KARYOTYPES OF SELECTED BATS
(ORDER CPHROPTERA)
by
Jerry Lee Osborne
A Thesis Submitted to the Faculty of the
DEPARTMENT OF ZOOLOGY
In P artial Fulfillm ent of the Requirements
For the Degree of
MASTER OF SCIENCE
In the Graduate College
THE UNIVERSITY OF ARIZONA
1965
STATEMENT BT AUTHOR
This th e sis has been submitted in p a r tia l fu lfillm en t of
requirements for an advanced degree a t the University of Arizona
and i s deposited in the U niversity Library to be made availab le
to borrowers under rules, of the Library,
Brief quotations from th is th e sis are allowable without
sp ecia l permission, provided that accurate acknowledgment of
source i s made. Requests for permission for extended quotation
from or reproduction of th is manuscript in whole or in part may
be granted by the head of the major department' or the Dean of
the Graduate College when in h is judgment the proposed use of
the m aterial i s in the in te re sts of scholarship. In a l l other
instances, however, permission must be obtained from the author.
SIGNED
APPROVAL BT THESIS DIRECTOR
This th e sis has been approved on the date shown below g
’ E, L» CQCKRTM
Professor of Zoology
Acknowledgments
Various people have helped in the preparation of th is th e s is .
My particular thanks go to Dr. E, L. Cockrum for h is many suggestions,
comments, and c ritic ism s.
I acknowledge with deep appreciation Dr.
¥ . J. McCauley who read the manuscripts and gave me the b en efit of much
needed constructive criticism and Dr. ¥. R. Ferris for his suggestions
and the use of much of the equipment needed for th is study.
I gratefu lly acknowledge my fellow graduate students who have
helped me in obtaining the animals used in th is study and a sp ecia l
note of thanks i s given to Jerry Nagle and h is w ife, S a lly , for their
constant encouragement and assistance in preparing th is th e sis.
I am
grateful to Jim Patton for h is assistance and h is knowledge of the
technique used in th is study.
I wish to thank Ruth Darden for her
help in the preparation of the illu s tr a tio n s .
To my w ife, Marilyn, I give my very deep appreciation and
thanks for without her help th is th e sis would not have been p ossib le.
iii
Table of Contents
Page
Xjlst of Tables
9qooooooooooossooop^oooooooooooooooooooooo^
l i s t 0,f P lates OOOOCtSOOPOt&OOPOOO^POOOOOOOQpOPOOOOOOOPOO'O^ *vl x
L is t Of PlgUi^eS
A bstract
o o t a a o t i a o o e o o i ^ d d o p d ^ o o p p O i d O O O b O i d O i p o d p o o o t i
o o o o o o o o o o o p e o o p o o o o o o ^ p o o o ^ o o o d o o o e o o p o o o o o o c
Introduction
o o o o o o p o o o o o o o p e o o p d o p p ^ o o a o o o o o o o o o o o o o o o o
M aterials and Methods
P e S U lltS
1%
1
o o o p o o p p o o o o o p a o o o o o p o o o o o p o o o o o o o o
3
o o o o o o o o o p o o o p & o o o o o o o o o o o o o o o p p o a o o c ' o c p o o d o o o
7
Pteronotus da^yi o o o o o d p © c p p p e e 6 o o » o e o o o o o e » p p e o p »
Macrotus /wa te r ho n sli d © o o e d o © © p o o d o © © o o o e o A o e o © o o o d
Choerongotera£ Bie^cxcanns o < ) © o o < > ' © o p o o o o o t > o c t o © o © © o p o ©
LeptonyeterIs nxvalxs o o d p o o © i ! > p o © o p © o © o o o £ j o o c ? o o < ? © o o
Myotis thysanodes o p o o o © © o o t s d o o p o © © o o o o © o p © « ® o o e o ©
MyotXS VOlanS o o o p o o d o o o p o o o o a o p o o d o o o © o © o o o © © o o o o
Myotis fo rtid e n s © o © G < s e © < i i o © o & o © o o © © o © © © o o © o © © o a o o o
Pl^onpc VlVeSl o o O ' £ > o a o o © © o p p | © p © © © ' ( > o e o o © P © « > o o o © © © . o i i j
P ip is tr e lln s hesperns © o o o © © © © © © © © © © © © © © © © © © © © © © © © ©
Eptesicus fnscns © © © © o © © » © © © © © © © © © © o © o © © © © < s i o © © © o o ©
la slu m s cine reus © © © © © © © © © © © © © © © © © © © © © © © © © © © © © o o ©
L aslo n y cteris noctxvagans © © © © © © © © © © © o © © © © © © © © © © © © © ®
Tadarida bras H ie ns is © © © © © < & © © © © © © © © © o o o © © © © © © © © © © * ©
Tadarlaa fetnorosacca © o © o o o o © o © o o o © © © © o 9 o © o o o © o © o © o
D iscu ssion
;
VXlX
©©©©©©©©©©©©©©©©©©©©©©©©©©©©©©ooo©©©©©©©©©©©
Mechanisms of Chromosomal Evolution A ffecting
the Karyotype ©©©©©©©©©©©o©©©©©©©©©©©©©©©©©©©©©©
Fusion and Fragmentation (Robertsonian Ifariatiori) ©
Non-Robertsonian Variation ©©©©©©oo©©©©©©©©©©©©
Polyploidy and Polyson^ ©a©.©©©.©©©©©©©*©©©©**
Interpreting Data Derived From Karyotypes o©©©oo©©©©o©©
Evolutionary Trends and Relationships
Within the Family V espertilionidae o©o©ooo©®oo©©©©©®
C ytological Evidence Suggests the Taxonomic
..^Revision of Pisonyx vivas i 0 » o 00 ©6oo©©©©o©ooo
iv
T
y
^
10
10
11
12
1 2
13
lii
llj,
13
16
16
13
18
18
18
19
19
23
25
"
Table of Contents (continued)
Page
Evolutionary
Within the
Evolutionary
Within the
Trends
Family
Trends
Family
Smmnary and Oonclusxons
and Relationships
Molossidae 6 « o o e Q o b Qo < ? « o o e e 0 0 o o o 0 =
and Relationships
Phyllostomatidae o o o 0 0 o o 0 0 0 o o 0 0 0 = =a
26
27
oooebooboboooboooooooeooooobotioooe
2^
Appendix A
eoeooooeoooooooooooeoooooooooooooooeooooooeoo
31
Appendix B
o o o o o o e o o o o o e o o o o o o Q o e o o o b b o o o b e o o e ^ Q b o o o e o o o
A ppend IX C o o o o o o o o o o o d o o o o o b o & o o o b o o o o o Q o o o o o o o t i o b o e e c y
53
L i t e r a t u r e C ite d
55
©©©©bodooooaoobooooeoooooobooooooeoeboob '
L ist of Tables
Table
I.
II.
Page
Summary of diploid numbers, autosomal types, sex
chromosomes, and fundamental numbers of the species
studied in the order C
b
i r o
p
t e r
Summary of chromosome numbers and fundamental
numbers in the Qhiroptera
a
8
21
III.
Evolutionary trends in the fam ily Yespertilionidae
based on diploid and fundamental numbers. . . . . . . . . . . . . . . . . . . 2h
IV.
Chromosome counts in the genus P i p i s t r e l l u s . . . . . . . . . . . . . . . . 25
V,
Evolutionary trends in the fam ily Phyllostomatidae
based on diploid and fundamental n u m b e r s . . . . . . . . . . . . . . . . . . . 28
vi
L ist of Plates
Plate
1,
Representative male karyotype of Pteronotns davyi o » » o»=«
31
2,
Representative female karyotype of Macrotus
waterhonsxi
o
e
o
e
o
o
o
o
o
o
e
. e
o
d
O
o
o
o
o
o
o
o
o
b
d
o
o
o
t i o
3<> Representative female karyotype of Choeronycteris
mexica nns
© 0 * ^ 0 0 0 0
li
=
o
o
e
e
o
o
e
e
Representative male
o
©
o
o
d
d
o
d
o
o
o
o
d
o
d
o
o
d
o
d
o
o
d
o
e
o
o
o
o
o
o
e
o
e
o
o
o
d
o
o
d
o
o
o
b
0
karyotype of Pizonyx v iv e s i
Representative female karyotype of P ip istre llu s
hesperus
o
o
o
.
32
i »
o
o
o
o
33
o
o
o
o
e
3l|
o
0
o
0
o
o
o
o
o
o
©
o
d
»
o
o
e
o
«
o
o
o
o
b
o
o
o
<
0
0
.
0
0
.
i o
o
o
«
o
o
o
Representative male karyotype of Lasionycteris
noctivagans
13.
.
111
= a o
36
=
e
=
o
=
=
=
a o
e
o
s
=
«
=
=
o
=
«
=
=
= a o
=
«
38
.
o
0 0 0 = 0
11. Representative female karyotype of Laslur us cinereus
o
0 0 0 0
0
10. Representative female karyotype of Eptesieua fuscus
.1 2 .
o
karyotype of Myotis fortldens 0. . 0 . 0 = 37
8, Representative male
o
o
o
karyotype of Myotis volans
Representative male
o
o
o
o
karyotype of Myotis thysanodes o 0 e o e o 35
6. Representative male
9.
o
Representative female karyotype of Leptonycteris
n iv a lis
o
7.
«
o
o
39
Uo
<,«= = J4.I
* « « ' » ' » ' »
I 4 .2
Representative female karyotype of 'Cadarida
b rasilien sxs ^= =o == o ==ooo = ooo = oo = === = = o = oo = = = o =o lt3
Representative female karyotype of Tadarida
femorosacoa oo = = = = = === = = = = =
= ==
o
o
o o
v ii
o o o
o
o o @
= =
o
o o o =
1^1^
L ist of Figures
Page
Figure
I,
Pteronotus davyig Photomicrograph of representative
chromosome spread yo<»oooeooobi >ooeo»o<>ooooodoooooo
2°
Macrotus w aterhousiig Photomicrograph of representative
chr omoso me s pr ead ob. ooooooooboboooobbObG)©ooooooa
U6
Choeronycteris mexicanus: Photomicrograph of representative “chromosome spread <, a e 0 0 0 <, 0 0 <, 0 0 0 „ 0 e 0 0 a 0 0 „
U6
Leptonycteris n iv a lis s Photomicrograph of representative
chromosome spread 000000000000000000000000000000
hi
Myotis thysanodes g Photomicrograph of representative
chromosome spread e ooooooooooooooooooooooooooooo
hi
Myotis voIanss Photomicrograph of representative
chromosome spread ooooe ooooooooooooooooooooooooo
h8
Myptis fortidenss Photomicrograph of representative
chromosome spread 0 00 0 0 0 0 0 0 0 0 0 0 0 oooeoooooooe oooo
U8
Piaonyx v iy e s ig Photomicrograph of representative
chromosome spread 00 000 000000000000000 0000000000
h9
P ip istrellu s. hesperuss Photomicrograph of representative
chromosome spread 0000000 00 *0000 0000 0000000000000
h9
10o Eptesicus fu scu s: Photomicrograph of representative
chromosome spread ooooooooooooooooooooooe ooooo.00
50
11o Laslurus cinereus s Photomicrograph of representative
chromosome spread 00 000000000000 00000 ooooooooooo
51
3»
he
' h,
60
7o
8,
9»
12,
13,
lit.
Laslonycteris noctlvagans? Photomicrograph of representative chromosome spread
oeo*«o«. oooo*o«o«ooo' »o
51
%darida b r a s ilie n s is % Photomicrograph of representative
cEromosbme spread 0000 o 00000 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ,
52
Tadarida femorosacca % Photomicrograph of representative
chromosome spreac oooooooooooooooooooooooooooooo
52
v iii
Abstract
Karyotypes of fourteen species of the order Chiroptera are
presented«, A comparative study of these karyotypes indicates that
centric fusion has played a major role in the evolution of Chiroptera
and that chromosome evolution in at le a s t two species has progressed
away from the main lin e while morphological and behavior parallelism
has been maintained.
While the data in most cases support the present c la s s ific a tio n
and phylogeny of Chiroptera, i t alsoi indicates that some revision may
be necessary.
Evolutionary trends and Interrelationships are discussed for
the three fam ilies;
V espertilionidae, Molossidae, and Fhyllostomatidae.
Introduction
Recent developments in cy to lo g ies1 techniques have caused an .
increased in te r e st in the study of mammalian chromosomes.
Yet, the
use of these techniques as a taxonomic aid has not become widespread
among mammalian taxonomists.
This can be attributed to the fa ct that
cy to lo g ies1 methods are not usually needed for id e n tific a tio n of
mammalian species and ordinarily w i l l simply substantiate the judgements
of an experienced sy stem stist.
However, the importance of these
techniques in interpreting the phylogeny of the mammalian species and
their in terrelation sh ip s should not be ignored.
Gain (1958) gives an
ex ce llen t summary of the use and importance of cytology in taxonomy.
Nuclear cytology has played only a slig h t role in the study of
the mammalian order Ghiroptera,
Prior to 191*8, the information
pertaining to the chromosomes of; bats was very in con sisten t.
Van der
Strich t (1910) reported a haploid number of 9'. or 10 for Vesperugo
noctula,
Athias ( 1912) reported an approximate haploid number of 16
for Rhinolophus hip posid eris,
Jordan (1912) and Hanee (1917) both
report chromosome counts of unknown sp ecies.
The only r e lia b le in for­
mation during th is period prior to 191)8 was reported by Painter (1925), '
In h is study, he correctly determined the diploid number of NyctinomUs:
mexicanus ( ^Tadarida b r a s ilie n s is ),
Makino ( 191)8), using testic u la r m aterial, determined the diploid
number of Pteropus dasymallus jnopinatus to be 38,
The chromosome
complement was reported to co n sist of th ir ty -s ix metacentric and two
1
terminal chromosomes„ This i s the only member of the suborder Megachiroptera that has been examined c y to lo g ie s lly .
The Japanese horse­
shoe b at, Rhinolophus ferrura-equinum, -Has a lso included in h is study
and a haploid number of 29 >ms reported for the sp ec ies.
In 19li9, Bovey studied fourteen species of Old World bats
representing three fa m ilie st
1, Rhinolophidae, three sp ec ies| 2.
Nycteridae, one sp e c ie s| and 3= V espertilionidae, nine sp e c ie s.
His
study represents the most recent in vestigation of chromosomes in bats
that has come to the atten tion of the author,
The present in vestigation has three objectives s 1. to make
available to other workers c y to lo g ies1 information regarding additional
species of Chiropteraj 2. to elucidate a few problems in the taxonomy
and phylogeny of Chiropteraj and 3. to present data that w ill be useful
in future stu dies concerning the evolution of mammalian chromosomes. .
In Chiroptera, the phylogeny i s complex and present interpretac­
tions are often ambiguous.
Cytotaxonomy can play a major role in
cla rify in g the relation sh ip s of many groups within th is Order.
present studyg
In the
four Species in the fam ily Pbyllostomatidae, seven
species in the family V espertilionidae, and two species in the family
Molossidae have been examined by c y to lo g ica l techniques.
The chromo­
some morphology of each i s described and illu stra te d in karyogram form
and selected photomicrographs are included.
Since only a few scattered species within the Order have been
studied by cy to lo g ica l methods, only trends in phylogeny can be shown.
The r esu lts generally v e rify the c la s s ic a l taxonomic c la s s ific a tio n but
in some instances show that minor revision s may be necessary. .
Materials and Methods
With the advent of the colchicine-hypotonic c itr a te method of
Ford and Hamerton (1956) the study of mammalian chromosomes has been
grea tly advanced.
The action of colchicine i s th ree-fold s
1 . i t acts
to prevent spindle formation and thereby causes an accumulation of c e lls
in metaphase$ 2. i t causes the chromosomes to shorten; and 3 . i t causes
the s is t e r chromatids to separate, showing the exact p osition of the
centromere.
The action -of the hypotonic c itr a te solu tion complements
the action of the colchicine by expanding the c e l l .
To date there are several good techniques u tiliz in g the
colchicine-hypotonic c itr a te method.
The one chosen for th is study is
described by Fatten (1965) and involves the following step s:
1,
In ject liv e animals intraperitoneally with colchicine
( 0.05 grams percent), 0,01 ml, per gram body weight.
2,
S acrifice the animal after seven hours and immediately
remove the complete humerus.
The epiphysis i s cut o ff and the bone
marrow removed from the shaft by flushing with 3 ml. of 1.0 percent
sodiumc itr a te .
3,
Pipette solution vigorously to break
up any c e l l clumps.
Incubate the resu ltant c e l l suspension a t 37° C for 30
minutes,
....
F ilte r solu tion through two layers of cheese cloth and
centrifuge a t 500 rpm for fiv e minutes.
5.
Pour o ff supernate without disrupting the button, add 3 ml.
of fresh ly prepared Carnoy8s fix a tiv e and allow c e lls to f ix in ac o l d ,
3
box for 30 minutes„
6.
Centrifuge a t $00 rpm for fiv e minutes and then wash several
times without disrupting the button,
7,
Pour o ff supernate and add about 0,7 ml, of fix a tiv e .
Re­
suspend c e lls by pip ettin g vigorously.
8,
Pipette droplets of c e l l suspension onto chemically clean
s lid e s and ig n ite the suspension immediately, follow ing the blaze-dry
method of Scherz (1962).
9.
10.
Allow slid e s to dry,
Stain in aceto-orcein for one and one h a lf hours.
Dehydrate in three, 30 second changes of absolute methanol
and mount.
After the slid e s were prepared each was scanned for w ell spread
metaphase p la te s.
Those c e lls that appeared in ta ct with the c e ll
membrane undamaged by the preparatory technique and with chromosomes
spread symmetrically were used to obtain the fin a l diploid number of the
sp ecies.
C ells that showed chromosomes widely separated and with other
c e lls in the v ic in ity were disregarded.
Any spreads that showed
chromosomes representing d ifferen t m itotic stages or reacting in d iffe r ­
ent ways to the sta in were also disregarded.
The c e lls selected to be
counted were then placed under 1280 X magnification and counted several
times by v isu a lly dividing the chromosome complement in to several groups.
Photomicrographs were taken of a t le a s t s ix spreads for each species
and then the chromosomal complement was again counted.
A Zeiss microscope equipped with a 3$ mm. camera was used and
photomicrographs were taken on Kodak high contrast copy film .
For karyogram a n a ly sis, in ta ct c e lls were chosen based on the
follow ing c r ite r ia s 1. c e lls showing the le a s t amount of overlapping
of chromosomes| 2. c e lls exhibiting the highest degree of chromosome
morphological acuity; and 3. c e lls representing m itotic metaphase.
In
describing the karyogram, the standardised terminology of Patton (1965)
was used.
In h is description, there are four types of chromosomes
based on arm r a tio s .
The median chromosome possesses a median centro­
mere and has an arm-ratio between 1 :1.0 and 1 :1, 09; the sub-median
chromosome has a submedian centromere and an arm-ratio between 1 :1,10
and 1 :1 . 99; the sub-terminal chromosome has i t s centromere subterminal
and an arm-ratio 1:2 or greater; the terminal chromosome has no v isib le
second arm.
Metacentrics refers to both median and submedian chromo­
somes while acrocentrics refers to both terminal and subterminal
chromosomes.
For karyogram a n a ly sis, the chromosomes were separated into
groups according to their, morphology and comparative s iz e .
In many
instan ces, a group contained a series of chromosome pairs that gradually
decreased in siz e without any obvious breaks in the se r ie s; thus the
group was not further subdivided.
The sex chromosomes were determined on the basis of the heteromorphic pair found in the males.
The non-homologons pair was designated
as the sex chromosomes and I - and T- determination was decided from the
female karyotype which con sists of a homomorphic X-complement.
Live bats used in th is study were collected in Japanese "mist"
nets and hand nets.
The Japanese "mist" nets were employed over water
holes and in the entrances of mine tunnels and caves.
Hand nets were
employed insid e mine tunnels and eaves where bats were found clinging
to the w a lls»
Specimens were id en tified on the b asis of external morphology
and sk u ll c h a r a cter istics« They are a l l now located in the museum a t
the U niversity of Arizona as part of the permanent c o lle c tio n .
Local­
i t i e s and catalog numbers are lis te d in Appendix C,
The taxonomy and c la s s ific a tio n of the bats used in th is study
was based on Hall and Kelson (1959)$ Simpson (19li9) and M iller (1906).
Results
Fourteen species (including four species in the fam ily Phyllostonmtidae, eigh t species in the fam ily V espertilionidae, and two
species in the family Molossidae) have been examined for chromosome
morphology and diploid number.
The numerical r esu lts from th is study
are lis te d in Table 1, and the morphological description of each species
karyotype i s given below.
Representative karyotypes and photomicro­
graphs are presented in Appendix A and B resp ectively.
Species Examined in the Family
Phyllostomatidae
Pteronotus davyig Subfamily Chllonycterinae
Two male individuals were examined and the sp ecies' diploid
number was determined to be !|0.
A representative karyotype (Plate 1,
Appendix A) shows three groups into which the chromosomal complement
was separated based on chromosome morphology and s iz e .
Group A co n sists of
s ix large pairs of approximately equal
sized metacentric chromosomes with eith er medial or submedia1 centro­
meres.
Group B co n sists of fiv e pairs of medium sized to small meta­
centric chromosomes with eith er medial or submedia1 centromeres.
Group C con sists of seven pairs of small to very small terminal
chromosomes.
Since only male specimens were studied, X and Y designation of
sex chromosomes was not p o ssib le .
Structurally, one of the sex
Table I ,
Summary of diploid numbers„ autosomal types3 sex chromosomes,
and fundamental numbers of the sp ecies studied in the order
Ghiroptera.
Family and
Species
Phyllostomatidae
Pteronotus davyi
2n
M
SM
NF*
cT $
38
u
18
(M T)
56
2 0
Macrotus
waterlaousii
1|0
2
18
SM T
60
3
Leptonyeteris
n iv a lis
32
lh
(SM SM)
60
0 3
Choeronycteris
mexicanus
16
it
?' ?
( 26)
Fespertilionidae
Myotis
thysanodes
3it
SM SM
50
it 1
3it
MT
50
1
1
3U (SM T)
50
1
0
3it
(SM T)
50
2 0
6
(SM T)
m
0 1
2
a
(SM T)
m
0
6
6
SM T
it6
2 1
6
SM T
30
2 0
MT
(5it)
2
2
(SM T)
(58)
0
1
ST
T
X-I
lit
2
16
16
=»«=>
csiaes,
b
it
It
14;
2
6
Myotis volans
ifU
2
6
Myotis
“TorEIdens
a
2
6
Pigonyx
vxvesx
a
2
6
o e e=»
P ip istre llu s
"Hesperus
28
lit
8
esaeo
50
«ae«ao.
Lasiurus
"cinereus
28
lit
Lasionycteris
nocMvigans
20
Molossidae
Tadarida
t r a s ilie n s is
—
0
2
1
Eptesicus
6
2
1|8
6
itO
CO
8
it
' it
2
Tadarida
36
* M 88 medial centromere| SM « submedia 1 centromere5 ST «= subterminal
centromere| T = terminal centromere.! HF = fundamental number.
chromosomes i s a small metacentric type with a submedial centromere
belonging to group B and the other a medium sized terminal chromosome
belonging to group Go
Macrotus w aterhousii s Subfamily Phyllostominae
Three specimens, one male and two females were studiedj the
sp e c ie s’ diploid number was determined to be !j.0o Fire morphological
groups can be e a s ily separated in the karyotype (Plate 2, Appendix A)«
Group A includes eigh t pairs of large to medium sized metacen tric chromosomes.
Pair it i s the only chromosome in the karyotype
with a medial centromere.
Group B includes one small pair of submedial chromosomes.
Group C includes one ch aracteristic pair of large subterminal
chromosomes,
Group D includes fiv e medium sized terminal chromosomes.
Group E includes three pairs of small to very small terminal
chromosomes.
The X-chromosome was determined to be a small submedian chromo­
some belonging to group B,
The T-chromosome, not shown in the female
karyotype, i s a terminal chromosome.
Ghoeronycteris mexicanuss Subfamily Glossophaginae
One female specimen was examined from which the species diploid
number of 16 was determined.
The karyotype (Plate 3 , Appendix A) is
e a s ily d iv is ib le into four groups.
Group A co n sists of two large pairs of chromosomes5 pair 1
includes two large subterminal chromosomes and pair 2 includes two
10
large submedial chromosomes.
Group B co n sists of two pairs of medium siz e chromosomes} one
• subterminal pair and one submedial pair.
Group C co n sists of two small pairs of medial chromosomes.
Group D con sists of two pairs of terminal chromosomes} one
large pair and one smaller pair.
Since only one female was examined the sex^-chromosomes could
not be determined,
Leptonycteris n iv a lis s Subfamily Glossophaginae
Three female specimens were examined and a diploid number of 32
was determined for the sp ecies.
The karyotype (Plate ij.s Appendix A)
co n sists of a l l metacentric chromosomes and i s d iv is ib le into two
groups based on s iz e .
Group A includes fiv e pairs of large metacentric chromosomes
with eith er medial or submedial centromeres.
Group B includes the remaining eleven pairs of metacentric"
chromosomes which range from medium to small in s iz e .
Sex chromosomes could not be determined because only females
were examined.
Species Examined in the Family
TTespertilionidae
Myotis thysanodes s
Three individuals were examined, two males and one female,
diploid number of Wi was determined for the sp ecies.
A
The karyogram
(Plate 5, Appendix A) shows three morphological groups that are e a s ily
discernable in the karyotype.
Group A i s composed of three large pairs of metacentric chromo­
somes with either submedia1 or medial centromeres.
Group B contains a sin g le pair of small metacentric chromosomes
with medial centromeres.
Group C includes seventeen pairs of medium to small sised term­
in a l chromosomes.
The submedial X-chromosome i s smaller in siz e than the chromo­
somes of group A and larger than the chromosomes in group B.
The Y-
chromoeoae i s the sm allest metacentric chromosome in the karyotype and
has a medial centromere.
Myotis volansg
Two individuals of th is
female.
species were examined, one male and one
A diploid number of lilt was determined for the sp e c ie s.
As
found in Myotis thysanodes, the autosomal complement could be e a s ily
divided into three morphological groups as shown in the sp e c ie s, karyOgram (Plate 6, Appendix A).
Group A includes three pairs of large metacentric chromosomes
with either medial or submedial centromeres.
Group B contains a sin gle pair o f small metacentric chromosomes
with medial centromeres.
• v-^-
Group C includes seventeen p a irs'o f large to small terminal
chromosomes =
The X-chromosome i s a medium sized metacentric type with a
medial centromere and i s e a s ily
recognized in the karyotype because of
i t s ch a racteristic s iz e , smaller than those of group A and larger than
12.
those of group B.
In contrast, the Y-chromosome i s a small terminal
chromosome of group C.
Myotis fo rtid en s:
The karyotypes of a l l three Myotis studied are morphologically
indistinguishable from each other.
The karyogram of Myotis fortidens
(Plate 7 ) Appendix A) shows chromosomes s lig h tly larger than those of
the other two species of Myotis but th is difference i s due to the use
of an ea rlier stage in m itotic d iv isio n .
The diploid number i s h h and the karyotype i s divided into
three groups.
Group; A con sists of three pairs of metacentric chromosomes with
the la r g e st, pairs 1 and 2, having submedial centromeres and the small­
e s t , pair 3, having medial centromeres,
.
■
Group B i s composed of a sin gle small pair of submedial chromo­
somes ,
Group G includes seventeen pairs of terminal chromosomes.
Since no female specimens were examined the X- and Y-chromosomes
were a r b itr a r ily assigned based on the findings of other sp ecies.
The
X-chromosome i s a medium sized metacentric type with a s lig h t ly sub^» ■'
medial centromere and i s e a s ily distinguished, because of s iz e , from
the remaining chromosomes.
The Y-chromosome i s a small terminal type.
Pizonyx v iv e s i g
Two male specimens were examined and the sp ecies diploid
number was determined to be Wi.
The karyotype (id e n tic a l to the Myotis
karyotype) was divided into three groups, (Plate 8, Appendix A).
Group A includes three large pairs of metacentric chromosomes
with eith er medial or submedia1 centromeres.
A ll three pairs are of
approximately the same s iz e .
Group B includes a sin gle small pair of submedia1 chromosomes.
Group G co n sists of seventeen pairs of terminal chromosomes
varying from the la rg est (pair number 5 ) to the sm allest (pair number
21 ).
The sex chromosomes are recognizable morphologically but could
not be designated as X or T since no females have yet been examined
c y to lo g ica lly .
I f , however, i t i s assumed that the same pattern is
followed as was found for the genus Myotls, from which they can not be
karyologically distinguished, we can a r b itra r ily designate the medium
sized metacentric sex chromosome as X and the smaller morphologically
undefined chromosome as Y.
'
P ip istre llu s hesperus:
One female specimen was examined from th is sp ecies and a
diploid number of 28 was determined.
The species karyotype (Plate 9>
Appendix A) was divided into four morphological groups.
Group A includes two large pairs of metacentric chromosomes $
pair 1 has medial centromeres and pair 2 has submedia1 centromeres.
Group B contains eigh t medium sized pairs of metacentric chromo­
somes with eith er medial or submedia1 centromeres.
There i s a s lig h t
decrease in s iz e from pair 3 to pair 10.
Group G includes a sin g le small pair of medial chromosomes.
Group D includes three pairs of terminal chromosomes that
ex h ib it a s lig h t decrease in s iz e .
Sex chromosomes could not be determined since only females
were examined.
Bptesicus fuscu sg
Two females were examined.
determined to be 50.
The species dip loid nrunber was
The karyotype of the species (Plate 10, Appendix
A) could only be divided into two morphological groups.
Group A con sists of one large pair of medial chromosomes which
represents the only metacentric type found in the karyotype.
Group B i s composed of the remaining twenty-four pairs of
terminal chromosomes = There is a gradual decrease in siz e of the
chromosome pairs without showing any breaks for further subdivision of
the group.
Sex chromosomes could not be distinguished since only females
were investigated.
Laslurus cinereus s
R esults obtained from two specimens, one male and one female,
show a diploid number of 28 for the sp ecies.
The representative karyo­
type i s illu stra te d (Plate 11, Appendix A) showing the three morpho­
lo g ic a l groupings into which the autosomal complement i s divided.
Group A includes six large pairs of metacentric chromosomes
with medial or submedia1 centromeres.
A ll pairs are of the same rela ­
tiv e siz e with the exception of pair number 6 which i s s lig h tly sm aller.
Group B includes four small pairs of metacentric chromosomes
with medial or submedial centromeres.
Group C includes three pairs of terminal chromosomes.
The 2-chromosome, as illu stra te d in the karyogram, i s a large
metacentric -with a submedial centromere and could be included in group
A based on siz e and morphology.
The Y-chromosome, not shown in the
female karyogram, was determined to be a small terminal chromosome.
Lasionycteris noctivagans:
Two in d ivid u als, both males, were studied and the species
diploid number was found to be 20.
The karyotype (Plate 12, Appendix A)
was divided in four autosomal groups.
Group A con sists of two large pairs of submedial chromosomes
which can be distinguished from each other by the more d is ta l location
of the centromeres in pair 1 and the more medial location of the centro­
meres in pair 2,
Group B includes three pairs of medium sized metacentric chromo­
somes which can be ind ividu ally id en tified by the location of the
centromeres.
Pair 3 is subterminal| pair U i s medial and pair 3 is sub­
medial.
Group C co n sists of an extremely small pair of medial chromo­
somes.
Group D contains three small pairs of terminal chromosomes. -- Sex chromosomes of th is sp ecies are determined morphologically
but a t th is time cannot be designated x and X since no females were
examined.
One i s a medium sized submedial chromosome and i s e a sily
id en tified due to i t s unique s iz e , smaller than the chromosomes in group
B and larger than those of group G.
The other sex chromosome appears
as a stained "spot* and i s assumed to be a terminal chromosome repre-senting the sm allest member of group D.
16
Species Examined in the Family
Molossidae
Tadarida b r a silie n s is g
Two specimens, one male and one female, have determined the
sp ecies diploid number to be 1|8°
The karyotype (Plate 13, Appendix A)
can be separated into three groups according to s iz e .
Group A con sists of a sin g le large pair of submedian chromo­
somes .
Group B includes three medium sized pairs of submedia1 chromo­
somes.
Group C includes nineteen pairs of acrocentric autosomes with
subterminal centromeres.
■
The Y-chromosome, not shown in the representative female karyo­
type, i s a medium sized subterminal acrocentric type.
The X-chromo­
some as shown in the karyogram i s a medium sized metacentric chromosome
with a submedia1 centromere.
Tadarida femorosacca g
The karyotype of th is species (P late lli. Appendix A) was deter­
mined from a sin g le female individual.
1|.8„
The species diploid number is
The karyotype i s divided in to four morphological groups which
e a s ily distin guish es i t from the karyotype of the related sp ecies,
Tadarida b r a s ilie n s is .
Group A i s composed of a sin g le large pair of submedial chromo­
somes.
Group B co n sists of four medium sized pairs of chromosomes with
both submedial and subterminal centromeres.
Group C i s composed of a sin g le pair of small subterminal
chromosomeso
Group D con sists of eighteen pairs of terminal chromosomes.
Since only a female specimen was examined, the sex chromosomes
of th is species could not be determined 6
Discussion
Mechanisms of Chromosomal Evolution
A ffecting the Karyotype
The various mechanisms of karyotype evolution were discussed
exten sively by White (19I45') and Swanson (1957)•
In th is study, we were
only concerned with those relevant to the changes in chromosome numbers
and/or a ffectin g the karyotype.
A.
Fusion and Fragmentation (Robertsonian V ariation).
Robert­
son (1916) , in h is studies on in s e c ts, f i r s t suggested the phenomenon
of fusion.
He suggested that the metacentric chromosomes may have
arisen by the apical fusion of two acrocentric chromosomes or according
to the formulas
V ^ l + 1.
This im plies that the reverse process, frag­
mentation or f is s io n , i s a lso possible but further in vestigation s have
shown l i t t l e supporting evidence in mammals.
Matthey (19^5, 19^9 ) has shown that centric fusion accounted
for a large part of the v isib le chromosome changes between d ifferen t
karyotypes of a llie d species of la c e r tilia n liz a rd s.
In h is work he
coined the term nombre fondamental (N .F .) or fundamental number for the
to ta l number of chromosomal arms in a complement,
B.
Non-Robertsonian Variation.
In th is group are a l l other
mechanisms a ffec tin g the structural changes of chromosomes including
the various types of tran slocation s, inversions, and d eletion -d u p licatio n fa ctors.
In th is study, only p ericen tric inversion s, ones which
could transform one metacentric chromosome into an acrocentric
18
■
19
;
chromosome or v ice-versa, would cause a noticeable a ffe c t in the species
karyotypeo
G» Polyploidy and polysomy are not thought to have played a
sig n ific a n t role in mammalian chromosomal evolution and no evidence for
eith er case has been found in the evolution, of the Ghiroptera,
Interpreting Data Derived From Karyotypes
Before interpreting the accumulative data in Table 2, a b r i e f . .
discussion concerning the r e lia b ilit y of fundamental numbers i s necessary.
The nombre fondamental can be determined in.two ways 3 ( 1) by counting
a l l arms in the complement including the sex chromosomes as Bovey has
done in h is study; or ( 2) by counting only the chromosomal arms of the
autosomes, excluding the sex chromosomes, as Maithey la ter defined the
term.
Another d if f ic u lt y in determining th e N .F . i s often encountered
in the study of mammalian chromosomes.’when, subterminal chromosomes are. ,
involved.
In some karyotypes, the complement includes chromosomes with,
extremely small arms which are d if f ic u lt to d etect by the present tech­
niques .
Therefore, the establishment of l.F , becomes a very d elicate
task and the value can be determined only in an approximate and often
arbitrary manner. ■
.■.-
In the present study the fundamental. number has been determined
according to the system proposed by Matthey in which a l l autosomal
chromosome arms are counted.
In cases where only female specimens were
examined, two was subtracted from the t o ta l number in the assumption
that the Y-chromosomes were term inals.
With only one exception, where.
.■
20
a l l chromosomes of the complement were of the metacentric type, a l l
determined Y-chromosomes were terminal chromosomes.
In interpreting the ranges of diploid and fundamental numbers
in Table 2, i t was assumed that a l l Robertsonian variation (fusion)
reduces the to ta l number of chromosomes without changing the to ta l
number of arms.
This i s to say, i f a l l species in a fam ily underwent
only Robertsonian variation we would expect to find a wide range in
diploid numbers but the fundamental number of each sp ecies would remain •
equal and unchanged.
I f any other structural change played a role in-
the chromosomal evolution of the group,-we would expect to find either
a decrease or increase in the fundamental number.
For example, i f a
•-
metacentrie chromosome was formed by a p ericen tric inversion instead of
by the cen tric fusion of two rods, there would be an increase of two .
arms in the karyotype without a lterin g the diploid number.
This can
obviously work in the reverse manner and create one rod shaped chromo- - .
some from a former metacentrie type thereby decreasing the N.F. by two
in the karyotype. ■In eith er case, the N.F, i s changed by two.
Therefore, changes in the N.F. can only be created by non*
Robertsonian variation but, on the other h a n d i t i s p ossible for non*
Robertspnian variation to counteract i t s e l f .‘and not change the N.F. in
the resultant karyotype, i . e . one p ericen tric inversion creating a
metacentrie chromosome and one creating a terminal type.
Assuming that both -Robertsonian and non-Robertsonian variations .
have played a role in the karyotype evolution,, we can express numer-
- .
ic a lly the degree to which each has been involved based on the range of
N.F. within the family and the range of the diploid numbers.
I f the
21
Table 2.
Species
Summary of chromosome numbers and fundamental numbers in
Chiroptera
_______ _______
'
'
1
2n ' At Ac NF
Investigator
Megachirptera
Pteropodidae
Pteropus dasymallus inopinatus
38
?
?
72
Makino 'W
Microchiroptera
Kycteridae
Nycteris sp.
h2
?
r
79
Bovey ®U9
58
?
?
62
Bovey 9U9
Rhinolophus ferrum-equinum
58
?
?
62
Bovey 8U9
Rhinolophus ferrum-equinum
58
?
?
?
Makino *li8
Rhinolophus hipposideros
k
?
?
60
Bovey ^ 9
38
22
Hi
56
Osborne *65
Phyllostominae
Macrotus waterhousil
ko
20
18
60
Osborne 1165
Glossophaginae
Leptonycteris n iv a lis
32
30
6o
Osborne 865
16
12
li
(26)
Osborne 865
10i
?
?
5ii
Bovey sii9
iili.
?
?
52-5L
Bovey 8li9
Myotis emarginatus
iiii
?
?
52
Bovey 8lt9
Myotis thysanodes
a
8
3U 5o
Osborne *65
Myotis voIans
iOi
8
3li
5o
Osborne ’65
Myotis fortidens
kh: 8
3U 5o
Osborne 165
Myotis daubentoni
h2
?
?.
51
Bovey %9
PiBonyx v iv e si
hb
8
3ii
5o
Osborne 865
P ip istr e llu s p ip is tr e llu s
b2
?
? (51)
Ehinolophid ae
Rhinolophus euryale
Phyllo s t omatid ae
Ghilonycterinae
Pteronotus davyi
Choeronycteris mexicanus
Vespertilionidae
Myotis myotis
■ Myotis mystacinus
Bovey 8li9
22
Table 2.
Species
Summary of chromosome numbers and fundamental numbers in
Chiroptera. ( continued)
2n Mt Ac NF
Investigator
UU ?
?
?
Bovey fk9
Plecotus auritus
32
?
?
51*
Bovey ”1*9
Barbastella barbastelius
32
?
?
51*
Bovey ”1*9
Eptesicus fuscus
50
2
-d*
Osborne ”65
Miniopterus sch reib ersii
h6
?
?
52
Bovey ”li9
Laslurus cinerius
28
20
6
1*6
Osborne ”65
L asionycteris noctivagans
20
10
8
30
Osborne ”65
hB
?
?
?
Painter ”25
P ip istr e llu s nathusii
Molossidae
Tadarida b ra silien ses
Note:
22
CO
(1*8)
28
CO
6
P ip istr e llu s hesperus
Osborne '65
TadarIda b ra silien ses
1)8 6
1*0 (51*)
Osborne ”65
Tadarida femorosacca
U8 12
36 (58)
Osborne ”65
Mt = metacentric chromosomes|
HF * fundamental number
i c = acrocentric chromosomes!
23
range of N.F, i s aero then Robertsonian variation has contributed 100
percent, on the other hand i f the range of N.F. i s equal to the range
of diploid numbers then each type of variation has contributed exactly
50 percent.
By interp olating the proportional degree of variations in
N.F. we can obtain the approximate percentage of Robertsonian variation
(fusion and/or fragmentation) and of non-Robertsonian variation.
Evolutionary Trends and Relationships
Within the Family Fespertilionidae
Looking a t the family Fespertilionidae as a whole, one can only
conclude that i t represents a very heterogeneous group.
The diploid
numbers of the seventeen species studied vary from the lowest count of
20 in Lasionycteris noctivagans to the highest count of 50 in Eptesicus
fuscus, giving a diploid range of 30 for the family.
By using the N.F.
as a criterio n for a n a ly sis, a range of 26 i s observed.
In eith er case,
the data indicate a very diverse group, a t le a s t karyotypically.
I f one separates the family in to two p arts, one part including
a l l the species except Lasionycteris noctivagans and the second part
including only L. noctivagans, one arrives a t a very d iffe re n t conclu­
sion .
We can ju s tify th is separation of L. noctivagans.
F ir s t, ana­
tomical and morphological studies by Winge (19I1I ) indicate that the
genus Lasionycteris is a very divergent member of the family F esp ertil­
ionidae and secondly, the cytologies1 evidence; presented here indicates .
that the genus occupies an isolated p osition a t le a s t karyotypically . ..
from the remaining genera studied.
By analysing the cyfcologieal data as previously described, the-,-vresu lts in Table 3 were obtained.
2k
Table I I I .
Evolutionary trends In the fam ily Vespertilionidae
based on diploid and fundamental numbers.
Family
Vespertilionidae
2n
%
N.F.
% R.7.
Non-R* Vo
Group 1
20-50
(range 30)
30-51*
(range 2U)
lilt. 7
55.3
Group 2
28-50
(range 22)
it6-5U
(range 8)
81.8
18.2
Notes
2n; = diploid number| N.F. = fundamental numberi R.7. =
percent of Robertsonian variation; Non-R.V. = percent of
non-Robertsonian variation. Group 1, includes a l l the
studied vesp ertilion id sp ecies; Group 2, includes a l l the
studied vesp ertilion id species except for L. noctivagans.
The cyto lo g ies1 evidence indicates that Robertsonian variation
has played the major role in the evolution of the vesp ertilion id bats
-with the exception of the sin g le genus L asionycterls.
The la tte r
appears to be karyotypically divergent and occupies a p osition isolated
from the remaining genera in the fam ily.
Farther evidence of th is hypothesis can be shown in the genus
P ip is tr e llu s.
Bovey8s study of P ip istre llu s p lp ls tr e llu s shows that
the karyotype of the species includes nine metacentrie chromosomes and
th irty-th ree rod shaped chromosomes for a diploid number of U2.
Since
h is study involved the male sex5 we can subtract one X-metacentric and
one T-acrocentric chromosome from the to t a l and have remaining eigh t
metacentrie and thirty-tw o acrocentric chromosomes for the species auto­
somal karyotype.
P ip istr e llu s hesperus, a New World representative of
th is genus, has an autosomal karyotype consistin g of twenty-two meta­
cen trie and four acrocentric chromosomes (see Table li).
By the mechan­
ism of centric fusion we can convert twenty-eight acrocentric
chromosomes to fourteen metacentric chromosomes in P. p ip is tr e llu s and
th e o re tica lly evolve P. hesperus which has a 2n number of 28, (twentytwo metacentric and four acrocentric chromosomes)«
Table 17.
Chromosome counts in the genus P ip is tr e llu s .
Species
2n
Met.
Acr.
P. p ip is tr e llu s
lt2
8
32
P. hesperus
28
22
it.
Note ? Met. «* metacentric chromosomes
Acr. ® acrocentric chromosomes
C ytological Evidence Suggests the Taxonomic
Revision of Pisonyx v iv e s i
M iller (1906) changed the generic name of Myotis v iv e s i to i t s
present taxonomic statu s Pizonyx v iv e s i.
This change in generic status
was based on the follow ing fa cts s 1 . the siz e of the foot r ela tiv e to
the tib ia exceeded that of any of the large-footed sp ecies of Myotis
and the extreme compression of the claws was unlike any member in the
related genus| 2. a glandular mass i s present near the middle of the
forearm and i s not found in any other c lo se ly related sp e c ie s.
other characters f a l l within the Myotis range.
A ll
The c y to lo g ica l data
presented here ind icates that the change by M iller in the generic
statu s of Myotis v iv e s i was not valid.and that Pizonyx v iv e s i should be
put back with the genus Myotis as f i r s t described by Menegaux (1901).
As shown in the karyograms of the three sp ecies of M yotis, M. thysa#
nodes, M. volans and M. fo rtid en s, and the karyogram of Pizonyx v iv e s i.
these four individuals have id en tica l karyotypes and c y to lo g ie s lly
are inseparable.
Each species has a diploid number of lUt and ident­
ic a l chromosomal complements composed of ttio median, s ix submediah,
and thirty-four terminal chromosomes a The s iz e relation sh ip s between
the four chromosome complements appear approximately equal when the
karyotypes from the same m itotic stages are compared.
Evolutionary Trends and Relationships
Within the Family Molossidae
Only two species of the genus Talarida were studied, both shew
in a diploid number of 1|8.
However, both karyotypes show morph­
o lo g ica l differences which can e a s ily be used for sp ecies id e n t if i­
cation.
Tadarida b r a silie n s is shotjs a unique karyotype consistin g
of eigh t submedia1, th ir ty -e ig h t subterminal and no terminal chromo­
somes.
The subterminal chromosomes a l l have extremely small second
arms which makes the concept of N.F. d if f ic u lt to apply.
the arms were to be
c o u n te d
I f a l l of
s the resu ltant N.F, would equal 92 as com­
pared to the N.F. of T. femorosacca, (5 8 ).
By not counting the sub­
terminal arms, an N.F. of 5h i s obtained and i s more c lo se ly related
to that found in T. femorosaeca which has only four subterminal chrome
somes.
This strik in g difference may simply be due to the technique
used in preparing the chromosome m aterial or might be explained by
a se r ie s of p ericen tric inversions.
In eith er case further study on
the cytology of the molossids i s needed for a better understanding of
the mechanisms involved in their evolution.
2'7
Evolutionary Trends and Relationships
Within the Family Phyllostomatidae
Within the family Phyllostomatidae, Ghoeronycteris mexicanus
exh ib its a unique karyotype with a diploid number of 16.
This repre­
sents the lowest diploid number that has thus far been determined in
the order Ghiroptera* Leptonycteris n iv a lis , the only other species
studied belonging to the same subfamily, Glossopbaginae, has a diploid
number of 32 and the en tire complement.i s composed of metacentric
chromosomes.
The karyotype of Choerohycteris mexjcanus i s composed of
s ix pairs of m etacentric, two pairs of subterminal, and two pairs of
terminal chromosomes.
<
Pteronotus davyi and Macrotus w aterhousii, the
two remaining phyllostomatids studied, each representing separate sub­
fa m ilies, show diploid numbers of 38 and it© resp ectiv ely .
From th is
data, we can only assume that Ghoeronycteris mexicanus represents a
karyotypically divergent form in the fam ily Phyllostomatidae while
maintaining sim ilar morphological c h a ra cteristics.
The other cyto­
lo g ie s ! explanation i s that th is species represents a d iffe re n t
evolutionary lineage and shows extreme morphological convergence in
adaptation and behavior mechanisms.
By assuming that Ghoeronycteris mexicanus i s a divergent form
and represents an isolated p osition in the evolution of the p h yllosto-
•
matida, we can speculate as to the mecbanidm involved in the evolution
of the remaining phyllostomatids by examining the karyotypes of the
other three species studied.
This w ill give us the r e su lt in Table 5
:
28
•which ind icates that Robertsonian variation has played the major role
in the evolution of the phyllostomatids and that Choeronycteris
mexieanus probably represents an isolated position within th eir evolu­
tio n .
On the other hand, i f we do not accept the theory that Cfaoeronycteris mexieanus represents a divergent form then the r e su lts are
reversed and non-Robertsonian variation has played the major evolution­
ary r o le .
Table 7.
Evolutionary trends in the fam ily Phyllostomatidae
based on diploid and fundamental numbers.
Family
Phyllostomatidae
%
2n
N.F.
%
R.V.
Ron~R„¥.
Group 1
16-80
(range 28)
26-60
(range 38)
29.3
70.7
Group 2
32-80
(range 8 )
56-60
(range 8 )
75=0
25=0
Note g Group 1-, includes a l l the studied phyllostomatid sp ecies3
Group 2, excludes Choeronycteris mexieanus.
Summary and Conclusions
Improved c y to lo g ica l techniques have been used in a study of
fourteen species of the order Chiroptera representing the fam ilies s
Phyllostoraatidae 3 V espertilionidae$ and Molossidae,
Diploid numbers,
fundamental numbers, and chromosome morphology have been determined for
each species and corresponding karyograms have been illu s tr a te d .
The r esu lts from th is study haw provided the b a sis for the
follow ing conclusionss
1.
Cytological information has in most cases agreed with and
further substantiated previous work in the taxomony and pbylogeny of
Chiroptera.
2.
Karyotype differences can be seen in a l l sp ecies examined
with the exception of two genera, Myotls and Pissonyx.
3»
The three species of Myotis g M»- thysanodes, M. volans, and
M, fortidens have sim ilar karyotypes and. can not be distinguished from
each other«
It.
Cytological evidence suggests that Pizonyx v lv e s i should
be Myotls v iv s s i.
5.
Centric fusion has played the major role in the evolution
of Chiroptera.
6.
Centric fusion can be demonstrated th e o r e tic a lly as the
mechanism for the evolution of the New World P ip istr e llu s hesperus from
the Old World P ip istr e llu s p ip is tr e llu s .
29
- . .
7.
Lasionycteris noctivagans represents a divergent genus
occupying an Isolated p osition in the phylogeny of the fam ily Vespertilio n id a e .
8.
Ghoeronycteris mexicanus shows extreme karyotypic diver­
gence in the fam ily Phyllostomatidae,
Appendix A
8X»* %n
n
XXXN*V
wy
» W
VV4#
1
o
•a
6
o
0
7
9
10
ii
V
«#
12
a# a#
16
1.
13
#
w
17
+
#
15
¥
18
Representative male karyotype of Pteronotus davyi
(2l4,OOOX)
X X K A Kx HX
5
6
7
K
*
W
K
c KA
B
10
8
X
X
Dou \,\> uu oo
00
16
E Vv
17
Plate 2.
V18 V
- 19 ~
Representative female karyotype of Macrotus waterhousii
( 2k , 000X)
(Hi
in :
Plate 3.
Representative female karyotype of Choeronycteris
tnexicanus (2li,OOOX)
a
m
1
1
2
0
d
n
H
u
6
U
*
8
7
K
8
K
Q
s
X
IK X $1
10
12
11
k
%
13
*
114
.
15
16
Representative female karyotype of Leptonycteris
n iv a lis (2k,000%)
35
t? n
KK
n
i
M
Y
u
MV
5
irU
<I U
vt>
V V
6
7
8
# V
HU
10
9
U U
Vv
u u
11
12
0 V
VV
15
16
13
Ih
VV
wV
V w
17
18
19
V
*
20
% ^
21
Plate 5.
Representative male karyotype of Myotis thysanodes
(24,000%)
1
2
3
*M
X
c u V xr
ty U
V O t/v w u o w
B
14
x
9
10
11
IS V
bh#
13
^
w
18
m*
+
20
Plate 6.
15
^
17
**
V V WV
111
V
12
16
#* w
19
m
21
Representative male karyotype of Myotis volans
(2 k ,0 0 0 X )
M
-
m
a
n
B
h
= yu
Ut f
U
6
9
0 0
13
Vv
17
Plate 7.
8
7
VO
av
0 0
10
n
12
W
111
yu
18
U0
VV
IS
16
Vv
19
*
*
20
Representative male karyotype of I^Iyotis fortidens
(2l| ,000X)
8
A U
8
8 8
2
B /
Q
X
c
0
<
1.
V 0
V
10
9
U
Ui#
V
xz o
Q
0
n
o
U V
V
w
<1 0
11
V v
1-L4
)l
V V
u v
17
18
u
12
u
Vw
-LJ>
19
20
« »
21
Plate 8.
Representative male karyotype of Pizonyx v iv e si
(2U,000X)
39
- ## Xtf
b X8 XX
3
5
6
MX *& XM
7
C
8
9
10
* X
11
D
U u
12
Plate 9.
w» v
13
vy
lit
Representative female karyotype of P ip istr e llu s
hesperus (2^,000X)
A * 8
1
b VU9UVUUU
2
3
b
5
00 0000«o
6
7
8
9
uo u» ww wu
WOWOuu uv
10
11
114
w
15
13
16
17
» v yw uv
18
U
19
U
22
Plate 10.
12
w
20
¥
23
0
0
214
21
*
*
25
Representative female karyotype of Eptesicus fuscus
(2ti,000X)
m i ix ii
n x*
IX
1
2
3
5
6
x x xx
8
VV
11
Plate 11.
k
#
*
12
xx
9
X
X
10
WV
13
Representative female karyotype of Laslu m s cinereus
(2i;,000X)
8
1
3
v
#
#
#
7
12.
¥
¥
8
R epresentative male karyotype of L asion ycteris
noctivagans (2l^,000X)
( (
u
8 1
3
2
9 9
n
14
n
5
6
# if
#
8
Q
7
1o
13
« »
VV
16
17
# V
20
13.
» V
21
v •
11
10
« «
M
7
M
14
13
VV
18
V
22
v
19
V
W
23
Representative female karyotype of Tadarida
b r a silie n sis (2U,000X)
Uh
n
b yy
C
D
sev*j»#v
V «r
o v V u VO VII
t) V v u v v v v
v v v v «#« v v
10
11
12
13
Ik
vVW«#«* «#•
W
P late lli.
15
16
17
19
20
21
#
23
18
22
2k
R epresentative female karyotype of Tadarida femorosacca
(2li ,000X)
Appendix B
Figure 1.
Pteronotus davyi: Photomicrograph of represen­
ta tiv e chromosome spread (9,600X)
Figure 2.
Macrotus waterhousii: Photomicrograph of repre­
sentative chromosome spread (9>600X)
Figure 3•
Choeronycteris mexicanus: Photomicrograph of
re p re se n ta tiv e chromosome spread (9,600X)
Figure k»
L eptonycteris n i v a l i s : Photomicrograph of rep re ­
se n ta tiv e chromosome spread (9,600X)
Figure 5>. Myotis thysanodes: Photomicrograph of re p re ­
se n ta tiv e chromosome spread (9,600X)
Figure 6 .
Myotis volans: Photomicrograph of representative
chromosome spread (9>600X)
&
5
f t
#
Figure 7.
#
Myotis f o r tid e n s : Photomicrograph of represen­
ta tiv e chromosome spread (9>600X)
Figure 8.
Pizonyx v iv e s i: Photomicrograph of re p re se n ta tiv e
chromosome spread (9>600X)
Figure 9.
P ip is tr e llu s hesperus: Photomicrograph of re p re ­
se n ta tiv e chromosome spread (9>600X)
50
Figure 10.
Eptesicus fuscus: Photomicrograph of represen­
ta tiv e chromosome spread (9,600X)
* *#
Figure 11.
Lasiurus cinereus: Photomicrograph of repre­
sentative chromosome spread (9,600X)
■8
)
P
Figure 12.
L asionycteris noctivagans: Photomicrograph of
re p re se n ta tiv e chromosome spread (9>600X)
52
Figure 13.
Tadarida b r a s ilie n s is : Photomicrograph of repre­
sentative chromosome spread (9>600X)
Figure lit.
Tadarida fem orosacca: Photomicrograph of rep re
se n ta tiv e chromosome spread (9>600X)
Appendix G
Pterono-bus davyis Two specimens examined | (2^1), Gueve de la
Tigre, Sonora, Mexicoj UA 13^20, Minos A rm olillo, Sonora, Mexico.
Macrotus w te r h o u s iii
UA
Three specimens examined $ UA 1 3 h l2 ,
UA 1 3 h l6 p Gueve de la Tigre, Sonora, Mexico.
Ghoeronycteris mexicanusg
One specimen examined| UA 11530,
Madera Canyon, Pima Co., Arizona.
Leptonycteria n iv a lis g Three specimens examinedj UA 13U21,
Minos Arm olillo, Sonora, Mexico; (5 7 ), (5 8 ), Colossal Gave, Pima Co.,
Arizona.
Myotis thyaanodess
Three specimens examined; UA 11536, UA 11537
(225), Madera Canyon, Pima Co., Arizona.
Myotis volans:
Two specimens examined; UA 11519, UA 11522,
Santa Catalina M ts., Pima Co., Arizona.
Myotis fo rtid en s: One specimen examined; UA 11538, Colima,
Mexico.
Pizonyx v iv e s i;
Two specimens examined§• UA 13l|25, UA 13h26,
San Carlos %y, Sonora, Mexico.
P ip istr e llu s hesperus t
One specimen examined; UA 13U22, Minos
A rm olillo, Sonora, Mexico.
Eptesicus fuscuss
Two specimens examined; UA 13h2k, Pima Co.,
Arizona; ( l 6h ), Tucson, Pima Co., Arizona.
Laslurus cinereuss
Two specimens examined; UA 11517, UA 11520,
Santa Catalina M ts., Pima Co., Arizona.
Lagionycteris noctivaganss Two specimens examined| UA 11518,
UA 13b233 Santa Catalina Mts, 3 Pima Co., Arizona«
Tadarlda b r a s ilie n s is ;
Two specimens examined j UA 13ltl8, (252),
Cueve de la Tigre, Sonora, Mexico,
Tadarida femorosacca % One specimen examined| UA 13519, Minos
A rm olillo, Sonora, Mexico.
Literature Cited
Athias, Mo.;: 1912« Sur le s d iv isio n s de maturation de I ’oeuf des
mammiferes. Arch. In st, Bact. Com, P est. Lisbonne, 3»
Bovey, R. 19U9® Les chromosomes des Chiropteres e t des In sectivores,
Revue Suisse Zoologie Geneve, £6s371-!i.20o
. 19ii9b. Les chromosomes des Chiropteres.
ogie Geneve, 56:335.
Revue Suisse
Cain, A. Jo 1958. Chromosomes and th eir taxonomic importance.
Linnean Soc. London, 169th sessio n , pp. 125-128.
Proc.
Ford, C. E ., and J. L. Hamarton. 1956/ A co lch icin e, hypotonicc itr a te , squash sequence for mammalian '.chromosomes. Stain Technol­
ogy. 31s2b7-25l.
H all, E. R ., and K. R, Kelson. 1959. The Mammals of North America.
Vol. I . Ronald Press Co., New York. pp. 79-217.
Hanee, R. T.
12
1917=
.
Jordan, H« E. 1912,
Anz. IjO.
The fix a tio n of mammalian chromosomes.
Anat. Rec.
Notes on the spermatogenesis of the bat,
Anat.
Miakino, S. I9U8. A study of the chromosomes in two sp ecies of b a ts.
(Ghiroptera). B iol. B ull, 9^:275-282.
1956, A Review of the Chromosome Numbers in Animals
State College Press, Ames, Iowa. pp. 264-265.
Matthay, R, I9it5. L8evolution de la formula chromoeomiale chez le s
vertebras, Experientia. I , 50-56 and 78- 86.
1951. The chromosomes of the vertebrates.
IV sl59-l80.
Advances in
is.
Menegaux, M« A, 1901, Description d ’une variete e t d 1une espic
nouvelles de Chiropteres rapportees du Mexique par M. Diquet.
Nat, H ist. Mus, B u ll. P aris, pp. 323-327=
M iller, G. S, 1907= The Families and Genera of Bats,
Nat. Mus., no, 57=
*+'■-
B ull, U. S.
Painter, T. S, 1925= A comparative study of the chromosomes of
mammals, Amer, N at., Vol. 59# pp. 385-W8.
56
Patton, Jo Lo 1965thus (Rodentias
Cytotaxonomy of the pocket mice, genus PerognaHeteromyidae). M=S- th e s is , Univ. of Arizo
Robertson, ¥<, R. B. 19l6« Chromosome studies* I . Taxonomic rela tio n ­
ships shown in the chromosomes of Tettigidae and Acrididae, 7shaped chromosomes and their sign ifican ce in Acrididae, Locustidae,
and G ryllidae s chromosomes and variation* Jour, Morph* 271
179-331*
Sobers, R* G* 1962* Blaze-drying, by Ign itin g the f ix a t iv e , for
improved spreads of chromosomes in leu cocytes, Stain Technology*
37:386.
Simpson, G* G* 19^5* %e princip les of c la s s ific a tio n and a c la s s­
ific a tio n of mammals* Bull. Amer. Mus. Nat. H ist. 85?54-61.
Swanson, C* P. 1957*
New Jersey.
Cytology and Cytogenetics.
Prentice-Ha11 I n c .,
7an der S trich t, 0. 1910. La structure de 1'oeuf des mammiferes.
Mem. Acad. Roy. Belgique, Ser. 2, 2.
White, M. J. D. 1954. Animal Cytology and Evolution (2nd e d . ) .
Univ. Press, Cambridge.