  in  Trinidad — Cecropia  and  Piper  (Fig.  2) — are  eaten  in 
significant  quantities  by  tanagers  and  honeycreepers  (Snow  and  Snow,  1971).  For 
example,  Eisenmann  (1961)  recorded  24  species  of  birds  feeding  on  Cecropia  fruit 
in  Panama.  Potential  mammalian  competitors  include  monkeys,  marsupials, 
rodents,  and  procyonids.  Monkeys  may  be  especially  important  in  eating  large 
quantities  of  unripe  fruit  (Daubenmire,  1972). 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


435 


Nectar  and  pollen. — Diurnal  birds  and  insects  seldom  compete  with  bats 
for  nectar  and  pollen,  because  many  flowers  are  adapted  for  pollination 
at  one  time  of  day  and  might  not  be  open  or  produce  nectar  at  other  times  (for 
example,  Bauhinia  pauletia,  Heithaus  et  al.,  1974;  Janzen,  1968).  Thus,  most 
flowers  used  by  Neotropical  hummingbirds  (Wolf,  1970;  Snow  and  Snow, 
1972)  are  not  visited  by  bats.  However,  Baker  et  al.  (1971)  have  shown  that 
Ceiba  acuminata  may  be  pollinated  by  both  bats  and  hummingbirds,  because 
these  flowers  open  at  night  but  continue  to  secrete  nectar  the  next  day  and  are 
visited  by  both  kinds  of  animals.  Balsa  ( Ochroma )  flowers  open  at  dusk 
(Fig.  2)  and  are  visited  by  phyllostomatid  bats  and  sphingid  moths.  Some 
flowers  used  by  bats  are  destroyed  when  monkeys  or  insects  eat  them.  Alvarez 
and  Gonzalez  Q.  (1970)  concluded  that  little  or  no  competition  for  flowers 
occurs  among  six  genera  of  glossophagines  in  Mexico.  Heithaus  et  al.  (1975) 
also  found  nectarivorous  bats  to  feed  as  generalists  with  high  dietary  overlap. 

Insects. — Seasonally  rapid  recruitment  rates  for  insects  (as  when  a  hatch 
is  under  way)  may  enable  bats  to  partition  temporally  a  common  resource 
during  the  night.  Additional  partitioning  is  possible  because  insect  taxa  differ 
in  periodicity  of  night-time  activity,  at  least  in  temperate  zones  (Williams, 
1935,  1939;  Lewis  and  Taylor,  1965).  Potential  night-time  competitors 
include  spiders,  tree  frogs,  caprimulgiform  birds,  owls,  night  monkeys, 
marsupials,  rodents,  and  procyonids. 

Roost  space. — We  know  of  no  published  evidence  of  phyllostomatids 
competing  for  roost  space.  However,  such  competition  frequently  may  be 
provided  by  the  more  colonial  taxa.  On  Barro  Colorado  Island,  Panama,  a 
group  of  Phyllostomus  hastatus  displaced  a  hollow  tree  colony  of  Carollia 
perspicillata  (S.  Graetz,  personal  communication).  F.  J.  Bonaccorso  observed, 
on  the  same  island,  a  displacement  of  C.  perspicillata  and  Saccopteryx  bilineata 
from  their  hollow  tree  roost  by  a  colony  of  Desmodus  rotundas. 

Predation  on  Bats 

Little  is  known  about  causes  of  phyllostomatid  mortality  or  the  food  habits 
of  their  potential  predators.  Reviewers  of  temperate  zone  data  judge  that 
predation  on  bats  is  opportunistic  but  seldom  regular  (Allen,  1939;  Gillette 
and  Kimbrough,  1970).  In  the  New  World  tropics,  predation  on  bats  may 
well  be  more  important,  in  view  of  the  general  prominence  of  biotic  interactions 
and  the  dominant  numbers  of  bats  in  mammal  faunas.  In  Haiti,  27  of  147  prey 
items  of  a  Hispanolean  barn  owl  ( Tyto  glaucops)  were  phyllostomatid  bats 
(Wetmore  and  Swales,  1931).  In  Panama,  three  species  of  owls  have  killed 
bats  in  our  nets.  Arboreal  opossums,  procyonids,  and  snakes  may  wait  for 
bats  visiting  resource  trees.  Opossums  ( Didelphis  virginiana  and  Philander 
opossum)  are  known  to  eat  bats  (Campbell,  1925;  Rice,  1957;  our  observations). 
The  bat  falcon,  Falco  rufigularis,  may  specialize  on  bat  prey.  The  largest 
phyllostomatid  bats  ( V ampyrum  spectrum,  Chrotopterus  auritus,  and 
Phyllostomus  hastatus)  are  suspected  or  known  to  eat  smaller  bats  (Goodwin 
and  Greenhall,  1961 ;  Valdivieso,  1964;  Greenhall,  1968). 


436 


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An  important  clue  to  the  role  of  predation  on  phyllostomatids  may  be 
the  inverse  relationship  between  moonlight  and  flight  activity  of  Desmodus 
rotundus  (Crespo  et  al. ,  1972;  Turner,  1975)  and  Artibeus  jamaicensis 
(Morrison,  1975).  Our  qualitative  observations  at  several  Central  American 
sites  are  that  phyllostomatid  foraging  is  characteristically  maximal  when 
no  moonlight  is  incident  and  minimal  under  full  moonlight.  To  explain 
avoidance  of  moonlight  as  a  response  to  heightened  predator  success  would 
be  uncomplicated  for  bats  that  feed  on  plants;  the  behavior  of  predatory  bats 
must  additionally  account  for  the  possibility  of  similar  responses  by  their  own 
prey  species. 


Acknowledgments 

We  appreciate  the  helpful  criticisms  of  R.  B.  Foster,  D.  J.  Howell,  E.  Leigh, 
J.  J.  McManus,  P.  A.  Opler,  and  C.  M.  Simon,  and  the  art  work  of  N.  Halliday 
and  S.  J.  Scudder.  Contributions  of  unpublished  data  by  G.  W.  Frankie,  E.  R. 
Heithaus,  D.  J.  Howell,  B.  K.  McNab,  D.  Morrison,  and  N.  Smythe  have 
enabled  us  to  explain  much  more  tropical  ecology  than  is  possible  from  the 
presently  available  literature.  Our  own  data  were  acquired  under  the  following 
grant  support  :  National  Institutes  of  Health  Biomedical  Sciences  grant  no. 
RR  7021-07  from  the  University  of  Florida  Division  of  Sponsored  Research 
to  S.  R.  Humphrey;  National  Science  Foundation  grant  no.  GB-36068  to 
J.  H.  Kaufmann;  and  Smithsonian  Tropical  Research  Institute  grants  to  F.  J. 
Bonaccorso. 


Literature  Cited 

Allee,  W.  C.  1926.  Measurement  of  environmental  factors  in  the  tropical  rain-forest  of 
Panama.  Ecology,  7:273-301. 

Allen,  G.  M.  1939.  Bats.  Dover  Publ.,  New  York,  368  pp. 

Allen,  L.  H.,  Jr.,  E.  Lemon,  and  L.  Muller.  1972.  Environments  of  a  Costa  Rican 
forest.  Ecology,  53: 102-1 1 1. 

Allen,  P.  H.  1956.  The  rainforests  of  Golfo  Dulce.  Univ.  Florida  Press,  Gainesville, 
417  pp. 

Alvarez,  T„  and  L.  Gonzalez  Q.  1970.  Analis  polinico  del  contenide  gastrico  de 
murcielagos  Glossophaginae  de  Mexico.  An.  Esc.  Nac.  Cien.  Biol.,  Mexico, 
18:137-165. 

Arata,  A.  A.  1972.  Thermoregulation  in  Colombian  Artibeus  lituratus  (Chiroptera). 
Mammalia,  36:86-92. 

Arata,  A.  A.,  and  C.  Jones.  1967.  Homeothermy  in  Carollia  (Phyllostomatidae: 

Chiroptera)  and  the  adaptation  of  poikilothermy  in  insectivorous  northern 
bats.  Lozania,  14: 1-10. 

Baker,  H.  1970.  Two  cases  of  bat  pollination  in  Central  America.  Rev.  Biol.  Trop., 
17:187-197. 

Baker,  H.,  R.  W.  Cruden,  and  I.  Baker.  1971.  Minor  parasitism  in  pollination 
biology  and  its  community  function:  the  case  of  Ceiba  acuminata.  BioScience, 
21:1127-1129. 

Baker,  J.  R„  and  Z.  Baker.  1936.  The  seasons  in  a  tropical  rain-forest  (New  Hebrides). 
Part  3.  Fruit-bats  (Pteropidae).  J.  Linnean  Soc.  London,  40:123-141. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


437 


Barkalow,  F.  S.,  Jr.,  R.  B.  Hamilton,  and  R.  F.  Soots,  Jr.  1970.  The  vital  statistics 
of  an  unexploited  gray  squirrel  population.  J.  Wildlife  Mgt.,  34:489-500. 

Barlow,  J.  C.,  and  J.  R.  Tamsitt.  1968.  Twinning  in  American  leaf-nosed  bats 
(Chiroptera:  Phyllostomatidae).  Canadian  J.  Zool.,  46:290-292. 

Bates,  M.  1945.  Observations  on  climate  and  seasonal  distribution  of  mosquitoes  in 
eastern  Colombia.  J.  Animal  Ecol.,  14:17-25. 

Baynton,  H.  W„  H.  L.  Hamilton,  Jr.,  P.  E.  Sherr,  and  J.  J.  B.  Worth.  1965.  Temperature 
structure  in  and  above  a  tropical  forest.  Quart.  J.  Royal  Meteorol.  Soc., 
91:225-232. 

Bonaccorso,  F.  J.  1975.  Foraging  and  reproductive  ecology  in  a  community  of  bats  in 
Panama.  Unpublished  Ph.D.  Dissertation,  Univ.  Florida,  1 19  pp. 

Bradbury,  J.  W.,  and  S.  L.  Vehrencamp.  1976.  Social  organization  and  foraging  in 
emballonurid  bats.  I.  Field  studies.  Behav.  Ecol.  Sociobiol.,  1:337-381. 

Bradshaw,  G.  V.  R.  1962.  Reproductive  cycle  of  the  California  leaf-nosed  bat, 
Macrotus  californicus.  Science,  136:645-646. 

Burns,  R.  J.  1970.  Twin  vampire  bats  born  in  captivity.  J.  Mamm.,  51:391-392. 

Campbell,  C.  A.  R.  1925.  Bats,  mosquitoes,  and  dollars.  The  Stratford  Co.,  Boston, 
vii  +  262  pp. 

Carpenter,  R.  E.,  and  J.  B.  Graham.  1967.  Physiological  responses  to  temperature 
in  the  long-nosed  bat,  Leptonycteris  sanbornu  Comp.  Biochem.  Physiol., 
22:709-722. 

Carter,  D.  C.  1970.  Chiropteran  reproduction.  Pp.  233-246,  in  About  bats 
(B.  H.  Slaughter  and  D.  W.  Walton,  eds.),  Southern  Methodist  Univ.  Press, 
Dallas,  vii  +  339  pp. 

Caughley,  G.  1966.  Mortality  patterns  in  mammals.  Ecology,  47:906-91 8. 

Christian,  J.  J.  1971.  Fighting,  maturity,  and  population  density  in  Microtus 

Pennsylvania is.  J.  Mamm.,  52:556-567. 

Cockrum,  E.  L.  1955.  Reproduction  in  North  American  bats.  Trans.  Kansas 

Acad.  Sci.,  58:487-511. 

Cockrum,  E.  L.,  and  E.  Ordway.  1959.  Bats  of  the  Chiricahua  Mountains,  Chochise 
County,  Arizona.  Amer.  Mus.  Novit.,  1938:1-35. 

Crespo,  R.  F.,  S.  B.  Linhart,  R.  J.  Burns,  and  G.  C.  Mitchell.  1972.  Foraging 
behavior  of  the  common  vampire  bat  related  to  moonlight.  J.  Mamm., 
52:366-368. 

Croat,  T.  B.  1974.  A  case  for  selection  for  delayed  fruit  maturation  in  Spondias 
(Anacardiaceae).  Biotropica,  6:135-137. 

Dalquest,  W.  W„  and  D.  W.  Walton.  1970.  Diurnal  retreats  of  bats.  Pp.  162-187, 
in  About  bats  (B.  H.  Slaughter  and  D.  W.  Walton,  eds.),  Southern  Methodist 
Univ.  Press,  Dallas,  vii  +  339  pp. 

Daubenmire,  R.  1972.  Phenology  and  other  characteristics  of  tropical  semi-deciduous 
forest  in  north-western  Costa  Rica.  J.  Ecol.,  60:147-160. 

Dwyer,  P.  D.  1971.  Temperature  regulation  and  cave-dwelling  in  bats:  an  evolutionary 
perspective.  Mammalia,  35:424-455. 

Easterla,  D.  A.  1972.  Status  of  Leptonycteris  nivalis  (Phyllostomatidae)  in  Big 
Bend  National  Park,  Texas.  Southwestern  Nat.,  17:287-292. 

Eisenmann,  E.  1961.  Favorite  foods  of  Neotropical  birds:  flying  termites  and 
Cecropia  catkins.  Auk,  78:636-638. 

Faegri,  K.,  and  L.  van  der  Pijl.  1971.  The  principles  of  pollination  ecology.  Pergamon 
Press,  Oxford,  291  pp. 

Fittkau,  E.  J.,  and  H.  Klinge.  1973.  On  biomass  and  trophic  structure  of  the 
central  Amazonian  rain  forest  ecosystem.  Biotropica,  5:2-14. 

Fleharty,  E.  D„  and  J.  R.  Choate.  1973.  Bioenergetic  strategies  of  the  cotton 
rat,  Sigmodon  hispidus.  J.  Mamm.,  54:680-692. 


438 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


Fleming,  T.  H.  1971.  Artibeus  jamaicensis :  delayed  embryonic  development  in  a 
Neotropical  bat.  Science,  171:402-404. 

Fleming,  T.  H„  E.  T.  Hooper,  and  D.  E.  Wilson.  1972.  Three  Central  American 
bat  communities:  structure,  reproductive  cycles,  and  movement  patterns.  Ecol¬ 
ogy,  53:555-569. 

Fleming,  T.  H.,  E.  R.  Heithaus,  and  W.  B.  Sawyer.  1977.  An  experimental  analysis  of  the 
food  location  behavior  of  frugivorous  bats.  Ecology,  58:619-627. 

Foster,  R.  B.  1973.  Seasonality  of  fruit  production  and  seed  fall  in  a  tropical 
forest  ecosystem  in  Panama.  Unpublished  Ph.D.  dissertation,  Duke 
University,  156  pp. 

Fournier,  L.  A.,  and  S.  Salas.  1966.  Algunas  observaciones  sobre  la  dinamica  de 
la  floracion  en  el  bosque  tropical  humedo  de  Villa  Colon.  Rev.  Biol. 
Trop.,  14:75-85. 

Frankie,  G.  W„  H.  G.  Baker,  and  P.  A.  Opler.  1975.  Comparative  phenological 
studies  of  trees  in  Tropical  Wet  and  Dry  Forests  in  the  lowlands  of  Costa  Rica. 
J.  Ecol.,  62:881-909. 

Gardner,  A.  L.  1977.  Feeding  habits.  Pp.  293-350,  in  Biology  of  bats  of  the  New  World 

family  Phyllostomatidae.  Part  II.  Spec.  Publ.  Mus.,  Texas  Tech  Univ., 
1 3: 1-364  pp. 

Gillette,  D.  D.,  and  J.  D.  Kimbrough.  1970.  Chiropteran  mortality.  Pp.  262-283, 
in  About  bats  (B.  H.  Slaughter  and  D.  W.  Walton,  eds.),  Southern  Methodist 
Univ.  Press,  Dallas,  vii  +  339  pp. 

Goodwin,  G.  G.,  and  A.  M.  Greenhall.  1961.  A  review  of  the  bats  of  Trinidad  and 
Tobago.  Bull.  Amer.  Mus.  Nat.  Hist.,  122:191-301. 

Greenhall,  A.  M.  1956.  The  food  of  some  Trinidad  fruit  bats  ( Artibeus  and 
Caroll ia).  J.  Agric.  Soc.  Trinidad  and  Tobago,  869:1-23. 

- .  1965.  Sapucaia  nut  dispersal  by  greater  spear-nosed  bats  in  Trinidad. 

Caribbean  J.  Sci.,  5:167-171. 

- .  1968.  Notes  on  the  behavior  of  the  false  vampire  bat.  J.  Mamm., 

49:337-340. 

Hales,  W.  B.  1949.  Micrometeorology  in  the  tropics.  Bull.  Meteorol.  Soc.,  30:124-137. 

Handley,  C.  O.,  Jr.  1966.  Checklist  of  the  mammals  of  Panama.  Pp.  753-795, 
in  Ectoparasites  of  Panama  (R.  L.  Wenzel  and  V.  J.  Tipton,  eds.),  Field  Mus. 
Nat.  Hist.,  Chicago,  xii  +  861  pp. 

- .  1967.  Bats  of  the  canopy  of  an  Amazonian  forest.  Atas  do  Simposio  sobre 

a  biota  Amazonica,  5:21 1-215. 

Hayward,  B.  J.,  and  E.  L.  Cockrum.  1971.  The  natural  history  of  the  western  long- 
nosed  bat,  Leptonycteris  sanborni.  WRI-SCI  (Western  New  Mexico  Univ.), 
1:75-123. 

Heithaus,  E.  R.,  P.  A.  Opler,  and  H.  G.  Baker.  1974.  Bat  activity  and  pollination  of 
Bauhinia  pauletia:  Plant-pollinator  coevolution.  Ecology,  55:412-419. 

Heithaus,  E.  R.,  T.  H.  Fleming,  and  P.  A.  Opler.  1975.  Foraging  patterns  and 
resource  utilization  in  seven  species  of  bats  in  a  seasonal  tropical  forest.  Ecology, 
56:841-854. 

Herreid,  C.  F.,  Jr.  1967.  Mortality  statistics  of  young  bats.  Ecology,  48:310-312. 

Holdridge,  L.  R.  1967.  Life  zone  ecology.  Tropical  Science  Center,  San  Jose, 
Costa  Rica,  206  pp. 

Howell,  D.  J.  1972.  Physiological  adaptations  in  the  syndrome  of  chiropterophily 
with  emphasis  on  the  bat  Leptonycteris  Lydekker.  Unpublished  Ph.D. 
dissertation,  Univ.  Arizona,  217  pp. 

- .  1974.  Bats  and  pollen:  physiological  aspects  of  the  syndrome  of  chiropterophily. 

Comp.  Biochem.  Physiol.,  48A:263-276. 

Howell,  D.  J.,  and  D.  Burch.  1974.  Food  habits  of  some  Costa  Rican  bats.  Rev. 
Biol.  Trop.,  21:281-294. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


439 


f 


J 


Howell,  D.  J.,  and  N.  Hodgkin.  1976.  Feeding  adaptations  in  the  hairs  and  tongues 
of  nectar-feeding  bats.  J.  Morph.,  148:329-336. 

Humphrey,  S.  R.  1975.  Nursery  roosts  and  community  diversity  of  Nearctic  bats. 
J.  Mamm.,  56:321-346. 

Humphrey,  S.  R.,  and  J.  B.  Cope.  1970.  Population  samples  of  the  evening  bat, 
Nycticeius  humeralis.  J.  Mamm.,  51:399-401. 

- .  1976.  Population  ecology  of  the  little  brown  bat,  Myotis  lucifugus,  in 

Indiana  and  north-central  Kentucky.  Amer.  Soc.  Mamm.  Spec.  Publ.,  4:1-81. 

Immelmann,  K.  1971.  Ecological  aspects  of  periodic  reproduction.  Pp.  341-389, 
in  Avian  biology  (D.  E.  Farner  and  J.  R.  King,  eds.),  Academic  Press,  New  York, 
l:xix  +  586  pp. 

Janzen,  D.  H.  1967.  Synchronization  of  sexual  reproduction  of  trees  within  the 
dry  season  in  Central  America.  Evolution,  21:620-637. 

- .  1968.  Reproductive  behavior  in  the  Passifloraceae  and  some  of  its  pollinators 

in  Central  America.  Behavior,  32:33-48. 

- .  1 97 1  zz.  The  fate  of  Scheelea  rostrata  fruits  beneath  the  parent  tree:  predispersal 

attack  by  bruchids.  Principes,  15:89-101. 

- .  19716.  Escape  of  Cassia  grand  is  L.  beans  from  predators  in  time  and  space. 

Ecology,  52:964-979. 

- .  1973.  Sweep  samples  of  tropical  foliage  insects:  effects  of  seasons,  vegetation 

types,  elevation,  time  of  day,  and  insularity.  Ecology,  54:687-708. 

Janzen,  D.  H.,  and  T.  W.  Schoener.  1968.  Differences  in  insect  abundance  and 
diversity  between  wetter  and  drier  sites  during  a  tropical  dry  season.  Ecology, 
49:96-110. 

Janzen,  D.  H.,  G.  A.  Miller,  J.  Hackforth-Jones,  C.  M.  Pond,  K.  Hooper,  and  D.  P.  Janos. 

1976.  Two  Costa  Rican  bat-generated  seed  shadows  of  Andira  inermis  (Legumi- 
nosae).  Ecology,  57:1068-1075. 

Jones,  C.  1972.  Comparative  ecology  of  three  pteropid  bats  in  Rio  Muni,  West  Africa. 
J.  Zool.,  London,  167:353-370. 

Kaufmann,  J.  H.  1962.  Ecology  and  social  behavior  of  the  coati  ( Nasua  narica)  on 
Barro  Colorado  Island,  Panama.  Univ.  California  Publ.  Zool.,  60:95-222. 

Klite,  P.  D.  1965.  Intestinal  bacterial  flora  and  transit  time  of  three  neotropical 
bat  species.  J.  Bacteriol,  90:375-379. 

LaVal,  R.  K.  1970.  Banding  returns  and  activity  periods  of  some  Costa  Rican  bats. 
Southwestern  Nat.,  15:1-10. 

LaVal,  R.  K.,  and  H.  S.  Fitch.  1977.  Structure,  movements  and  reproduction  in  three 
Costa  Rican  bat  communities.  Occas.  Papers  Mus.  Nat.  Hist.,  Univ.  Kansas, 
69:1-28. 


Lewis,  T.,  and  L.  R.  Taylor.  1965.  Diurnal  periodicity  of  flight  by  insects.  Trans. 
Royal  Entomol.  Soc.,  116:393-479. 

Liat,  L.  B.  1970.  Food  habits  and  breeding  cycle  of  the  Malaysian  fruit-eating  bat, 
Cynopterus  brachyotis.  J.  Mamm.,  51:174-177. 

Linhart,  S.  B.  1973.  Age  determination  and  occurrence  of  incremental  growth 
lines  in  the  dental  cementum  of  the  common  vampire  bat  ( Desmodus  rotundas). 
J.  Mamm.,  54:493-496. 

Lord,  R.  D.,  F.  Muradali,  and  L.  Lazaro.  1976.  Age  composition  of  vampire  bats 
( Desmodus  rotundas)  in  northern  Argentina  and  southern  Brazil.  J.  Mamm., 
57:573-575. 

MacArthur,  R.  H.  1969.  Patterns  of  communities  in  the  tropics.  Biol.  J.  Linnean 
Soc.,  1:19-30. 

- .  1972.  Geographical  ecology.  Harper  and  Row,  New  York,  xviii  +  269  pp. 

McNab,  B.  K.  1969.  The  economics  of  temperature  regulation  in  Neotropical  bats. 
Comp.  Biochem.  Physiol.,  31:227-268. 

- .  1971.  The  structure  of  tropical  bat  faunas.  Ecology,  52:352-358. 


440 


SPECIAL  PUBLICATIONS  MUSEUM  TEXAS  TECH  UNIVERSITY 


- .  1973.  Energetics  and  the  distribution  of  vampires.  J.  Mamm.,  54:131-144. 

- .  1976.  Seasonal  fat  reserves  of  bats  in  two  tropical  environments.  Ecology, 

57:332-338. 

Mares,  M.  A.,  and  D.  E.  Wilson.  1971.  Bat  reproduction  during  the  Costa  Rican 
dry  season.  BioScience,  2 1 :47 1  -477. 

McManus,  J.  J.  1977.  Thermoregulation.  Pp.  281-292,  in  Biology  of  bats  of  the 
New  World  family  Phyllostomatidae.  Part  II  (R.  J.  Baker,  J.  K.  Jones,  Jr., 
and  D.  C.  Carter,  eds.).  Spec.  Publ.  Mus.,  Texas  Tech  Univ.,  13:1-364. 

Miguela,  P.  1969.  Bioenergetics  of  pregnancy  and  lactation  in  European  common 
vole.  Acta  Theriol.,  14:167-179. 

Morrison,  D.  W.  1975.  The  foraging  behavior  and  feeding  ecology  of  a  Neotropical  fruit 
bat,  Artibeus  jamaicensis.  Unpublished  Ph.D.  dissertation,  Cornell  Univ.,  Ithaca, 

101  pp. 

Mutere,  F.  A.  1968.  The  breeding  biology  of  the  fruit  bat.  Rouse tt us  aegyptiacus 
E.  Geoffroy  living  at  0°  22'S.  Acta  Tropica,  25:97-108. 

- .  1970.  The  breeding  biology  of  equatorial  vertebrates:  Reproduction  in  the 

insectivorous  bat,  Hipposideros  caffer,  living  at  0°  27 'N.  Bijd.  Dierk.,  40:56-58. 

Nellis,  D.  1971.  Additions  to  the  natural  history  of  Brachyphylla  (Chiroptera).  Car¬ 
ibbean  J.  Sci.,  11:91. 

Osmaston,  H.  A.  1965.  Pollen  and  seed  dispersal  in  Cblorophora  excelsa  and  other  Mor- 
aceae,  and  in  Parkin  filicoidea  (Mimosaceae),  with  special  reference  to  the  role  of  the 
fruit  bat,  Eidolon  helvum.  Commonwealth  Forest.  Rev.,  44:96-103. 

Pijl,  L.  van  der.  1972.  Principles  of  dispersal  in  higher  plants.  Springer-Verlag,  Berlin, 
2nd  ed.,  161  pp. 

Ratcliffe,  F.  N.  1931.  The  flying  fox  ( Pteropus )  in  Australia.  CSIRO  Bull.,  53:1-81. 

Rice,  D.  W.  1957.  Life  history  and  ecology  of  Myotis  austroriparius  in 
Florida.  J.  Mamm.,  38:15-32. 

Richards,  P.  W.  1973.  The  tropical  rain  forest.  Sci.  Amer.,  229:58-67. 

Robinson,  D.  1971.  Costa  Rican  mammals.  Pp.  1-6,  in  Handbook  for  tropical  biology 
in  Costa  Rica.  (C.  E.  Schnell,  ed.),  Organization  for  Tropical  Studies,  San  Jose, 
Costa  Rica. 

Ross,  A.  1967.  Ecological  aspects  of  the  food  habits  of  insectivorous  bats.  Proc.  Western 
Found.  Vert.  Zool.,  1:205-263. 

Route,  C.  S.,  and  B.  P.  Glass.  1970.  Comparative  gastric  histology  of  five  North  and  Cen¬ 
tral  American  bats.  J.  Mamm.,  51:455-472. 

Rumney,  G.  R.  1968.  Climatology  and  the  world's  climates.  MacMillan,  New  York, 
656  pp. 

Saenz,  J.  A.,  and  M.  Nassar.  1972.  Toxic  effect  of  the  fruit  of  Passiflora  adenopoda  D.  C. 
on  humans:  Phytochemical  determination.  Rev.  Biol.  Trop.,  20:137-140. 

Schoener,  T.  W.  1969.  Optimal  size  and  specialization  in  constant  and  fluctuating  environ¬ 
ments:  an  energy-time  approach.  Pp.  103-114,  in  Diversity  and  stability  in  eco¬ 
logical  systems,  Brookhaven  Symposia  in  Biology,  22:264  pp. 

Shannon,  C.  E.,  and  W.  Weaver.  1949.  The  mathematical  theory  of  communication. 
Univ.  Illinois  Press,  Urbana,  125  pp. 

Slater,  J.  A.  1972.  Lygaeid  bugs  (Hemiptera:  Lygaeidae)  as  seed  predators  of  figs.  Bio- 
tropica,  4: 145-15 1. 

Smythe,  N.  1970.  Relationships  between  fruiting  seasons  and  seed  dispersal  methods  in  a 
Neotropical  forest.  Amer.  Nat.,  104:25-35. 

- .  1974.  Insect  sampling.  Pp.  147-157,  in  Smithsonian  Tropical  Research  Institute 

Environmental  Monitoring  Program,  Smithsonian  Institution  Environmental  Sci¬ 
ence  Program,  409  pp. 

Snow,  D.  W.  1965.  A  possible  selective  factor  in  the  evolution  of  fruiting  seasons  in  tropi¬ 
cal  forest.  Oikos,  15:274-281. 


BIOLOGY  OF  THE  PHYLLOSTOMATIDAE 


441 


- .  1971.  Evolutionary  aspects  of  fruit-eating  by  birds.  Ibis,  1 13:194-202. 

Snow,  B.  K.,  and  D.  W.  Snow.  1971.  The  feeding  ecology  of  tanagers  and  honeycreepers 
in  Trinidad.  Auk,  88:291-322. 

- .  1972.  Feeding  niches  of  hummingbirds  in  a  Trinidad  valley.  J.  Animal  Ecol., 

41:471-485. 

Spencer,  A.  W.,  and  H.  W.  Steinhoff.  1968.  An  explanation  of  geographic  variation  in 
litter  size.  J.  Mamm.,  49:281-286. 

Studier,  E.  H.,  and  D.  E.  Wilson.  1970.  Thermoregulation  in  some  Neotropical  bats. 
Comp.  Biochem.  Physiol.,  34:251-262. 

Studier,  E.  H.,  V.  L.  Lysengen,  and  M.  J.  O'Farrell.  1973.  Biology  of  Myotis  thysanodes 
and  M.  lucifugus  (Chiroptera:  Vespertilionidae) — II.  Bioenergetics  of  pregnancy 
and  lactation.  Comp.  Biochem.  Physiol.,  44:467-471. 

Tamsitt,  J.  R.  1967.  Niche  and  species  diversity  in  Neotropical  bats.  Nature, 
213:784-786. 

Tamsitt,  J.  R.,  and  D.  Valdivieso.  1965.  Reproduction  of  the  female  big  fruit-eating  bat, 
Artibeus  lituratus  palmarum,  in  Colombia.  Caribbean  J.  Sci.,  5:157-165. 

Turner,  D.  C.  1975.  The  vampire  bat.  Johns  Hopkins  Univ.  Press,  Baltimore,  145  pp. 

Tuttle,  M.  D.  1970.  Distribution  and  zoogeography  of  Peruvian  bats,  with  comments  on 
natural  history.  Univ.  Kansas  Sci.  Bull.,  49:45-86. 

Valdivieso,  D.  1964.  La  fauna  quiroptera  del  Departamento  deTC'undinamarca,  Colombia. 
Rev.  Biol.  Trop.,  12:19-45. 

Vehrencamp,  S.  L.,  F.  G.  Stiles,  and  J.  W.  Bradbury.  1977.  Observations  on  the  foraging 
behavior  and  avian  prey  of  the  Neotropical  carnivorous  bat,  Vampyrum  spectrum. 
J.  Mamm.,  58:469-478. 

Vogel,  S.  1969.  Chiropterophilie  in  der  neotropischen  Flora.  Neue  Mitteilungen  III. 
Flora,  Abt.  B,  158:289-323. 

Wetmore,  A.,  and  B.  H.  Swales.  1931.  The  birds  of  Haiti  and  the  Dominican  Republic. 
Bull.,  U.S.  Nat.  Mus.,  155:1-483. 

Williams,  C.  B.  1935.  The  times  of  activity  of  certain  nocturnal  insects,  chiefly  Lepi- 
doptera,  as  indicated  by  a  light-trap.  Trans.  Royal  Entomol.  Soc.,  83:523-555. 

- .  1939.  An  analysis  of  four  years  captures  of  insects  in  a  light-trap.  Part  1.  General 

survey;  sex  proportion;  phenology;  and  time  of  flight.  Trans.  Royal  Entomol  Soc., 
89:72-132. 

Wilson,  D.  E.  1971.  Food  habits  of  Micronycteris  hirsuta  (Chiroptera:  Phyllostomatidae). 
Mammalia,  35: 107-1 10. 

Wilson,  D.  E.,  and  J.  S.  Findley.  1971.  Spermatogenesis  in  some  Neotropical  species  of 
Myotis.  J.  Mamm.,  52:420-426. 

Wilson,  D.  E.,  and  D.  H.  Janzen.  1972.  Predation  on  Scheelea  palm  seeds  by  bruchid 
beetles:  seed  density  and  distance  from  the  parent  palm.  Ecology,  53:954-959. 

Wilson,  D.  E.,  and  E.  L.  Tyson.  1970.  Longevity  records  for  Artibeus  jamaicensis  and 
Myotis  nigricans.  J.  Mamm.,  51:203. 

Wolf,  L.  L.  1970.  The  impact  of  seasonal  flowering  on  the  biology  of  some  tropical 
hummingbirds.  Condor,  72: 1-14. 


8240  040 


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.  1 
No.  2 
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No.  10 
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No.  12 

No.  13 

No.  14 

No.  15 
No.  16 


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  Didelphis  (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- 
positae,  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  mites  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  bats  of  the 
New  World  family  Phyllostomatidae.  Part  I.,  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  Astyanax:  environment,  distribution,  and  evolution,  89  pp., 
21  figs .  $5.00 

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 

Francke,  O.  F.  1978.  Systematic  revision  of  diplocentrid  scorpions  (Diplocentridae) 
from  circum-Caribbean  lands,  93  pp.,  146  figs . $7.00 

Carter,  D.  C.,  and  P.  G.  Dolan.  1978.  Catalogue  of  type  specimens  of  Neotropical 
bats  in  selected  European  museums,  136  pp . $8.00 

Baker,  R.  J.,  J.  K.  Jones,  Jr.,  and  D.  C.  Carter,  eds.  1979.  Biology  of  bats  of  the 
New  World  family  Phyllostomatidae.  Part  III.,  441  pp . $20.00 


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