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NOAA Technical Memorandum NMFS 

January 1981 

An Annotated Computerized Bibliography 

of the Use of Karyotypic Analysis 

in the Subspecific Taxonomy of Mammals 

Gary L. Worthen 

N OAA-TM-N M FS-SW FC-9 

U.S. DEPARTMENT OF COMMERCE 

National Oceanic and Atmospheric Administration 

National Marine Fisheries Service 

Southwest Fisheries Center 
NOAA Technical Memorandum NMFS 

This TM series is used for documentation and timely communication 01 preliminary results imterim reports, or 

special purpose information and have not received complete formal review editorial control or detailed editing 

January 1981 

An Annotated Computerized Bibliography of the Use of Karyotypic 

Analysis in the Subspecific Taxonomy of Mammals 

Gary L. Worthen 

Research Associate 

Biomedical Laboratory 

Exceptional Child Center 

Utah State University 

Logan, Utah 94322 

This report was prepared by Gary L. Worthen, Utah State University, 

under contract No. NOAA/79-ABC-00205 for the National Marine Fisheries 

Service, Southwest Fisheries Center, La Jolla, California. The statements, 

findings, conclusions and recommendations herein are those of the author 

and do not necessarily reflect the views of the National Marine Fisheries 

Service. Dr. William F. Perrin of the Southwest Fisheries Center served 

as Chief Official Technical Representative for this contract. 

U.S. DEPARTMENT OF COMMERCE 

Philip M Klutznick, Secretary 

National Oceanic and Atmospheric Administration 

Richard A. Frank, Adminisirator 

National Marine Fisheries Service 

Terry L Leitzell, Assistant Administrator for Fisheries 
"- 

I_ 

_. 

w 

NOAA Technical Memorandum NMFS 

This TM rsrm 4 uucl for docufnen(8Don and bMy cmrnunranon of prel8rnmary results. mmnm reporm. or 

fmibl purpose tntormabon. and have not received complete forma revnru. ed1t0n.l conva. 01 d.oIyd edag. 

January 1981 

An Annotated Computerized Bibliography of the Use of Karyotypic 

Analysis in the Subspecific Taxonomy of Mammals 

Gary L. Worthen 

Research Associate 

Biomedical Laboratory 

Exceptional Child Center 

Utah State University 

Logan, Utah 94322 

This report was prepared by Gary L. Worthen, Utah State University, 

under contract No. NOAA/79-ABC-00205 for the National Marine Fisheries 

Service, Southwest Fisheries Center, La Jolla, California. The statements, 

findings, conclusions and recomnendations herein are those of the author 

and do not necessarily reflect the views of the National Marine Fisheries 

Service. Dr. William F. Perrin of the Southwest Fisheries Center served 

as Chief Official Technical Representative for this contract. 

U.S. DEPARTMENT OF COMMERCE 

Philip M. Klutznick. Secretary 

National Oceanic and Atmospheric Administration 

Richard A. Frank, Administrator 

National Marine Fisheries Service 

Terry L Leitzell. Assistant Administrator for Fisheries 
TABLE OF CONTENTS 

List of Fiqures and Tables .................. .iii 

Introduction. .......................... 1 

Literature Search ani' Information Retrieval Techniques .... 2 

Discussion ........................... 6 

Description of Computerized Listing ............... 12 

Computerized Listing of Articles. ................ 22 

Publication Date List. .................... 22 

List of Authors and Co-Authors ................ 24 

Key-Word-In-Context Listing. ................. 28 

Bibliographic Citation Listing ................ 47 

Abstracts of Articles ...................... 65 

Appendix A - Articles Relating to Cytogenetics of 

Subspecific Taxa ................. 119 

Appendix B - Articles Relating to Cytogenetic Analysis 

Used to Substantiate the Existence of a Full 

Species Where Only a Subspecies was Previously 

Thought to Exist ................. 129 

Appendix C - Articles Relating to Cytogenetics of 

Sibling Species. ................. 132 

Appendix D - Articles Relating to Cytogenetics of Taxa 

at Specific (or higher) Levels .......... 135 

Appendix E - Articles Relating to Cytogenetics of 

Domestic Mammals ................. 142 

Appendix F - Articles Relating to Individual and Intrapopulation Variability, and Sibling Diversity . . 145 

Appendix G - Articles Relating to C-Band (Constitutive 

Heterochromatin) Vari abi 1 ity ........... 148 

Appendix H - Articles Relating to Speciation and 

Chromosomal Evolution. .............. 152 

ii 
LIST OF FIGURES AND TABLES 

Figure 1. 6001ean logic used to find articles pertipent to the 

use of karyotypic analysis in subspecific taxonomy of 

mamnials, displaying the root word (or pol Lion thereof) 

used in the search and some of the more !ommon words 

scanned by the search program under eix, -oot word (see 

text for details) ........... ......... 5 

Figure 2. An example of a series of indexing ident'fication codes, 

showing their two principal components (see text for 

details) ......................... 13 

Figure 3. An example of the publication date listing, showing 

the arrangement of publication date and indexing 

identification code (see text for details) ........ 15 

Figure 4. An example of the author listing, showing the 

arrangement of authors and co-authors in relationship to 

the indexing identification code (see text for details) . . 16 

Figure 5. An example of the key word listing, showing the 

arrangement of the alphabetized key words, title field, 

wrap-around title field, end of title indicator, and 

indexing identification code (see text for details) .... 19 

Figure 6. An example of the full bibliographic citation list, 

showing the four major elements of each citation and 

the indexing identification code (see text for details) . . 21 

Table 1. Words excluded from the key-word-in-context listing in 

the annotated computerized bibliography .......... 17 

iii 
I 

"I 

LNTRQDUCTLON 

As one of the task objectives to be completed in fulfilling our 

contract (#NUAA/?9-ABC-00205) on the Stock Assessment of Pacific 

Ocean Delphinids by G md C Chromosome Banding, we agreed to 

compile an annotated computerized bibliography on the use of 

karyotypic analysis in the subspecific taxonomy of mamnals. 

We stated under "k9search Design" on page 7 of our original grant 

proposal, that: 

. . .an annotated bibliography of articles will be 

constructed discussing the use of karyotypic analyses in 

subspecific taxonomy of mammals. 

include a computerized bibliographic search of titles and 

abstracts, references found in citation in known relevant 

articles, and manual searches. Any available article found 

in such a reference will be obtained and a brief abstract 

written for use as an annotation. All bibliographic entries 

dill be. . .keypunched, verified, and run through a computer 

program which will be developed by our staff for a €36700 

computer to display the entries. The end product of this 

process will be a listing of each key word in the title of 

the article (or other key words we may select) in 

alphabetical order in the context of the remainder of the 

title. 

Abstracts and should be very useful to both our research 

staff and to NMFS personnel. In addition, a list of authors 

(including junior authors) of articles wi 11 appear in 

3lphabetical order. A third section (which, to save expense, 

Tay be typed rather than listed on the computer) will appear 

with a full citation of each article followed by a brief 

annot at ion. 'I 

Techniques used wi 11 

This format is similar to what is found in Biological 

This document, then, is a summary of the most pertinent and most 

recent literature related to that subject. It should be noted, 

however, that a document such as this one is a dynamic entity, and will 

be, by its very nature, in a continual state of flux. Continual 

additions will be made as new literature is published, or as other 

articles in the existing literature come to light. Articles will also 
2 

be removed from the bibliography, as they become outdated 

substantially contribute to the scientific advancement of 

and no longer 

the rield. 

This document, then, is an attempt to capture at this moment in time 

the most relevant literature related to the use of '.aryotypic analysis 

in the subspecific taxonomy of mammals. 

As a result of the dates of available listin95 from the search 

service (see the following section), this bibliog: ;,)hy covers primarily 

the period from 1969 to May of 1980, which coinc;c':s nicely with the 

time frame of the bulk of relevant cytogenetic work in the subject area 

(see Discussion). However, as a result of other search techniques 

(described below), some articles are included as far back as 1959. 

article that was found by any of the search techniques is included in 

this bibliography regardless of the language in which it was written. 

Literature Search and Information Retrieval Techniques 

Any 

A computerized literature search was conducted in the Biosis 

Previews, which is a computerized literature search service that 

abstracts from Biological Abstracts, Biological Abstracts/RRM, and 

Bioresearch Index in their entirety. 

will list more articles on a given subject than a hand search in the 

above sources, as all articles or abstracts are not available in those 

In actuality the Biosis Previews 

sources, but are available to the computer service. 

are broken down into two major sections--one running from 1969 to 1973 

Biosis Previews 

inclusive, and the other running from 1974 to present. In May of 1980 

an update on the most recent Biosis Preview was conducted in order to 

assure that the most recent articles would be included in this 

'bibliography. 

46 

The most recent articles are included, because the 

articles are placed on the Biosis Preview computer index before they 

are actually printed in reference form. 
3 

Several key words were used in the search to isolate articles 

relevant to the subject at hand. These key words were then linked 

together with Boolean logic. Three major components were used in the 

logic system: 

(1) A component referring to subspecific taxonomy and related areas. 

In addition to terms relating strictly to subspecific taxonomy and 

subspeci at ion, terms re1 at ing to species and speci at ion were also 

added. This addition was made because of the frequent intermixing 

within articles of both subspecific and specific taxonomy (without 

clear designation in the title which taxon was being discussed). 

The species component was added also for a second reason--that 

being a very low yield (when linked to the other components) of 

articles relating strictly to subspecific taxonomy. 

(2) A component referring to a suite of cytogenetic terms intended to 

isolate all relevant articles on any form of karyotypic analysis. 

(3) A component used to isolate articles referring to mammals. 

In this situation, the use of a concept code was extremely 

useful, inasmuch as it would be impossible to list all the terms 

which referred to mammals in titles of articles. By using 

the concept code, all articles referring to mamnals were In 

scanned. addition, the term "mammal" was also scanned. 

In all three components of the logic system, whenever applicable, 

I 

a code was used (in this case a question mark); such that a root word 

could be listed, and all words containing that word would be retrieved 

in the bibliographic search. For example, the word "mammal" 

+ 

immediately followed by a question mark (MAMMAL?) would yield all 

articles which contained the words "mammal ,'I "mammals," "mammalogy," 

"mammalian," etc. Within each of the three logical components, each of 
4 

the words was linked together with l'or" such that the presence of any 

one of the words in the group registered the article into that logical 

component. In contrast, each of the logical components was linked 

together with "AND" such that an article had to be registered in each 

of the logical components to be referenced for this bibliography. 

Figure 1 gives a graphic presentation of the Boolean logic used in the 

original computer search. 

This computerized literature search formed the nucleus, and the 

majority, of articles which are referenced here. However, as each 

article was obtained, the Literature Cited section in that article was 

scrutinized in an attempt to find other related articles. In addition, 

some articles were added strictly as a result of the author's 

familiarity with cytogenetic literature. Through a combination of 

these procedures, a comprehensive list of articles relating to the use 

of karyotypic analysis on subspecific taxomony of mammals has been 

constructed. 

Articles were obtained either from the Utah State University 

Library, the University of Utah Library, or the Interlibrary Loan 

facility at Utah State University. 

abstracted art cles are on file in the Biomedical Laboratory of the 

Exceptional Ch Id Center at Utah State University. 

Xerox copies of each of the 

The great majority of articles found were in English; however, a 

few were in foreign languages. In most cases, an English abstract was 

provided, or enough of the article could be read to understand its 

contents. In only one instance was it necessary to have an article 

translated by staff external to our facility. 
5 

INTERSUBSPECIFIC? 

In tersubspecifi c 

Intersubspeci fically 

(or) 

Intraspecific 

lntraspeci fi cal ly 

INTRASPECIFIC? 

(or) 

INTRASUBSPECIFIC? 

Intrasubspeci f i c 

Intrasubspeci fical ly 

(or) 

(or). 

(or) 

(or) 

(or) 

SPECIATION 

SPECIES 

SUBSPECIATION 

SUBSPECIES 

SUBSPECIFIC? 

Subspecific 

Subspeci f i cal ly 

/ 

C-BAND? 

C-band 

C-banded 

C- ba nd i n g 

C-bands 

(or) 

CHROHOSOM? 

Chmsomal 

Chromosomally 

Chromosome 

Chromosomes 

(or) 

CYTOGENET? 

Cytogenetic 

Cytogenetical 

Cytogenetically 

Cytogenetics 

(or) 

CYTOLOG? 

Cytological 

Cytological ly 

Cytology 

(or) 

Gband 

Gbanded 

G-banding 

6- bands 

(or) 

Heterochrornati c 

Heterochrornati n 

6-BAND? 

HETEROCHROMAT? 

(or) . KARY OGRAH? 

Karyogram 

Karyograrns 

(or) 

KARYOLOG? 

Karyol ogi c 

Ka ryol og i ca 1 

Karyological ly 

Karyology 

(or) 

Karyotype 

Karyotypes 

Karyotypic 

Karyotypically 

KARYOTYP? 

\ / 

WL? 

Ham1 

Ham 1 i an 

bmmalogi cal 

Hamloqy 

Mamls 

CONCEPT CODE = 62520 

referring to 

mamnals without 

the word "mamnal" 

actually appearin

in the title 

Figure 1. 

of karyotypic analysis in subspecific taxonomy of mamnals, displaying 

the root word (or portion thereof) used in the search and some of the 

more comnon words scanned by the search program under each root word 

(see text for detai 1 5). 

Boolean logic used to find articles pertinent to the use 
6 

DISCUSSION 

The articles abstracted in this computerized bibliography cover a 

wide range of subject matter. 

intended scope of this bibliography, center around the use of 

kar.yotypic analysis in subspecific taxonomy of mmals. Some articles, 

however, are only peripherally related to that subject in its narrowest 

interpretation. They do, however, have relevance to that subject in 

a broader sense. 

Most of the articles, by virtue of the 

All types of mammals for which literature could be found are 

covered in this bibliography, and no attempt was made to select for or 

against any group of mammals. Many of the mechanisms presented in this 

bibliography are not commonly found in cetaceans, but are comnon to 

other mammalian qroum (such as rodents). 

irrelevance) of these articles to cetaceans will not be discussed 

here--such discussion will be saved for the final report. 

The relevance (or 

Because of the time profile involved, the articles covered in this 

bib1 ioqraph-y, cover a wide range of cytogenetic techniques and schools 

of thought. 

kar.yotyping was by the use of gross karyotypes. 

that only nmber, shape, and general configuration of the chromosome 

complement could be ascertained. Karyotypic work in the ~O'S, however, 

is much more refined because of a major scientific breakthrough in the 

early 70's which allows for the banding of chromosomes. 

manner, homologous chromosomes can be completely matched. 

of this breakthrough, the old gross karyotypes had to be entirely 

re-examined in liqht of the new banding data. 

For example, during the 1960's the common method of 

Such methodology meant 

In this 

As a result 

In many cases, 
7 

the old interpretations were found to be incorrect. These erroneous 

conclusions resulted because it was frequently impossible to tell from 

the gross karyotype, which chromosomes had undergone centric fusion (a 

process by which two chromosomes are united to form one larger 

chromosome). As chromosome nunber, size, shape, and arm ratio were the 

only indicators of what had evolutionarily transpired, many incorrect 

assumptions were made. For example, if three pair of acrocentric 

chromosomes (A, 6, and C) existed in some ancestral type, where 

chromosomes B and C were identical in size and shape, interpretational 

problems could result. If in one evolved species, chromosome A fused 

to 6, and in another closely related species A fused to C, the 

resulting chromosomes would, with gross karyotypinq, appear identical. 

If these chromosomes were the only delineator between the two species, 

the two species may be incorrectly synonynized as one, when in reality 

the two karyotypes were quite distinct. 

would elucidate that fact. 

The bandinq protocols now used 

Although the abstracted articles comprise more or less of a 

continuum, I have somewhat arbitrarily categorized each article into 

one or more subject qroupinqs. Each of these groupinqs constitutes an 

appendix to this report, and all articles dealing with a particular 

grouping can be found listed in the appropriate appendix (A throuqh 

HI. 

The bulk of the articles covered in the biblioqraphy, in the broad 

sense, refer to karyotypic or cytogenetic analysis of subspecific taxa. 

These papers are listed in Appendix A, and include articles on a wide 

range of animal qroups ranqing from rodents, lemurs and deer to piqs 

but weighted heavily with articles coverinq various species of rodents. 
3 

example, larqe suite of articles exists on chromosomal 

polporphisrn sf various geoqraphic forms of the black rat (Rattus 

rattus). 

Various approaches have been used to analyze the diverse 

subspecific taxa being studied. 

number (exemplified by the series of papers by Yosida, -- et al.on black 

rats), and many include a comment on the fundamental number of 

chromosome arms (for example, see Rausch and Rausch, 1972). Other 

articles (for example, Greenbaum, Baker and Ramsey, 1978) use a 

combination of G and C bandinq pattern analysis to elucidate 

Many of the papers are on chromosome 

e, Sharma 

tutive 

chromosomal evolution; while still other papers (for examp 

and Garg, 1975) heavily utilize C-bandinq to analyze const 

heterochromatin. 

In many instances, c-floqenetic analysis has been used to 

substantiate the existence of a full species where only a subsoecies 

was previously thought to exist. Articles covering this subject are 

listed in ADpendix 6, as well as in appropriate positions in other 

appendices. In this category, studies on rodents aqain predominate the 

literature; however, articles on artiodactyls and insectivors are also . 

found. The conce'pt of sibling species or a super species containing 

many closely related forms, has also received wide recognition. These 

articles are listed in Appendix C. 

Articles abound in the literature on karyotypes and cytoqenetics 

of taxa at specific or higher levels. 

articles have been omitted here for reasons of brevity, space 

limitations, and because they are out of the scope of the present work. 

A few of the articles, however, have been included when they: (1) show 

The great majority of these 
9 

relevance to one of the other major cateqories in the biblioqraphy, (2) 

when they show a parallel in technique or interpretation to subspecific 

taxonomy, (3) when they show clear-cut relevance to one or more of the 

other articles contained in the bibliography, (4) when they show 

particular interest to the cytoqenet ics of cetaceans, or (5) when they 

serve as an example of a particular type of paper which is being 

discussed. 

karyotype for a particular species of mammal. 

example, Haiduk, Bickham, and Schmidley 1979) describe karyotypes for 

several species within a particular qenus. Other articles (such as 

Schroder and Van Der Loo 1979) compare the karyotypes of several genera 

within the same family. 

relating to the cytoqenetics of taxa at specific or higher levels can 

be found in Appendix D. 

Many articles (such as Arnason 1970) simply present a 

Other articles (for 

A synopsis of the few abstracted articles 

A section (Appendix E) has been included on the cytoqenetics of 

domestic mammals because of certain parallels between selected breedinq 

in domestic animals and evolution of subspecies per se. Another 

section (Appendix F) has been included on individual variation, interpopulational variabiitv, and sib1 ing diversity. These articles cover a 

wide ranqe of taxa includinq rodents, laqomorphs, pigs, chiropterans, 

and primates (includinq man). Articles included in this group show the 

potential scope of variability within some animal groups below the 

subspecies level (in some cases within a sinqle individual). 

-- 

Because of its relevance to cytogenetic interpretations of 

mammalian taxonomy, a separate section has been included on 

constitutive heterochromatin (C-band) variability in mammals (ADpendix 

G). This section perhaps has the greatest relevance to cetacean 

chromosomal classification inasmuch as most of the variation found 
within cetacean chromosomes are found in variation in regions of 

constitutive heterochromatin. This phenomenon will be discussed in 

greater detail in the final report. A general pattern is frequently 

apparent within mammalian groups for C-banding to show more variability 

between populations than corresponding G-banding. 

The articles in the remaining category (listed in Appendix H) 

cover cytological aspects of speciation and chromosomal evolution. 

Bush, .Case, Wilson and Patton (1977) state that speciation rate is 

strongly correlated with the rate of chromosomal evolution. Bush 

(1975) states that the highest rates of chromosomal evolution occur in 

parapatric situations and that the lowest chromosome evolution rate 

occurs in allopatric siutations where speciation occurs by subdivision. 

In the latter case there is little or no directed chromosomal 

evolution. In allopatric situations where the founder effect is the 

rule, and also in sympatric situations, the rate of chromosomal 

evolution is intermediate. Schroeder, Antoni, and Vanderloo (1978) 

also indicate that speciation is encouraqed by geographic proximity and 

that geographically isolated species and subspecies have no need to 

develop divergent karyotypes. Nevo and Cleve (197'8) also indicate that 

as a general rule, habitat specialists display significantly lower 

genetic variability than do habitat generalists. Arnason (1972 and 

1974) indicates that cetaceans are characterized by a remarkable 

stability of their karyotypic patterns. Most of the existing 

variability is found within the C-band areas of constitutive 

heterochromotin. Arnason (1972) further points out that the orders 

Insect ivora and Rodentia conversely, both display great karyotypic 

variability and would thus represent a different mode of speciation 

than that found in cetaceans. 
11 

Chromosomal evolution historically has been considered a 

unidirectional evolutionary process in the direction of centric fussion 

of small acrocentric chromosomes to form large metacentric chromosomes. 

Lawlor (1974) suggests that cytological evidence provided by studies of 

manmalian chromosomes suggests that “fission” - a process by which 

biarmed chromosomes divide to create small acrocentric chromosomes (and 

thus create a higher diploid number) - has also played an important 

role in chromosomal evolution. 

evolutionary change in the number of chromosomal arms. 

Evidence suggests no obvious trend of 

Other subject categories could be split out from the articles in 

the bibliography. I felt, however, that those that have been covered 

are the most important. If information on other subtopics covered in 

the bibliography is desired, use of the key-word-in-context listing 

will help in the location of that material. 
12 

DESCRIPTION OF COMPUTERIZED LISTING 

The program which generated the computerized listing for this 

bibliography was written in Cobol for a Burroughs 6700 computer. A 

slightly revised version of the program was used to make these actual 

runs on the Utah State University Computer Center's Burroughs 6800, now 

in operation. 

The computerized portion of the bibliography contains four 

separate listings which serve to expedite the location of references 

according to different criteria. Included are elements in the 

bibliography listed according to: (1) date of publication, (2) name of 

author or authors (principal and junior), (3) title of articles 

referenced by key words, and (4) complete bibliographic citation. 

Throughout each of the listings generated by the computerized 

bibliography program, an indexing identification code can be found to 

facilitate cross-referencing between the listings. This identification 

code consists of an author identification field and a numerical index 

field. The author identification field consists of an eight-column 

field bearing as much of the principal (or sole) author's name as space 

permits. If the author's last name exceeds eight characters, the first 

eight letters of the name will appear. In the case of shorter names, 

one or more initials will follow the complete last name of the author, 

as space permits. 

Immediately following (and adjacent to) the author field is a 

4-digit numerical index field. The index number corresponds to the 

numerical position in the full bibliography of that reference. Using 

the example in Figure 2, the first entry (by Craig-Holmes) will be the 

24th reference to appear in the alphabetical citation list. 
13 

indexing identification code . 1 

numerical index field 

CR41G-HUO024 

11- author 

Figure 2. 

identification codes, showing, their two principal 

components (see text for details). 

An example of a series of indexing 
14 

The first list that appears on the printout displays all 

references in the bibliography chronologically according to the date of 

publication. Within each date, the entries are listed alphabetically 

according to the author index field. 

its complete corresponding indexing identification code (see Figure 

Each date entry is followed by 

3). 

The next list which appears on the printout is an alphabetical 

compilation of all authors and co-authors in the left-hand 20-character 

field, fol lowed by the index identification code previously mentioned. 

If an author listed in the first field is the sole or principal author 

of a reference, the name will appear in both the author and 

identification fields. 

be an example. 

In Figure 4 the first line, for Selander, would 

This list is meant to be used primarily to locate a reference for 

which the principal author is not known, or if the work of a particular 

author is of special interest. For example, if the work of T. Sharma 

is being researched, one would find his name in the left column, 

identify him as co-author with R. Ramen, whose name appears in the 

author identification field, and then find the full citation by using 

the identification code (see line 2 in Figure 4). All authors and 

co-authors are listed in this manner. 

In the next list, key words in the titles of articles are arranged 

alphabetically for easy reference to specific subject areas. All words 

that appear within a title, except words statistically found to be of 

little use in suggesting the subject matter of a title (articles, 

conjunctions, and such words as "several," "relative," and "typical'l), 

are included in this list (see Table 1 for a full list of excluded 
15 

I' indexing identification code 

pub1 ication date 

1978 ELDER# F0030 

1978 GR€€N0AUOOSS 

1978 GREEN8AUO036 

1978 KULIEVt 0048 

I978 SCHRODEROO72 

1978 VbLbEZt 008Z 

1978 kHITEt MOO80 

1979 BRUERE, 0013 

1979 DUTRILLA0029 

1979 HAIOUKi 0037 

1979 ROOllVAt 0070 

1979 SASAKIp 0071 

1979 SCHROOEHnO73 

1979 YONENGA-OO@b 

1978 NEVOt EeOObl 

Figure 3. An example of the publication date listing, 

showing the arrangement of publication date,and indexing 

identification code (see text for details). 
16 

indexing i denti f i cati on 

r 

Figure 4. An example of the author listing, showing the 

arrangement of authors and co-authors in relationship to 

the indexing identification code (see text for details). 
-11- 

L 

L 

LTable 1. Words excluded from the key-word-in-context listing in the 

annotated computerized bibliography 

A 

ABILITY 

ACTIOM 

ACT I VI TY 

ADDITIONAL 

ADMINlSltRED 

AFFECT 

AFFECTED 

AFFECTING 

AFTER 

AGAINST 

ALL 

ALONG 

FLTERNATING 

IMOYE 

AN 

ANALYSES 

ANALYSIS 

ANALYI ICAL 

AYD 

ArJIMAL 

ANIMALS 

ANOTHER 

APPARATUS 

APPRDPRIATE 

AREA 

ARC AS 

AR9 A ti CEPE NT 

ARTIFICIAL 

AS 

ASPFCTS 

AT 

ATTACHED 

AVER4GE 

BASED 

BASIS 

bkFORE 

BETHEEN 

BUT 

BY 

CALCULATION 

CAPACITY 

CEQT 

CERTAIN 

CHARACTERIS1 1CS 

L HE HI C A L 

CL3SLD 

CULLECT ION 

COrMLrrlS 

COYMDN 

CONYUNIC AT IOM 

CO'IPUTATIONS 

CONCERNING 

CONDI TION 

COI4NLCT ION 

C 0:JS I DER A 1 I ON 

CO:JSIOERATIONS 

CONSTRUCTION 

C OM T k I NED 

troTn 

COWLETE 

COI4YIhENT 

C 0 N Y 1 NU 0 US 

DATA 

DE 

CEGPEE 

DFCREES 

DEh 

DER 

UETERMINATION 

OETtRYINATIONS 

UC TERHINING 

DIE 

DI FFERENT 

DO 

DOES 

CUE 

DURING 

EFr ECT 

t1s 

ELEMENTS 

EnPHASIS 

LQUILIBRATION 

ESTIMATION 

EVALUATION 

EVIDENCE 

E X PE 4 IME tiTAL 

CXPERIMFNTS 

E X T CIlE 

EXTREYELY 

F ACTOR 

FACTORS 

FEW 

FIRST 

fOR 

+ OUND 

F RUM 

f UWCT ION 

F UPTHE4 

GENUS 

UXGH 

1 

11 

111 

1 MPL IC AT 10'4s 

1~POHTArtCE 

IN 

INFLUEhCE 

I NS T Pur€ til 

I kS TQUYEr+T AL 

1 NTE RNA T I ON AL 

INTERPRETATION 

INTO 

INVFST IGAT ION 

INVESTIGATIOWS 

1s 

ISLArdD 

11s 

IV 

L 

LA 

LbSOHATORY 

LAM 

LE 

LEVEL 

LITTLE 

LOW 

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words). 

of the printout, following a blank space in column 20. As much of the 

title to which the key word belongs as will fit on one line is printed. 

Key words within a title appear beginning in the 21st column 

An asterisk indicates the end of an article title. 

Following any asterisk is the beginning of the title which, when 

the title is short enough, wraps with the first 19 columns of the 

title. Where a title is longer than one line of printout can 

accommodate, a portion of the title is omitted. Enough of the title is 

present, however, to facilitate assessment of the importance of the 

article by seeing the key word in context. 

is another blank space and the indexing identification code which 

corresponds to the reference's appearance in the full bibliography. 

Following the title field 

A title will appear within this listing as many times as the 

number of key words within it (which totals the number of words in the 

title minus the trivial words that have been excluded as indicated 

above). The title, "CHROMOSOME POLYMORPHISM IN INBRED SUBSPECIES OF 

PEROMYSCUS MANICULATUS," for instance, will appear first with 

"CHROMOSOME" being the isolated key word in column 21; again with 

"INBRED" isolated; later with "MANICULATUS" as the key word, and so on. 

Figure 5 illustrates the position of the elements from an excerpt of 

this list. 

The final listing on the printout is an alphabetical (by author) 

bibliography of complete citations. Each citation consists of five 

elements, each including one or mre lines. The first element is the 

cross-reference indexing identification code which appears in the upper 

right-hand corner of each citation. The identification codes 

correspond to those found on the preceding lists. Next, the authors 
19 

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are listed. If there are more than three authors, the author list will 

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located on a separate line near the right-hand side of the list. 

The title of the article follows the date on the next line and 

appears directly below the author list. 

characters, it will appear on two or more lines. The last element of a 

If the title exceeds 60 

citation in this bibliography is the source. 

from which the reference came is indented and appears directly below 

the title of the article. Volume, number (where applicable), and pages 

are found on the same line to the right of the journal name. In some 

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a 
ABSTRACTS OF ARTICLES 

The abstracts of the articles in this section follow standard 

bibliographic order, which is the same order that is followed in the 

Bibliographic Citation section of the computerized listing of articles. 

For each abstract a full bibliographic citation is given. 

Wherever possible the abstracts themselves were drawn from the 

literature: the rationale being that an author of an article would be 

the best person to abstract that article because of his familiarity 

with subject content. Where an author's abstract was not available an 

author's summary was frequently used. When an abstract or a sumnary 

were not available from the author, either the abstract contained in 

Biological Abstracts or an abstract written by myself was used. The 

abstracting source (which is bracketed) immediately follows each 

abstract and is denoted as: 

"authors I abstract ,I' "authors ' summary," "abstract from Biological 

Abstracts," or "abstract by GLW" (Gary L. Worthen). 

"author's abstract ,'I "author Is summary, It 

In the right margin of the first line of each abstract listing is 

a three-digit number which has been added to facilitate crossreferencing within the text. This number corresponds to the last three 

digits of the indexing identification code used throughout this work. 

4. 
Arakaki, 0. T., I. Veomett, and R. S. Sparkes 1970. Chromosome 00 1 

polymorphism in deer mouse sibl ings (Peromyscus maniculatus) . Experientia 26:425-426. 

Siblinq offspring of P. maniculatus subspecies from the same 

parents have been found to'demonstrate chromosome polymorphism. 

Comparison between chromosomes of the sibl ing animals was made based 

on the nunbers of large acrocentric and submetacentric chromosomes; 

and mal 1, metacentric and submet acentric chromosomes (3 groups) . A 

constant diploid nunber of 48 was confirmed, although the nunber 

acrocentric chromosomes varied between 12 and 16 among the animals 

studied. [Abstract by GLW) 

Arnason, U. 1970. The karyotype of the grey seal (Halichoerus 002 

grypus) . Hereditas 64937-242. 

The somatic chromosomes of the grey seal, Halichoerus r us 

Fabr., and their autoradiographic pattern were studied in cu prye tures of 

lung tissue. 

and 14 female cells. 

2n=32. 

interrelationships among the species so far studied of the family 

Phocidae. [Author's abstract] 

An idiogram was made based on measurements of 6 male 

The chromosome nunber of the grey seal is 

Some comnents were made on the karyological 

Arnason, U. 1972. The role of chromosomal rearranqement in mamnalian 003 

speciation with special reference to Cetacea and Pinnipedia. Hereditas 

70(1):113-118. 

The karyotypes of 17 cetaceans and 18 pinnipeds are surveyed. 

Within the Cetacea only two different chromosome nunbers have been 

found, viz. 2n = 42 and 2n = 44. Within the Pinnipedia three different 

chromosome nunbers have been described, viz. 2n = 32, 2n = 34 and 

2n = 36. The material indicates that the orders Cetacea and Pinnipedia 

are characterized by a remarkable stability of their karyotypic 

patterns. Factors cooperating towards karyotypic stabi 1 ity may be 

sought in the reproductive biology of these animals - late sexual 

maturity and small progeny - as well as in their ecoloqy - qood 

mobility in an environment without distinct niches. 

the main mode of speciation should be a allopatric. 

In such animals, 

The orders Insectivora and Rodentia on the other hand which both 

display great karyotype variability would accordingly represent a 

different mode of speciation. 

have early sexual maturity and larqe proqeny; their mobility is often 

limited and the distribution discontinuous. 

Insectivora and Rodentia stasipatric speciation should be frequent. 

[Authors' abstract] 

c 

The species belonging to these orders 

Consequently, in II 
Arnason, U. 1974. Comparative chromosome studies in Cetacea. 

Hereditas 77:l-36. 

The karyology was studied in 9 species of cetaceans, viz. in 5 

Comparisons were made between the 

odontocetes and 4 mysticetes. Chromosome measurements and idiograms 

of 8 species are presented. 

karyotypes of the different species on the basis of conventional 

staining methods as well as by autoradiography and by Q-, G-, and 

C-banding techniques. All cetaceans so far studied have 2n=44, 

except the sperm and pygmy sperm whales, which have 2n=42 and 

karyotypes entirely different from those of the 2n=44 species. 

of the latter, except the killer whale, have closely concordant 

karyotypic morphology. Specific attention was payed to the 

C-heterochromatin, both because of its large amount and its striking 

pattern. 

total chromosome length in the odontocetes to 25-30% in the 

mysticetes. In both of them the distribution of C-heterochromatin in 

the karyotypes was mainly interstitial and terminal and to a lesser 

extent centromeric. 

observed between the homologues of the same pair. The pronounced 

karyotypic agreement between the odontocetes and mysticetes, both in 

general chromosome morphology and C-band pattern, is incompatible 

with the theory of a diphyletic origin of the Odontoceti and 

Myst icet i . 

All 

The amount of C-heterochromatin varied from 10-15% of the 

Size heteromorphism in C-bands was frequently 

[Author's abstract] 

004 

Arrighi, F. E., A. D. Stock, and S. Pathak 1974. Chromosomes of 005 

Peromyscus (Rodentia, Cricetidae). V. Evidence of percentric 

inversions. Chromosomes Today 5:323-329 

Cytoloqical studies were conducted on three species of Peromyscus 

(deer mouse). G-bandinq revealed structural stability in euchromatin, 

whereas constituti&! heterochromatin was found to be very flexible in 

amount and position among species and within a species. [Abstract by 

GLW) 
(L 

P 

111 

Badr, F. M. and R. S. Badr 1970. The somatic chromosomes of a wild 0

population of rats: nunerical polynorphism. Chromosoma 30:465-475. 

The karyotype and quantitative characteristics of a wild 

population of rats, Rattus rattus, were studied. Individuals of the 

population were clas3fEfmree distinct groups, each with a 

characteristic chromosome number of 38, 42 and354 respectively. 

frequency distribution of the three groups of rats in the sample 

studied was as follow: group I with 38 chromosomes formed 14X, group I1 

with 42 chromosomes formed 54% and group I11 rats have had chromosome 

nunbers varying between 50-60 formed 32%. The rats with 38 chromosomes 

had two pairs of marker chromosomes (2 long metacentric pairs). 

of group I11 were characterised by having a marked decrease or complete 

absence of short metacentric chromosomes with a simultaneous increase 

in the frequency of short telocentric chromosomes. Group 11 rats had 

more or less the chromosomal characteristics established for laboratory 

rats studied by previous workers. The total chromosomal length of 

somatic cells in either group 1 and I1 were found to be similar. The 

notable chromosomal polynorphism in number was explained in terms of 

centromeric fusion or dissociation. [Authors' abstract] 

The 

Those 

Baker, R. J., W. J. Bleier, and W. R. Atchley 1975. A contact zone 

between karyotypically characterized taxa of Uroderma bilobatum 

(Mammal ia: Ch iroptera) . System. 2001. 24( 2)- 

A total of 191 specimens of Peters' tent-making bat, Uroderma 

bilobatum, were collected from the zone where two chromosomal races, 

representing two subspecies, meet. 

four had 2n=39, one had 2n=40, one had 2n=41, one had 2n=42, 14 had 

2n=43, and 82 had 2n=44. This chromosomal variation is best 

explained as resulting from hybridization between two cytotypes with 

the 2n=38 and the 2n=44 being the parental (pure) types, the 2n=41 

individual being of the F1 type and the 2n=43, 42, 40, and 39 

individuals representing backcross or F2 products. The two parental ' 

cytotypes were not found to be sympatric at any individual collecting 

station. The zone of hybridization is approximately 200 kilometers 

long on the Pacific versant of Honduras, eastern El Salvador and 

northwestern Nicaragua. 

were reproductively active a1 though the frequency of reproductive 

activity was not so great as in individuals with parental cytotypes. 

Measurements of the cranial and wing morphology of chromosomally 

intermediate individuals indicated that in general there was a 

correspondence between chromosome number and phenet is simi 1 ar ity to 

one or the other of the two parental stocks. Chromoshal and cranial 

and wing morphological data suggest that there is considerable gene 

flow between the two cytotypes and that chromosomal divergence has 

occurred in the absence of speciation. differentiation between the two subspecies serves as a caveat to 

those cases where specific recognition is based on chromosomal 

distinctness of a1 lopatric samples. 

Uroderma. [Authors' abstract] 

Eighty-eight specimens had 2n=38, 

Some chqmosomal ly intermediate individuals .. 

The magnitude of chromosomal 

(Karyotypes; Chiroptera; 

0
Baker, R. J. and J. L. Patton, 1967. 

variation of North American vespertilioned bats. 

Karyotypes and karyotypic 

3. Mmlogy 

48(2) ~270-286. 

Karyotypes of 32 species of North American vesperti 1 ioned bats 

are described. Individual, population, subspecific, specific, and 

generic karyotypic variation are discussed. 

taxonomic tool and phylogenetic indicator in bats is discussed. 

[Authors' abstract] 

The use of karyotypes as 

0

Baverstock, P. R., C. H. S. Watts, J. T. Hogarth, A. C. Robinson, 0

and J; F. Robinson 1977. Chromosome evolution in Australian rodents 

A total of 219 wild caught specimens representing 12 of the 

currently recognised 13 species and subspecies of Australian Rattus 

have been karyotyped. No two species possessed karyotypes in comnon, 

most species and several subspecies differing markedly in chromosome 

nunber. While the diploid nunber varied from 2n=32 to 2n=50, the 

fundamental nunber (FN) varied only from 60 to 62, suqqesting that 

Robertsonian rearrangements have played a major role in karyotypic 

evolution in the group. - Karyotypically the Australian species of 

Rattus fall into two qroups - the R. lutreolus group and the R. 

-us group. Of the karyotypic-forms encountered in the former 

group, that of R. lutreolus is probably most ancestral because it is 

identical to thzt of many Asian species of Rattus. Other karyotypic 

forms in the R. lutreolus qroup can be deri- follows: 

(1) E. tunneri tunne.yi and R. t. culmorum by a single fixed 

pericentric inversion: 

assimilis and R. f. coracius-by two fixed fusions:-(3T R.'leucop% 

cooktownesis by tfiree fixed fusions: and (4) R. leucopus leucopus 

by four fixed fusions. 

vi1 losissimus may possess the most ancestral karyotyoewith 2n=50 

(FN=60), from which R. s. colletti (2n=42; FN=60) is derived by four 

fusions and R. s. s&dTdum; FN=60) by nine fusions, four of 

which appear-to 6e homolosous with those R. s. colletti. - The 

11. The Rattus qroup. Chromosoma (Berl.) 61, 227-241. 

That of 

(2) 8. fuscipes fuscipes, R. f. qreyi, R. f. 

Of the R. sordidus groups, R. s. 

- 

karyotypic data are in accord with Taylor-ana Horner's (1973) 

suggestions that (1) R. t. tunneyi and R. t. culmorum belong to one 

species; (2) R. - lut.-luTeo- R. l;t.-velutinus belong to one 

species; (3) R. leu. leucopus and R. leu. cooktownen.sis belong to one 

species and (4) R. f. fuscipes, R.-f. greyi, R. f. assirnilis and R. 

f. coracius belong to one species. However, The-large karyotypicaifference between R. s. sordidus and R. s. coletti and R. s. 

vi1 losissimus may ixdiyate that these qrouis -to diTferent 

bioloqical species. - Supernunerarv or 6-chromosomes were found in R. 

-- - 

-- --- 

f. assirnilis and R. t. tunne i. 

Iieterozyqous for centdion. [AuthoTs'-abTGZZJA"sing1e R. t. culmorm was 
Belcheva, R. G. and D. T. Peshev 1979. Intersubspecific sex 010 

chromosome difference in Citellus citellus L. (Rodentia, Sciuridae). 

Experientia 35(5) :595-596. 

The autosomal karyotypes of all subspecies studied of Citellus 

citellus from Bulgaria do not differ. 

is different in 1 of the subspecies where it seems to have undergone a 

pericentric inversion. [Authors' abstract] 

The X chromosome, by contrast, 

Benado, M., M. Aguilera, 0. A. Reig and F. J. Ayala 1979. 01 1 

Biochemical genetics of chromosome forms of Venezuelan spiny rats of 

the Proechimys guairae and Proechimys trinitatis superspecies. 

Genetica 50( 2) :89-98. 

Spiny rats from Venezuela show an extensive karyotypic 

diversification (2n = 24 to 2n = 62) and little morphological 

differentiation. This study reports genetic distance, heterozygosity 

and polynorphism based upon 22 loci in semispecies and allospecies of 

the Proechimys quairae superspecies from N Central Venezuela, as 

compared with Pmys urichi, a member of the Proechim s trinitatis 

guairae complex are included, each characterized by karyotses of 2; = 

'-(Fundamental Nunber = 72), 2n = 48 (FN = 72), 2n = 50 (FN = 72), and 

2n = 62 (FN = 74). Proechimys urichi has a distinctive karyotype of 2n 

= 62 (FN = 88). The overall memue of Nei's genetic identity index 

for all pair-wise comparisons is I = 0.942 + 0.011. 

within the P. guairae complex is I = 0.969 T 0.033. 

between P. urichi and members of that compTex is I = 0.889 + 0.011. 

Within tFe Pmrae complex, increased genetic diverqence 7s 

correlated with higher karyotypic divergence. Heterozvqosity varies 

from H = 0.059 to H = 0.153, with a mean value of H = 0.088 + 0.014. 

The mean percent of pol_ynorphic loci is P = 18.2 + 3.9 after-the 

'0,95%' polymorphism criterion, and P = 20,5 + 5.7 after the '0.99%' 

criterion. These results are compared with similar data from fossorial 

and non-fossorial rodents. Spiny rats are non-fossorial, forestdwelling rodents which have undergone a speciation process with little 

genetic divergence and extensive chromosome rearrangements. [Authors'. 

abstract ] 

superspecies from eastern Venezuela. Four chromosome + orms of the P. 

Mean identity 

Mean identity - 

Bianchi, N. O., 3. Paulete-Vanrell, and L. A. De Vidal Rioja 1969. 012 

complement with 38 chromosomes in two South American populations of 

Rattus rattus. Experientia 25:1111-1112 

The chromosomal complements of 16 specimens of Rattus rattus 

from two populations, one from Argentina and the other from Brazil, 

are observed to be 2n=38. 

believed to have been derived from a European ancestor havinq 2n=42. 

[Abstract by GLW] 

The South American populations are 
Bosma, A. A. 1976. Chromosomal Ploymorphism and g-banding Patterns in 01

the wild boar (Sus - scrofa L.) from the Netherlands. 

46 : 391-399. 

Genetica 

Cytogenetic examination of 15 wild pigs (Sus scrofa L.) from the 

Nether1 ands has revealed intrapopulation po1ymKh-r the diploid 

chromosome nunber. Eleven pigs showed 2n = 36 chromosomes, three pigs 

showed 37 chromosomes, and one piq showed 38 chromosomes. The cause of 

these differences in chromosome nunber is discussed. 

With the aid of a Giemsa banding technique it is demonstrated that 

the "extra" submetacentric chromosome (chromosome No. la) of the wild 

boar is homologous with chromosomes Nos. 15 and 17 of domestic piqs. 

[Author's abstract] 

Bosma, A. A. 1978. The chromosomal g-bandinq pattern in the wart hog, 01

Phacochoerus aethiopicus (Suidae, Mammal ia) and its imp1 ications for 

the systematic position of the species. Genetica 49(1) :15-19. 

It appears from the G-banding patterns that the karyotypes of the 

wart hog (Phacochoerus aethiopicus), the European wild boar (Sus 

scrofa) and the domesticscrofa) - are very similar, tF 

differences in karyotype between these species consisting of 

Robertsonian translocations only. 

sugqests a close phylogenetic relationship between the genera 

Phacochoerus and - Sus. 

This close similarity in karyotype 

[Author's abstract] 

Bowers, J. H., R. 3. Baker, and M H. Smith 

electrophoretic, and breeding studies of selected populations of deer 

1973. Chromosomal, 

mice (Peromyscus manicul atus) and bl ack-eared mice. (P. me1 anot is) . - 

Evolution 27(3) ~378-386. 

An unusual amount of chromosomal polymorphism and geographic 

variation in chromosomes and electrophoretic pattern have been 

reported for Peromyscus maniculatus. Karyoloqical, electrophoretic 

and breeding data indicate that populations of Peromyscus from the 

Chiricahua, Pinaleno and Santa Catal ina Mountains in southern Arizona 

are conspecific with P. melanotis, not with P. maniculatus. These 

data also argue againzt the assumptions that-monomorphism in the 

isolated populations arose by drift. Rather they support a model of 

centr i f ugal spec i at ion. [ Authors ' summaryj 

01
Brown, R. J. 1974. A comparative study of the chromosomes of three 016 

species of shrews,, Sorex bendirii, Sorex trowbridqii, and Sorex 

vagrans. Wasmann J.m.32(033K 

Three subspecies of Sorex trowbrid ii, (S. t. trowbridqii, S. t. 

humboldtensis, S. t. montere -79- ensis ave ideztiyal karotypes (2: =-34, 

. bendirii karyotyped had 2n = 54 and 

FN = 70. Seven subspecies of S. YaGm-v. vagras, S. v. 

halicoetes, S. v. pacificus, S. v. yaquin<e,-S. v. permiliensis, S. v. 

bairdi, S. v. sztom k%-oTypmhoi izter- and 

intrapopzlaTiomiation. With the exception of 1 individual of S. 

v. bairdi with a diploid nunber of 53 (attributed to Robertsonian 

Tus-11 subspecies have a diploid nunber of 54. 

nunber varies from 58 and 59 in S. v. pacificus and S. v. a uinae at 

northern end. Graphically intermediate populaTio~s, S. v. bairdi and 

S. v. permilliensis have fundamental numbers varying from 58-62. The 

XarEtypic differences in the S. va ras complex are presunably due to 

38). The singTe specimen o 

-r .- 

- --+T 

-- 

- 

The fundamental 

the southern end of the ranqe to-62-and 63 in S. v. SetT'sus 4L91F- at t e 

pericentric inversions. [ AbsTract -+ rom Biological Abstracts] 

-- 

Bruere, A. N:, H. M. Chapman, P. M. Jaine, and R. M. Morris 1976. 

Origin and significance of centric fusions in domestic sheep. J. 

Heredity 67:149-154. 

The karyotypes of 731 sheep of various breeds were studied and 

considered in association with previous chromosome studies of 

domestic sheep. A high incidence of the T2 translocation was found 

in two pedigree flocks of New Zealand Romney sheep. 

flocks was established over 100 years ago and it is suggested that 

this translocation originated in the Romney Marsh breed of sheep in 

England. 

52tlt2 was reported for the first time. 

One of these 

A naturally occurring double translocation heterozygote 

A further flock of sheep of the Perendale breed was found with a 

hiqh incidence of dicentric chromosome fusion that was identified as 

the t translocation. The apparently comnon Occurrence of 

discussed in relation to karyotype evolution among both domestic and 

wild sheep. [Authors' smrnary] 

chrom a some polymorphism, due to centric fusions, in domestic sheep is 

017 
Bruere, A. N. and R. D. McLaren 1967. The idiogram of the sheep with 01

particular reference to secondary constrictions. Can. J. Genetic Cytol. 

9: 543: 553. 

Peripheral leucocyte cultures from 22 normal sheep aged less 

than two years have shown that the modal diploid chromosome number 

(54) was present in 87.44% of cells counted from a total of 1831. 

The modal number of 54 chromosomes is verified for five previously 

unreported breeds of sheep, includinq the Scottish Blackface, Grey 

Face, Clun Forest, Welsh Montain and Soay, Karyotype observations 

showed that the short arms of the X chromosome are noticeably larger 

than those on other members of the acrocentric group of chromosomes, 

and that the Y chromosome is clearly submetacentric. Secondary 

constrictions were seen on a number of the 333 "photographic" 

karyotypes prepared from the best metaphases of 3280 counted cell s. 

The incidence of these was highest on the six large metacentric 

chromosomes and suggested sites of regular occurrence, which may be 

associated with nucleolus formation. An increase in the incidence of 

secondary constrictions was recorded in preparations made from 

cultures in which hypotonic sodiun citrate was used instead of 

hypotonic Hanks balanced salt sol ut ion. [Authors' summary] 

Bruere, A. N., D. L. Zartman, and H. M. Chapman 1974. The 

significance of the Gbands and C-bands of three different 

Robertsonian translocations of domestic sheep (Ovis aries). 

Cell Genet. 13:479-488. 

Cytogen. 

G-banding of three different Robertsonian translocations of 

domestic sheep (MI, MII, and MIII) has shown that the arm components 

of each are nonhomologous. 

translocation chromosomes showed heavy blocks of centric 

heterochromatin which were distributed evenly on either side of the 

centromeric constriction and which suggested a dicentric structure. 

The C-band pattern of the MI1 translocation was reqularly exocentric, 

suggesting that it may be formed by reciprocal translocation. 

need for caution on the use of the term "Robertsonian translocation" 

is discussed. The mounts of centromeric heterochromatin in each of 

the translocation chromosomes is apparently greater than in the 

regular metacentrics, which suggests that they are of a more recent 

oriqin. [Authors' abstract] 

The C-bands of the MI and MI11 

The 

0
Buettner-Janusch, J. and A. E. Hamilton 1979. Chromosomes of 020 

Lemuriformes: IV. Karyotype evolution in Lemur fulvus collaris (E. 

Geoffroy 1812). 

diploid nunber of 52; the nombre fondamental (n%bWchromosome 

arms) is 64. 

19 pairs of acrocentric autosomes and acrocentric sex chromosomes. 

Comparisons of the 2N = 52 complement with the 2N = 50 and 2N = 51 

karyotypes published by Hamilton, et al. (1977) supports the view that 

hybridization occurs naturally betGEe?i-animals with diploid numbers of 

50 and 52. 

h. J. Phys. Anthropol. 5m 363-366. 

The G[Giemsa]-banded karyotype of a feral L. fulvus collaris has a 

The karyotype consists of 6 pairs of biarmed autosomes, 

[Abstract from Bioloqical Abstracts] 

Bunch, T.D. 1978 Fundamental karyotype in domestic and wild species 021 

of sheep: Identity and ranking of autosomal acrocentrics involved in 

biarmed formations. J. Heredity 69:77-80. 

The fundamental karyotype of Ovis is described according to 

G-band analysis of Ovis viqnei, 0. mon, 0. orientalis, 0. musimon, 

-- 0. dalli, 0. canadex,nies (to inclzde the Massey'ii,md 

T3 translocations) and O.-nivico'la. The acrocentrics involved in 

their biarmed fusions were identified from G-banded idioqrams of 

Capra. 

centric fusions were: 

lower and upper arms, respectively. The siqnificance of partial 

homology in the M4 and T3 biarmed chromosomes as an evolutionary 

occurrence leading to speciation is discussed. [Author's sumnary] 

Acrocentrics involved in the Ml, M2, M3, M4, T1, T2 and T3 

115, 3/10, 419, 11/17, 8/30, 12/15, and 11/29, 

Bunch, T. D. and W. C. Foote 1976. Chromosomes, hemoqlobins, and 

transferrins of Iranian domestic sheep. J. Heredity 67:167-170. 

Twelve breeds of Iranian domestic sheep were cytoqenetically 

analyzed. The diploid chromosome number of 2n=54 is identical to 

that of most breeds of domestic sheep, which is comprised of 3 pairs 

of metacentric and 23 pairs of acrocentric autosomes. The sex 

chromosomes consist of a large acrocentic X and a small bi-armed Y. 

Hemoglobin analysis by isoelectric focusing for these 12 breeds ' 

resulted in the identification of 2 protein fractions: Hb AB and B. 

The A allele was observed in 7 percent of the 506 sheep sampled, in 6 

of the 12 breeds, and then only in the heterozygous form. 

Transferrins were analyzed by starch-gel electrophoresis and 

comparisons made between breeds. Nine alleles, pooled frequency of 

occurrence in descending order, 6, C, D, A, M, E, G, P, and I were 

identified and resulted in 19 phFnoTypFs. Tfs -A, E, C, and IT were 

observed in all breeds, whereas I was found in 2, 'ii; in 1, M yn 8, E 

in 6, and P in 2. Significant dTfferences in aIIeTic occu7rence of 

Tfs A, B, c, and D were observed in seven breeds. 

- 

--- [Authors' sumary] - 

022 
Bunch, T. D., W, C. Foote, and J. 3. Spillett 1976. Translocations 02 

of acrocentric chromosomes and their implications in the evolution of 

sheep (Ovis). -Cytogen. Cell Genet. 17:122-136. 

Cytogenetic evidence suggests that the caprids (sheep and qoats) 

evolved from a common ancestor with a 2n=60 karyotype. Although 

goats (Capra) retained the primitive 2N=60 karyotype, sheep (Ovis) 

underweEsequent i a1 reduction in the nunber of chromosomesF 

means of acrocentric translocation. The formation of the first 

metacentric autosome (Ml) occurred in the aoudad (Amnotragus) and 

urial (0. viqnei), resulting in a 2N=58 karyotype. 

homolog&s, which implies both genotypes arose from a comnon 

ancestor, possibly a rupicaprid. Based on G-bands, acrocentric 

chromosomes 1 and 7 of the 2n=60 karyotype formed the Kl. The X 

chromosome, which is the second longest acrocentric in the 2n=60 

karyotype, became the longest acrocentric in hotra us and Ovis 

(2n=58). The second pair of metacentrics to -+- evo ve, which isnked 

in the M3 position of the 2n=54 karyotype, resulted from the 

translocations of acrocentric chromosomes 4 and 14 or 15 in the 2n=60 

karyotype. The M2 was the third pair of metacentrics to be formed 

and resulted from the translocat ions of acrocentric chromosomes 3 and 

12 or 13 in the 2n=60 karyotype. The G-bands of all 2n=54 karyotypes 

are homologous, which indicates origin from a comnon ancestor. 

Evidence is presented that suggests a prezygotic selection is 

bringing about a reduction in diploid chromosome numbers. The 

possible roles of fission and fusion in the karyotypic evolution of 

- Ovis are discussed. [Authors' abstract] 

Th e G-bands are 

Bunch, T. D., C. F. Nadler, and L. Simnons 1978. G-band patterns, 02

evolutionary relationships with sheep and goats. J. Heredity 

hemoglobin, and tranferrin types of the bharal: Chromosomal 

69 ~316-320. 

G-band patterns of the bharal (Pseudois nayaur), 2n = 54, were 

compared with those of wild sheep (Omi stonei), 2n = 54, and 

the Persian wild goat (Capra hircusnn-. 

longer segments of the biarmedmosomes of Pseudois were similar 

to those of the longer biarmed segments of Ovis, whereas the shorter 

segments differed. Biarmed chromosomal segments had G-band 

homologies with specific acrocentric autosomes of Capra and were 

ranked as follows in descending order of relative lengths: Pseudois 

1:4/13; 2:1/27 and 3:3/29; and for Ovis 1:1/5; 2:3/10 and 3:nm 

ratios and relative lengths of the biarmed chromosomes were compared. 

The Y chromosome of Pseudois is a small biarmed chromosome that 

resembles those of Capra and Ovis. 

indistinguishable from Hb B of Ovis using starch-gel electrophoresis. 

The transferrin "zone pair" of -dois migrated more slowly in 

starch-gel electrophoresis than do any of the known transferrin types 

ir? sheep and goats. 

Patterns of the 

- 

Hemoglobin B was observed in the bharal and was 

We refer to this. transferrin as Pseucivjs Tf A. 

ev-olutir, i41e~+~i43n-. r&u;thars' 

smrrryT 
Bush, G. L. 1975. Modes of animal speciation. Annual Rev. Ecol . Syst. 6~339-364. 

The author presents an excellent treatise on speciation in 

animals. 

chromosomal rearrangements in speciation. Only this portion of the 

paper is abstracted here. 

occurs by subdivision, there is little or no directed chromosomal 

evolution. 

generally not associated with speciation. In allopatric situations 

where speciation occurs by the founder effect, and also in sympatric 

situations, chromosome rearrangements may or may not be associated with 

speciation. In parapatric situations, chromosome rearrangements are 

frequently associated with speciation. These broad generalities apply 

to the entire animal kingdom, and are not limited to any particular 

group (such as mammals). [Abstract by GLW] 

A portion of his discussion deals with the role of 

In allopatric situations where speciation 

Any chromosomal rearrangements that may be present are 

025 

Bush, G. L., S. M. Case, A. C. Wilson, and J. L. Patton 1977. Rapid 026 

speciation and chromosomal evolution in mammals. Proc. Natl. Adac. 

Sci. USA 74(9):3942-3946. 

To test the hypothesis that population subdivision into small 

dernes promotes both rapid speciation and evolutionary changes in gene 

arrangement by inbreeding and drift, we estimated rates of speciation 

and rates of chromosomal evolution in 225 genera of vertebrates. 

Rates of speciation were estimated by considering the number of 

living species in each genus and the fossil record of each genus as 

well as information about extinction rates. Speciation rate was 

strongly correlated with rate of chromosomal evolution and average 

rates of speciation in lower vertebrate genera were one-fifth those 

in mammalian genera. 

speciation rates may generally have small effective population size 

(Ne), whereas large Ne values may be associated with 

karyotypical ly uniform genera and slow rates of speciation. 

Speciation and chromosomal evolution seem fastest in those genera 

with species organized into clans or harems (e.g., some primates and 

horses) or with limited adult vagility and juvenile dispersal, patchy 

distribution, and strong individual territorial ity (e.g., some 

rodents). This is consistent with the above hypothesis regarding the 

evolutionary importance of demes. [Authors' abstract] 

Genera with high karyotypic diversity and rapid 
Caire, W. and E. G. Zimmerman 1973. Chromosomal and morphological 02

variation and circular overlap in the deer mouse, Peromyscus 

maniculatus, in Texas and Okl ahoma. Syst . Zoo1 . 24( 1) :89-95. 

Mitotic chromosomes were analyzed from 74 specimens representing 

three subspecies of Peromyscus maniculatus from Oklahoma and Texas. 

Two distinct chromosomal forms were found. One form, occurring 

predominantly north of the Red River in Oklahoma and representinq - P. 

- m.ozarkiarum, had 11 pairs of subtelocentric, six pairs of 

metacentric or submetacentric and six pairs of acrocentric autosomes. 

A second form, occurring south of the Red River in Texas and 

representing P. m. pallescens, had ten pairs of subtelocentric, eiqht 

pairs of metacenTric or submetacentric, and five pairs of acrocentric 

autosomes. Both karyological types were found in the same habitat at 

two localities in north-central Texas, but no chromosomal h.ybrids 

were found in this area. Meiotic analysis of hybrids from laboratory 

crosses between the two types indicated partial sterility. A 

morphological analysis revealed that qene flow is restricted in 

eastern Okl ahoma, west-central Texas, and eastern Texas. The most 

plausible explanation for the observed variation is the existence of 

circular overlap. [Authors' abstract] 

Capanna, E., M. V. Civitelli, and R. Nezer 1970. The karyotype of 02

38-chromosome cment. Exper ient ia 26:422-425. 

the black rat (Rattus rattus L.): another population with a 

In one Italian population of Rattus rattus, the karyotype is 

constantlv characterized by a diploid commt of 2n=38. 

Differences in diploid nunber in populations of rats collected in 

other parts of the world are believed to be caused by independent 

centric fusions involvinq both Robertsonian and non-Robertsonian 

translocat ions. [Authors' abstract] 

Craig-Holmes, A. P., F. B. Moore, and M. W. Shaw 1973. Polymorphism 0

of hman C-band heterochromatin. I. Frequency of variants. Amer. J. 

Human Genet. 25:181-192. 

The constitutive heterochromatin of human chromosomes is subject 

to a high rate of variation. 

C-band variants were discovered. In two families, three of six 

var i ants were tr ansm i t ted . 

Amonq 20 unrelated individuals, 31 

The high frequency of variants in constitutive heterochromatin is 

presumably due to the highly repetitious DNA present in these regions. 

The polymorphic spectrum of C-band heterochromatin, therefore, should 

be dependent on the degree of heterogeneity of the repetitious families 

within a region as well as the complexity of the repeated sequences. 

[Authors' abstract] 
Craig-Holmes, A. P. and M. W. Shaw 1971. Polymorphism of human 

constitutive heterochromatin. Science 174(4010) : 702-704. 

Genetic polymorphism has been demonstrated in man for many 

characteristics including blood groups, serum proteins, tissue 

enzymes, and hemoglobins. A class of chromosomal polymorphism 

involvinq constitutive heterochromatin has now been found. 

the use of a special technique that permits visualization of 

heterochromatin, seven heterochromatin variants have been found among 

four individuals. 

variability of heterochromatin in the population. [Authors' 

abstract] 

Through 

These results suggest a very high frequency of 

030 

Davis, B. L. and R. J. Baker 1971. Chromosome morphology of North 031 

American Rattus rattus (L.) IMuridae). Cytologia 36:417-420 

Although the present distribution of Rattus rattus is 

cosmopolitan, the three subspecies reportem Northmerica were 

introduced by man in recent times. 

collected from feral populations. Karyotypic preparations were by in 

vivo culture of bone marrow, sodium citrate, Carnoy's fixative, blaz 

=method; stain was by Giemsa's blood stain. Specimens of the three 

subspecies all had the same karyotype, characterized by a diploid 

number of 2n=38. Subspecific assignment was made on the basis of 

characteristics of external morohology. 

Specimens for our study were 

[Abstract by GLW] 

Dev, V. G., 0. A. Miller, R. Tantravahi, R. R. Schreck, T. H. Roderick, 

B. F. Erlanger, and 0. J. Miller 1975. Chromosome markers in Mus 

musculus: differences in C-bandinq between the subspecies M. m. 

musculus and M. m. molossinus. Chromosoma 53: 335-344. 

patterns of metaphase chromosomes of two subspecies of Mus musculus 

were compared. 

molossinus (a su6species from southeast Asia) had similar q-bznd 

patterns along the length of the chromosomes, but differences were 

observed in the centromeric reqion of some chromosomes. The two 

subspecies had very different distributions of C-band material. 

Antibodies to 5-methylcytosine were bound to reqions of the chromosome 

corresponding to the C-bands in each animal. These findinqs support 

the idea that satellite DNA, which is concentrated in the C-band 

region, changes more quickly than bulk DNA. The interfertility of 

these two subspecies permits the development of a musculus strain 

carryinq normal marker chromosomes for genetic stumAuthors' 

abstract] 

032 

- 

-- 

-- 

Quinacrine (Q-band) and centromeric heterochromatin (C-band) 

M. m. musculus (the laboratory mouse) aTM. m. 
Duffy, P. A. 1972. Chromosome variation in Peromyscus: A new 

mechanism. Science 176:1333-1334 

Differences in total chromosome lengths between two 

karyotypically divergent qroups of Permoyscus maniculatus are taken 

as evidence for an addition-deletion mechanism of chromosomal 

variation in the species. The differences may be due in part to 

variation in the amount of constitutive heterochromatin present in 

the two karyotypes. [Author's abstract] 

033 

Dutrillaux, B., A.M. Fossee, and G. Chauvier 1979. Cytogenetic 034 

study of six species or sub-species of Mangabeys monkeys (Papiinae 

cercopithecidae). Ann. Genet. 22(2) :88-89 

sub-species are studied and compared. By comparison with the other 

Papiinae previously studied (Papio and Macaca) Lophocebus albiaena 

and L. aterrimus possess very similar karyotypes, differing at most 

by a-pericentric inversion of the Y chromosome. 

Chromosome banding patterns of six Mangabey species or 

The other four, 

Cercocebus torquatus torquatus, 

aaleritus and C. 4. chrvsoaaster aifTer bv a comDlexrearranaement of 

C. t. fuliginosus, C. galeritus 

I. 

Ghrornosome No TO 2nd by acquisition of heterochromatin on chromosome 

No 12. 

Lophocebus nor between the four Cercocebus. Cytoqenetic criteria are 

thus in agreement with the morphological, immunological and 

hematological data, separating the two genuses,[sic.] and placing 

Lophocebus closer to the Papio and Macaca than tflercocebus. 

IAuthors' abstract] 

No difference was detected nor [sic.] -between the two 

Dutrillaux, B. and Y. Rumpler 1977. Chromosomal evoluation in 03

Malaqasy lemurs: 11. Meiosis in intra- and interspecific hybrids in 

the genus Lemur. Cytogenet. Cell Genet. 18(4) :197-211. 

The chromosome analysis of meiosis in 4 lemurs, L. fulvus fulvus, 

- L. f. collaris, L. f. albocallaris and L. macaco and Parmr hybrid 

crosses is reporTed: In metaphase I, t'F-iv- and chain elements 

were detected and identified with T-banding. The absence of chain 

multivalents elements in the pachytene stages of hybrid meiosis, where 

a chain is detected later in diakinesis, may offer evidence on the 

possible existence of a 2-step pairing mechanism in meiotic homoloq 

pairing. Considerations about the role of the chromosome 

rearrangements in establishing a gametic barrier in speciation are 

developed. [Abstract from Biological Abstracts] 
Elder. F, F. B. 1978. Chr-1 evoP&~Oft of the cuttun rats, Genus 036 

Sigmodon (Rodentia, Muridae). 

The Chromosomes of five species of cotton rats were analyzed using 

C- and G-bandinq techniques. The chromosomal evolution was discussed 

in the context of known ecological and populational parameters of the 

species. Limited paeontoloqical and biochemical data suwest that 

chromosome evolution has proceeded rapidly, the time span from an 

ancestral form (Siqmodon hispidus) to the derived S. arizoniae being 

about 100,000 yenrm relationships were identified and 

biogeography and evolution of the species were discussed. A phylogeny 

based on the available data was proposed. 

Diss. Abstr. 39(5) :2171-8 

[Abstract by GLW] 

Evans, H.J., R. A. Buckland, and A. T. Sumner 1973. Chromosome 

homology and heterochromatin in goat, sheep, and ox studies by 

banding techniques. Chromosoma 42:383-402. 

Peripheral blood lymphocyte metaphase chromosomes of three 

Bovoidean species have been studied using Quinacrine flourescence and 

Giemsa banding techniques to give Q-, G-, and C-bandinq patterns. Qand G-banding characteristics, coupled with chromosome lenqth, 

enabled all of the chromosomes in each of the chromosome complements 

to be clearly distinquished, although some difficulties were 

encountered with the very smallest chromosomes. 

G-bandinq patterns between the species revealed a remarkable deqree 

of homoloqy of banding patterns. 

acrocentric autosomes of the domestic sheep (2n = 54) was represented 

by an identical chromosome in the goat (2n = 60) and the arms of the 

3 pairs of sheep metacentric autosomes were identical matches with 

the remaininq 6 goat acrocentrics. 

was evident for all but two of the autosomes in the ox (2n = 60). 

This homology between sheep metacentric and goat acrocentric elements 

confirms a previously suggested Robertsonian'variation. 

homology in G-banding patterns between these related species 

indicates that the banding patterns are evolutionarily conservative 

and may be a useful quide in assessinq interspecific relationships. - 

The centromeric heterchromatin in the autosomes of the three species 

was found to show little or no Q- or G-staining, in contrast to the 

sex chromosomes. This lack of centromeric staininq with the 

G-technique (ASG) contrasts markedlv with results obtained with other 

mammalian species. However, with the C-banding technique these 

regions show a normal intense Giemsa stain and the C-bands in the sex 

chromosomes are inconspicuous. 

heterchromatin in the sheep metacentric chromosomes is considerable 

less than in the acrocentric autosomes or in a newly derived 

metacentric element discovered in a goat. It is suggested that the 

pale G-staining of the centromeric heterochrornat in in these species 

miqht be related to the presence of GC-rich satellite DNA. 

abstract] 

A comparison of 

Each of the 23 differenct 

A similar interspecies homology 

The close 

The amount of centromeric 

[Authors' 

037 
Forejt, J. 1973. Centromeric heterchromatin polymorphism in the 

house mouse. Chromosoma 43( 2) :187-201. 

Polymorphism of Giemsa-specific centromeric heterochromatin 

(C.H.) has been described in the laboratory and wild mice. All 

examined wild mice and inbred mouse strains display some chromosomes 

with considerably reduced or enlarged C.H. regions. The quantity of 

C.H. could be an inherent property of a chromosome as inferred from 

(a) the finding of the identical C.H. pattern within inbred strains, 

(b) the finding that two genetically related inbred strains, C3H and 

CRA, separated from each other for more than 150 qenerations, possess 

the same tw chromosome pairs with tiny C.H. marker reqions. These 

chromosomes were identified as No. 1 (1.g. XIII) and No. 14 (1.9.111) 

by means of T(14;15)6Ca translocation, and C- and G-band analysis. 

The neutrality of C.H. polynorphism in murine genome is inferred from 

the "heterozygosity" for the C.H. variants found in all studied wild 

mice. The possible relationship of C.H. polynorphism to the 

centromere interference phenomenon is hypothesized. 

abstract ] 

[Author's 

0

irechette, A. G. and S. M. Jalal 1971. Karyological study of two 0

subspecies of the red back vole. Mammalian Chromosome Newsletter 

12 :38-39. 

Karyotypes of Clethrionomys qapperi qapperi Viqors and 

Clethrionomys gapperi lorinqi V. -were established from 

specimens collected within their respective geographical ranges from 

south-western Ontario ( Q u e t i c o P r o v i n c i a l Park) and northeastern 

North Dakota. 

Chromosomal counts revealed the same modal diploid number of 56 

for both subspecies. Karyotypic and idiogram analysis indicated the 

presence of one pair of large subtelocentrics, one pair of small 

metacentrics, 25 pairs of telocentrics, a telocentric X, and a 

subtelocentric Y. This karyotype study, at the subspecific level, 

differs from Mathey's (1956) who reported 56 acrocentric chromosomes. 

The presence of the small autosomal metacentric at the species level 

was confirmed. The NF ("nombre fondamentale") should thus be changed 

from 56 as previously recorded to 48 without consideration of the 

subtelocentric chromosomes. [Abstract by GLW] 
Garcia, M., R. Miro, L. Freitas, and J. Egozcue 1978. Bandinq 040 

patterns of the chromosomes of Cebus apella: Unstable chromosomes and 

pericentric inversion. Folia Primaton(3) 996-205. 

Quinacrine- and Giemsa-banding studies of the chromosomes of C. 

apella permitted the establishment of a pattern for the sp.ecies ad 

med an extreme chromosome unstability [sic.] in the specimens 

studied. The male showed a pericentric inversion in 1 of its 

chromosomes, while the female had 4 different cell lines, with multiple 

structural changes involving several pericentric inversions and 1 

deletion. The existence of a marked intra- and interspecific 

chromosome variability is indicated. 

pbs t r ac t s] 

- 

[Abstract from Biological 

Gileva, E. A. 1973. 6-chromosomes, unusual inheritance of sex 04 1 

chromosomes, and sex ratio in the Artic lemning (Dicrostonyx torquatus. 

torquatus Pal 1. 1779) . Dokl ady Akad. Nauk SSSR, 213(4) :952-955. 

The karyotype of the Arctic lemming is characterized by complex 

variability. In particular, the diploid number of chromosomes in the 

68 animals examined varied from 45 to 51, primarily because of the 

presence of B-chromosomes. 

Apparently, all male Arctic lemnings each have two true 

X-chromosomes. This situation differs in principle from the system of 

multiple X=chromosomes known in mamnals up to the present, where males 

have only one true X-chromosome, and the second is an autosome, lacking 

its homolog because of' its translocation into a Y-chromosome. It is 

known that polysomy with regard to the X-chromosome leads to sterility 

in male mammals. The normal fertility in male Arctic lemninqs may be 

due either to total genetic inactivation of one of the X-chromosomes or 

to peculiarities in the organization of the X-chromosome. 

The karyoloqical differences between the two forms discussed (D. 

- t. torquatus- and-the Nearctic forms of D. torquatus) are very larqe-iin 

- 0. t. stevensoni 2n = 34, NF = 54), and-therefore their inclusion in 

the-same species must be regarded as doubtful. [Abstract by GLW] 
Gladkina, T. S. 1972. Georqaphical variability of two subspecies of 042 

comnon vole (Microtus arval is). Zoo1 . Zh 51(2) :267-279. 

Investigations were performed during 5 generations on two comnon 

vole subspecies; M. arvalis caspicus Ogn. from the Kustanai region 

and M. a. dup1icaTus-em1 Born with different karyotypes 

from-thz Leningrad and Kal iningrad regions respectively. 

subspecies were distinguished by ecologo-physiological features of an 

adaptive character. However, adaptation mechanisms are different in 

them and are similar to those of rodent species populating the same 

regions. Temperature adaptations in subspecies are of a 

comparatively stable character. The geoqraphical forms of M. a. 

duplicatus did not have any distinctions as to the characters under 

investigation in spite of differences in caryotypes [sic.]. On the 

whole, a high capability for both phenotypical and gexypical 

variability is characteristic of comnon vole which ensures its fast 

adaptation to the new condition of tilled lands. 

The 

[Author's sumnary] 

Greenbaum, I. F., R. J. Baker, and J. H. Bowers 1978. Chromosomal 04

homology and divergence between sibling species of deer mice: 

Peromyscus maniculatus and Peromyscus melanotis (Rodentia, Cricetidae). 

Evolution 32(2) :334-341. 

Chromosomal homology and amount and location of heterochromatin, 

as revealed by G-banding and C-bandinq analyses, were examined for 

Perm scus manicul atus bairdi i and P. me1 anot is. These procedures zmh- etermination of interspecific homology of all euchromatic 

autosomal segments. Autosomally, the karyotypes of P. maniculatus 

bairdii and P. melanotis are related as follows: 

chromosomes are unchanged, five pairs differ in the presence (in P. m. 

bairdii) and absence (in P. melanotis) of heterochromatic short a7rns; 

and six pairs differ by pericentric inversions. The acrocentric 

condition of two pairs of chromosomes found to be polymorphic in - P. 

maniculatus is homologous to the acrocentric condition of the 

correspondinq pairs in - P. melanotis. 

12-pairs of 

Intraspecific polymorphism in our sample was restricted to two 

pairs of autosomes in P. m. bairdii. One polymorphism involves 

heteromorphism for a h<teyoc~c short arm, whereas the other may 

be either the result of a pericentric inversion, translocation of a 

small segment, or polymorphism for undetected heterochromatin. 

In P. maniculatus heterochromatin is centromeric on all 

chromosomes. 

heterochromatic (one pair is reported polymorphic for the 

heterochromatic short arm). 

and the Y are heterochromatic. The amount of C-band material in P. 

melanotis is reduced relative to that characteristic of the P. 

maniculatus examined. Autosomal heterochromatin in P. melanztis is 

confined to the centrpmeric regions. 

Additionally, the short arms of five autosomal pairs are 

In both species, the short arm of the X 

- 

- 

Heterochromatic arms and pericentric inversions are important in 

the evolution of the karyotype of the genus. [Authors' sumnary] 
Greenbaum, I. F., R. J. Baker, and P. R. Ramsey 1978. Chromosomal 044 

evolution and the mode of speciation in three species of Peromyscus. 

Evolution 32(3) :646-654. 

Analysis of G- and C-banding patterns of Peromyscus polionotus 

reveals intra-specific variation in three chromosome pairs (numbers 

16, 18 and 19). All three cases involve segments of heterochromatin. 

Chromosomal homologies (based on G-banding patterns) between P. - 

polionotus, P. maniculatus and P. melanotis suggest that the 

ancestral karyotype for the thrze species was composed of a large 

number of acrocentrics (probably near 30). The data indicate the P. 

maniculatus and P. polionotus have increased the number of biarmedchromosomes in tKeir karyotypes by additions of heterochromatin and 

pericentric inversions. Four inversion product are unique to P. 

maniculatus, whereas two inversions are shared by P. maniculatzs and 

P. polionotus. Peromyscus maniculatus and P. poliznotus also share 

aerived heterochromatic additions. These c6ta indicate that P. 

maniculatus and P. polionotus had a comon ancestor after P. 

melanotis diverq5d from the line. Chromosomal data for theTe three 

species are discussed in light of the centrifugal speciation model. 

As predicted by this model, P. maniculatus has the most derived 

karyotype and the most intraspecific variation. [Authors' sumary] 

- 

Gropp, A. A,, 3. Markwong, J. Marshall, and Y. J. Kim 1972. 

Robertsonian chromosomal variation in the lonqtailed tree mouse 

(Vandeleuria) . Z. Zool. Syst. Evolutionsforsch lO(3) :210-214. 

identified. The occurrence of an autosomal polymorphism of the 

Robertsonian type is suggested. The presence of Robertsonian variation 

may be linked with the existence of different local forms or subpsecies 

of this widely distributed taxon. 

Ab st r ac ts] 

In Vandeleuria oleracea the sex chromosomes could not be 

[Abstract from Biological 

045 
GroDD. A.. J. Marshall, and A. Markwong 1973. Chromosomal findings in 04

the' spiny-mice of Thai 1 and (qenus - Mus)-and occurrence of a complex 

intraspecific variation in - M. shortridqei. Z. Saugetierkunde 

38( 3) ~159-168. 

The karyotypes of M. shortridqei and M. ahari belonging to the 

subgenus Leggadil la and-Coelomys respectively, her radically from 

those of most members of the genus Mus. A complex and mixed type of 

intraspecific polynorphism is responsible for a numerical and 

structural variation of the chromosomal complement of M. shortridgei 

(2n = 46-49; N.F. = 48-52), but the occurrence of supeFnumerary 

autosomes seems to be the most prominent feature besides pericentric 

inversion. -On the contrary, M. pahari, is revealed by the study of a 

limited nunber of specimens beTonging to one colony, was shown to 

posses% a stable karyotype with 48 acrocentric chromosomes. [Authors' 

summary]. 

- 

Hafner, D. J., J. C. Hafner, and M. S. Hafner 1979. Systematic status 04

of kangaroo mice, genus Microdipodops: Morphometric, chromosomal, and 

protein analyses. J. Mamnalogy 60(1) :1-10. 

The systematic status of Microdipodops megacephalus and M. 

pall idus is investiqated at the genic, karyotypic, and morphologic 

levels at a purported hybrid zone. Instances of hybridization are 

shown to be either nonexistent or present at extremely low levels 

(<0.6%), and full specific status of the two taxa is affirmed. Amounts 

OT genic variability and pol-morphism are approximately equal for the 

species (H = 0.064) and fall well within the ranqe observed for other 

vertebrate species. Estimates of the timinq of the speciation event, 

based upon alloz-mic data, range from 1.2 to 6 million years before 

present. Possible means of ecological separation of the two sibling 

species in sympatry are suqqested. [Authors' abstract] 

Haiduk, M. W., J. W. Bickham, and 0. J. Schmidley 1979. Karyotypes O

of six sDecies of Oryzomys from Mexico and Central America. J. 

Mammalogy 60( 3) : 610-615. 

The karyotypes of six species of Oryzomys from Mexico and 

Central America were described. Data were analyzed in light of the 

groupings proposed by Gardner and Patton (1976); additionally, a 

prel iminary comparison of G- and C-banding was presented. 

b.~ GLWI 

[Abstract 
Halkka, L. and 0. Halkka 1974. Karyotype Q- and G-banding in three 04 9 

species of the genus Sorex. Hereditas 78(2):314. 

In Kuhmo. NE Finland. the Sorex species minutus. caecutiens and 

isodon, all with 2n=42, live apparently sympatrically in the forest 

-the river Kuusijoki. The qoniomitotic metaphase chromosomes of 

these species were analyzed with-the Q- and G-banding techniques. 

The chromosome sets of caecutiens and isodon were found to be 

quite similar. 

chromosomes are mostly meta- or submetacentric, the small ones mostly 

telo- or subtelocentric. 

has several telocentrics among the large chromosomes of the set, and 

accordingly the NFa is only 54. 

The NFa is 60 in both species, and the large 

In contrast to these two species, minutus 

A nunber of chromosome arms appear similar in banding patterns 

in all three species, but the parallelisms are much more frequent 

between caecutiens and isodon than between these and minutus. The 

diverqence between the 

from a nunber of pericentric inversions and possibly also from 

translocations. The direction of the change has perhaps been towards 

a more metacentric condition at the expense of telocentrics. If this 

is true, then the caecutiens and isodon sets could be divergent 

derivatives of a highly telocentric ancestral condition from which 

the minutus set deviates less than these two. 

and NFa 54 karyotypes pm results 

The chromosal differences are certainly sufficient to keep all 

three species reproductively isolated. Primarily, however, 

hybridization of these sympatric species appears to be prevented by 

psycho1 og i c a1 i sol at ion. [Authors ' abstract] 

Holden, H. E. and H. S. Eabry 1970. Chromosomes of Sylvilaqus 

floridanus and S. t r a n s i t i o n a l i s . J. Mammalogy 51(1) :166-168. 

The karyotypes of four species of S lvilagus were studied. 

Three species possessed a diploid chromosome + num er of 42; the fourth 

(S. transitionalis) had 2n=52. Because of the great difference in 

cbomosome number, it is difficult to explain an evolutionary 

mechanism; further, the high proportion of biarmed chromosomes 

belongs to the species with a higher diploid number. 

centric fusion theory can not explain the divergence. 

GLW] 

- 

Therefore, 

[Abstract by 

050 

c 
HSU, T. C. and F. E. Arrighi 1966. Chromosomal evolution in the 051 

genus Peromyscus (Cricetidae, Rodentia). Cytogenetics 5:355-359. 

The diploid number of species and subspecies in genus Peromyscus 

is 48. The number of total chromosome arms (fundamental number), 

however, varies from 56 to 96. Apparently the changes were 

accomplished by reciprocal translocations and pericentric inversions, 

but not by the Robertsonian process. [Authors' abstract] 

Hsu, T. C. and F. E. Arrighi 1968. Chromosomes of Peromyscus 

(Rodentia, Cricetidae) 1. Evolutionary trends in 20 species. 

Cytogenetics 7:417-446. 

The somatic metaDhase chromosome constitutions of 20 Peromvscus 

species (188 spec imens) were described and compared. 

(Ochrotomys) nuttal i, which has been considered by many mamnalogists 

as a member of a separate genus, Ochrotomys, has a diploid number of 

52. All other 19 species possess a diploid number of 48. The lowest 

Permyscu; 

number of total chromosome'arms is 56 IP. crinitus and P. bovlei) and 

the highest is 96 (P. eremicus and P. cZll'atus).Since--thnurii6er of 

centromeres remains-the same. Rober-fsonian fusion is not found in the 

genus Peromyscus. 

always a larqe, biarmed element, but the Y chromosome is extremely 

The X chromosome, with one doubtful exception, is 

variable in size as well as in centromeric position. 1ntraspecif:c 

pol-morphism is extensive in many species, and in some cases, e.g., 

-~ P. truei, cytological data strongly suggest that there may be more 

than one species under one name. The survey indicates that it may be 

extremely profitable to use Peromyscus cytology to analyze species 

relationships, population structure, population dynamics and other 

evolutionary processes. Phylogenetic relationships among the species 

thus far analyzed are discussed. [Authors'abstract] 

052 

Hsu, T. C. and F. E. Arrighi 1971. Distribution of constitutive 05

heterochromatin in mammalian chromosomes. Chromosoma 34:243-253. 

Using a special staining technique, a survey of the chromosomes 

of many mammalian species showed that constitutive heterochromatin is 

present in all cases and that the heterochromatin pattern appears to 

be specific and consistent for each chromosome and each taxon. 

Usually heavy heterochromatin is found in the centromeric areas, but 

terminal heterochromatin is not uncommon. Occasionally interstitial 

heterochromatin bands occur. In some species, such as the Syrian 

hamster and Peromyscus, many chromosome arms are completely 

heterochromatic. [Authors' abstract] 
Jalal, S. M.? R. W. Clark, T. C. HSU, and S. Pathak 1974. Cytological 05

differentiation of constitutive heterochromatin. Chromosoma 

48: 391-403 

The constitutive heterochromatin, as demonstrated by the C-band 

technique, may be subdivided into a number of categories when other 

characteristics are considered. The responses to f lourochromes QM and 

33258 Hoechst, the behavior following G band staining, the repetitive 

DNA content, and many other criteria are useful for the classification 

of heterochromatin. 

species are presented to demonstrate that within each karyotype there 

may be several different types of C bands. In general, a correlation 

may also be made between GC-rich satellite DNA and dull (or negative) Q 

flourescence, and between AT-rich satellite DNA and bright Q 

f lourescence. [Authors’ abstract] 

The heterochromatin patterns of three mammalian 

Kato, H., K. Tsuchiya, and T. H. Yosida 1974. Constitutive 

heterochromat in of indian munt jac chromosomes revealed by DNase 

treatment and a C-banding technique. Can. J. Genet. Cytol. 

16:273-280. 

A karyotype of 

vaginalis, was descr 

No. 3 homologous pai 

a female Indian muntjac, Muntiacus muntjak 

ibed. The karyotype was unique in m. 1 and 

rs were hetermorphic with respect to the size of 

their secondary constrictions. In these pairs, one of the homologs 

always had a longer secondary constriction than that on the 

corresponding homolog. Heterochromatin in the secondary constriction 

region was visualized with difficulty by a C-banding technique, but 

was demonstrated clearly by a DNase treatment followed by Giemsa 

staining, which also revealed the size difference of the secondary 

constriction. Centromeric constitutive heterochromatin of No. 1 

chromosome was also found to differ in size between the homologs. 

the basis of the hetermorphic character of No 3 chromosome, or an 

X-autosome complex, it was possible to confirm autoradiographically 

that X-inactivation had occurred at random. [Authors ’ abstract] 

On 

05
Kennerly, T. E.? Jr. 

allopatric species of pocket gophers. 

1959. Contact between the ranges of two 

Evolution 13~247-263. 

The biological dynamics of contact between the ranges of two 

allopatric species of pocket gophers, Geomys personatus and G. 

bursarius ,in southern Texas were investigated. Ninety-one szecimens 

of G. bursarius and 113 specimens of G. personatus were trapped. 

Fievd work proceeded irregularly between December 12, 1952 and April 

6, 1955. The ranges of these two species approach one another at 

seven localities. At one trapping site both species were collected. 

No abrupt ecological changes coincide with the area of approximation 

between the ranges of the two species. Both species appear to have 

similar ecological requirements thouqh different tolerances. 

Indurate soils are thought to constitute the most formidable barriers 

to dispersal in the area studied. Interspecific competition is 

thought to be larqely responsible for separation of ranges. No 

evidence of hybridization or introgression was observed. 

summary] 

[Author's 

056 

Korobitsyna, K. V., C. F. Nadler, N. N. Vorontsov, and R. S. Hoffman 35

1974. Chromosomes of the Siberian snow sheep, - Ovis nivicola, and 

implications concerning the origin of amphiberingian wild sheep 

(Subgenus Pachyceros) . Quat. Res. 4:235-245. 

The chromosomes of Ovis nivicola, described for the first time, 

exhibit 2n=52, the lowesmploid number to be reported for wild 

sheep and qoats. The new chromosomal data, together with a review of 

the fossil history of the genus, lead us to conclude that the 

bighorned wild sheep (subqenus Pachyceros) evolved their distinctive 

characteristics while isolated in the ice-free Reringian refugium, 

and then migrated southward into western North Anerica when the 

P Authors' abstract] 

lacial barriers melted, as first suggested by Cowan (1940). 
Kozlovskii, A. I. 1974. Karylogical differentiation of North Pmerican 058 

subspecies of hoofed lemminqs. Doklady Biol. Science 219:499-502. 

The hoofed or coll ared lemning Dicrostonyx torquatus Pall. 

occupies the circumpolar area. 

recognized; in North America, 12 subspecies are distinguished. 

In Eurasia, 4 subspecies are usually 

In Dicrostonyx tor uatus vinogradovi a diploid set of 28 

chromosomes, N.F. = _e_f 54 is ound. The chromosomes represent three 

morphological types: meta-, submeta-, and subtelocentric. The 

chromosome compliments in each groups are described. The population of 

hoofed lemmings from the Chaunsk region proved to be polynorphous, 

having a diploid nunber from 57 to 60 chromosomes. This polymorphism 

is related to variability in the number of microchromosomes. 

Both continental subspecies (0. t. chionopaes and D. t. torquatus) 

display an evident similarity of kzryztypes, in which satFlo- and 

telocentric chromosomes of the most diverse sizes predominate, along 

with a few rather small metacentric chromosomes. Chromosome 

polymorphism is observed in populations of these sub-species. Their 

chromosome sets are similar, although they are by no means identical. 

The differences are quite essential, though not as striking as the 

differences among the northeastern subspecies. The differences in 

diploid number (45-51; 57-60) and in the form of sex chromosomes are 

most important and most notable in the subspecies investiqated. 

A comparison of the chromosome sets of hoofed lemnings from 

Vrangel I Is1 and and from North America suggests their qenetic affinity, 

since the karyotypes have the same structure and a similar N.F., 

despite their somewhat diverse diploid numbers. 

Chromosome sets have been described for half of the 16 subspecies 

of Dicrostonyx torquatus from the Arctic territory. 

chromosome sets may be noted: the "Asiatic t.ype" which seems older -- 

Two types of 

perhaps even the oriqinal type (at present this type is known only in 

the two continental sub-species - D. t. chionopaes and D. t. torquatus; 

and the "North American type," represented in the 5 nonrarytic 

subspecies and in the hoofed lemninq of Vranqel' Island. It should be 

emphasized that the chromosome sets of the northeastern subspecies of 

hoofed lemmings represent extreme variants of these types. 

-- 

H_ybridization between 0. t. stevensoni and four other subspecies 

with the "North American t$e"-of chromosome set (exsul, nelsoni, 

rubricatus, richardsoni) does not proceed beyond therst qeneration. 

This indicates the plausibility of regarding the nonarctic subspecies 

as a group of close, but reproductively isolated, independent species 

within the superspecies Dicrostonyx torquatus. [Abstract by GLW] 
Kozlovskii, A. I., and V. N. Orlov 1971. Karyological evidence for 

species independence of Sorex isodon Turov (Soricidae, Insectivora). 

Zoo1 . ZH 50( 7) :1056-1062- 

Karyotypes of S. isodon from the Maqadan and Kemerovo Districts 

were studied and were ident-ical. 

autosomes and 1 pair of sex chromosomes, NF = 68. 

9 pairs were metacentric, 5 pairs submetacentric, and 7 pairs subteloand telocentric. X-chromosome was telocentric and measured 5.47% of 

the haploid female set length. 

submetacentric. Quantitative characteristics of chromosomes were 

obtained when measuring 10 chromosome sets. 

identification of individual chromosome pairs by polykaryograms are 

discussed. The karyological data confirm the species status of S. 

isodon. Differences in chromosome sets of S. araneus and S. isoaon are 

so sharp that it appears impossible to distTnquish comnon Teatures. By 

the karyotype structure, S. isodon is most closely related to S. 

caecutiens, - S. unguiculat%.,and. -- Vir. [Abstract from Biological 

Ab st r act s] 

The chromosome set contains 40 

Among the autosomes, 

Y-chromosome was the smallest 

Possibilities of 

95 

Kral, B., E. Von Lehmann, and J. Zejda 1972. The hybrids of two 06

subspecies of the red-backed vole (Clethrionomys glareolus Schreb) . Zool. Listy 21( 1) :43-61. 

The karyotype, body and skull dimensions as well as fecundity 

were examined in the subspecies Clethrionomys glareolus qlareolus 

Schreber, 1780, Clethrionomys glareolus garqanicus Hagen, 1958, and 

gar anicus differs from the nominate form by 

larger body and sku c.1 s9--- imensions, amonq which the length of hind foot 

their crosses. 

and the height of skull appear to be dominant in male line hybrids. 

The same holds for the inheritability of fur colour. In the 

karyotype of C. g. garganicus, the sex chromosome Y is distinctly 

acrocentric to f<eb’ly subtelocentric and is the smallest one of the 

set. Therein the karyotype of C. 9. qaraqnicus differs from that of 

the hitherto examined central Eiropean populations of the nominate 

form, showing a mediocentric sex chromosome Y. Also this character 

of C. q. garganicus is inheritable in the male line. Both subspecies 

show acertain degree of sexual differentiation; that is to say, the 

hybrids in the female line of C. q. garqanicus appear to attain 

sexual maturity later than tho7e Tn the female line of C. q. 

glareolus. 

- se was unsuccessful . [Authors ’ summary] 

The crossinq of individuals of the F2 qeneratizn inter 
Kuliev, G. N., G. K. Kuliev and S. I. Radzhabli 1978. Karyot-we 06 1 

differences between populations of the water vole Arvicola terrestris 

(Rodentia, Cricetidae). Zool. ZH 57(9): 1409-1411. 

Water voles from 4 populations of the Caucasus and the Novosibirsk 

Distinct [sic] (USSR) were studied by means of G[Giemsa]- and 

C[const i tut ive heterochromat i nl- methods of different i a1 stain inq of 

chromosomes. In all animals studied 2n = 36. Differences were found 

by the values of NF (nunber of chromosome arms) and the size of 

Y-chromosome. In water voles of 3 Caucasian populations and in those 

from the Novosibirsk District NF = 72 and Y-chromosome is the smallest 

acrocentric of the set. 

the Azerbaijan SSR NF = 66 and Y-chromosome is almost X larger. 

comparative analysis of the chromosomes showed that the differences in 

the nunber of chromosome arms are due to the absence of short fully 

heterochromatic arms in 3 pairs of autosomes of the water voles from 

the Pushkinsky population. Y-chromosome of A. terrestris consists 

almost completely of heterochromatic materiar, and differences in the 

size of this chromosome determine the differences in the amount of 

heterochromatin as we1 1. [Abstract from Biological Abstracts) 

In water voles from the Pushkinsky region of 

A 

Lakhotia, S. C., S. R. V. Rao, and S. C. Jhanwar 1973. Studies on 062 

rodent chromosomes VI. Co-existence of Rattus rattus with 38 and 42 

chromosomes in south western India. C.fim -3-410 

Chromosomes of two subsoecies of Rattus rattus. namelv. .,1 R. - - - r. 

Wrouqhtoni and R. r. rufescens, from Sagar, Mysore hate, SouTh 

Western India, Kave been examined. R. r. wrouqhtoni has 42 

chromosomes typical of Rattus rattus; fiilefescens shows 2n = 

38. The karyotype of tmter is similar To Thanescribed by other 

workers for Rattus rattus populations with 2n = 38 from other parts of 

the world. 

chromosomes from India. It is proposed that the 2n = 38 karyotype is 

evolved from the basic Rattus rattus karyotype with 2n = 42 by centric 

fusion and pericentric inversi.r[Authors' summary] 

This ismirst report of Rattus rattus with 38 

Law1 or, T. E. 1974. Chromosomal evol ut ion in Peromyscus . Evol ut ion 063 

28( 4) ~689-692. 

Mechanisms for chromosome evolution were discussed in light of 

cytological evidence for their occurrence. Centric fusion, a process 

responsible for the reduction of the diploid number by creatinq 

biarmed chromosomes from the fusion of acrocentrics, has been 

considered a unidirectional evolutionary process. Cytological 

evidence provided by studies of mammalian chromosomes suqgests that 

"fission" by which biarmed chromosomes divide to create acrocentrics 

(and thus create a higher diploid number), has also played an 

important role in chromosome evolution. Evidence suggests no obvious 

trend of evolutionary change in the number of chromosomal arms. 

[Abstract by GLW) 
Lee, M. R., D. J. Schmidly, and C. C. Huheey 1972. Chromosomal 

variations in certain populations of Perom scus boylii and its 

sytematic imp1 ications. J. Mamnalogy -33fqx9T-m 

The karyotype of Perm scus boylii attwateri is contrasted with 

three taxa Fave identical-karyotypes. CFrozosomal and morphological 

evidence indicate that attwateri is specifically separated from the 

bo lii-rowle i-utahensis assemblage. Limited samples from Mexican 

htd P. boylii imply that current taxonomic alignments in 

this species require reevaluation. 

that of Peromyscus truei truei and unlike those of northern 

popul ationsdP.ilis. - [Authors' abstract] 

that of P. b. boylii, --5 p. b. row eyi and P. b. utahensis. The latter 

difficilis Comanche is described and it appears to be * 1 entica to 

The karyotype of Perom scus 

064 

Lyapunova, E. A. and N. N. Vorontsov 1978. Genetics of Ellobius 965 

(Rodentia): I. Karyological characteristics of 4 Ellobius species. 

Genet i ka 14( 11) :2012-2014. 

Ellobius is a cytogenetically unique group of animals. A review 

is given on the problem of sex determination in E. lutescens (2n = 17 

in both males and females) and on the chromosome-divergence of the 

Ellobius genus. Karyotypes of 83 animals of 4 Ellobius spp. are 

studied. E. lutescens from 2 Armenian regions has 2n = 17 in 8 and 

2. 

morphometrically observed. A male karyotype of E. fuscocapillus (2n = 

36), the only species having heteromorphic X and-Y chromosomes, is 

described for the 1st time. 

the length of the Y chromosome 1.7% of the haploid set length. 

distribution of west and east karyotypes is described on the basis of 

the investigation of E. talpinus from 14 points. No intrapopulational 

chromosome polynorphiTm was observed i n a l l the 14 populations. 

alaicus, a chromosomal sibling species, has 2n = 52, NF = 56. 

originated from the east E. talpinus variant by means of Robertsonian 

fusion. E. alacius was found in the Alai valley and in the Alai 

mountains, Emnus and E. alaicus are characterized by sex 

chromosome Tsomorphism, which is unusual for other Ellobius spp. 

[Abstract from Biological Abstracts] 

No translocation of a sex chromosome on an autosome was 

The length of the X chromosome is 5.7% and 

The 

E. 

It- 
aMalygin, V. M. 1970. The systematics of the subspecies of the comon 066 

field mouse. 

Pochvovedenie 25( 5) : 89-91. 

Vestnik Moskovskogo Universiteta, Seriya V. Biologiya 

The cornon field mouse Microtus arvalis Pallas is a well studied 

species covered in more than 500- The forms which have been 

combined under the heading tkomnon field mouse” constitute a young 

group characterized by an active process of species formation. 

common field mice, then, 

consisting of a group of closely related (sibling) species. 

corrrnon field mice consisting of both the 46 and 54 chromosome varieties 

from 17 geographical locations in USSR were studied. The chromosomal 

makeup of the trans-Caspian field mouse was also studied. 

chromosome type common field mouse showed marked karyotypic differences 

in different portions of its geographic range. The 54 chromosome type 

is, again, completely different. 

The 

as a group, comprise a superspecies 

190 

The 46 

Among the hundreds of field mice caught in the wild, not a single 

sibling species hybrid was found. In order to prove the species 

independence of these forms individuals were hybridized to demonstrate 

the reproductive isolation between the sibl ing species. The hybrids 

have 50 chromosomes (23 chromosomes from 46 chromosome field mice and 

27 chromosomes from 54 chromosome species) and are sterile. 

Chromosomal differences between these sibl inq species is so great that 

in meiosis the conjuqation of homologous chromosomes is completely 

violated leadinq to total sterility in hybrids. 

The diploid complement of M. transcaspicus elaeus Thom. has 54 

chromosomes. In the hybrid males resulting from crossing 54 

chromosomal comnon field mice with transcaucusus mice, spermatogenesis 

progresses almost to the end, but there are no mature spermatozoons and 

the hybrids are sterile. 

analyses attest to the species independence of the transcaucusus field 

mouse. 

The results of karyotypic and hybridologic 

The discovery of sibling species as well as the description of 

several new karyotypic forms of the common field mouse totally alters 

the present concept of the species Microtus arvalis Pallas and its 

intra-species differentiation. The studv remhat because of 

synpatry M. a. duplicatus and M. a. trankaucasicus are without a doubt 

mixed subFpeFies and a radical-syytematic review of‘ these subwecies is - 

indicted. 

It is postulated that the 54-chromosome karyotype is ancestral and 

that the 46-chromosome form derived by centric fusions in the large 

chromosomes and pericentric inversions in the small chromosomes. 

[Abstract by GLW] 
Malygin, V. M. and V. N. Orlov 1974. Range of 4 species of voles 06

(superspecies Microtus arval is) by karyological data. Zool. ZH 

53(4) :616-622. 

Karyotypes of over 400 voles from 8 regions were studied and a 

distribution map of closely related species within the range of the 

superspecies M. arvalis is given. M. subarvalis occurs in eastern 

Europe, the UFalsrmeni a [USSRI. 

Kopetdag Mountains and M. ilaeus is an endemic of Tien-Shan. 

chromosome forms of M. arvmepl ace one another geographically. 

The M. a. forma obscTrus was found in Altai, South-West Siberia, 

UralT Cxucasus, in the basins of the Volga and Don, and the Crimea 

and M. a. forma arvalis in Europe. Ranges of sibling species, M. 

arvaTis-- M. sub-, are sympatric in the eastern Europe, txe 

=andanArmenia. 

were found in 8 regions. 

M. transcaspicus inhabits the 

8 

Contacts between the colonies of these species 

[Abstract from Biological Abstracts] 

Mascarello, J. T., A. D. Stock, and S. Pathak 1974. Conservatism in 06

the arrangement of genetic material in rodents. J. Mamnalogy 

55(4) :695-704. 

The Giemsa banding pattern of the karyotype of Neotoma micropus is 

compared with the banding patterns of seven other romecies that 

are progressively more distantly related to N. micropus. Results of 

comparisons imply that there is a tendency fzr arrangement of genetic 

material on chromosomes to be conserved. Results also lend support to 

systematic inferences based on characters of the glans penis and 

suggest that the type of approach employed here may be useful in 

characterizing phylogenetic relationships between genera, tribes, 

subfami 1 ies, and fami 1 ies of rodents. [Authors' abstract] 

Mascarello, J. T. and J. W. Warner 1974. Chromosome variations in 06

the plains woodrat: A pericentric inversion involving constitutive 

heterochromatin. Experientia 30(1) :90-91 

The chromosomes of the plains woodrat (Neotoma micropus Raird) 

were studies using C- and G-banding, and compared with the 

chromosomes of other members of the qenus Neotoma. G-bands revealed 

distinct homologies between the two largest autosomes in every case. 

The only regions not matching band-for-band were those regions having 

been stained positive for constitutive heterochromatin. C- and 

G-band patterns of the heteromorphic autosomal pair were interpreted 

as clear indications that the large submetacentric and large 

acrocentric chromosomes are homologous, except that one has sustained 

a pericentric inversion in the region containing constitutive 

heterochromatin. 

Neotoma micropus and N. floridana appear to be very closely 

related, both morphologicai-ly and karyotypically, and probably had a 

comnon ancestor. [Abstract by GLW] 
McFee, A. F. and M. W. Banner 1969. Inheritance of chromosome number 070 

in pigs. J. Reprod. Fert. 18:9-14. 

All possible crosses were made between European wild piqs with 

either 36 or 37 chromosomes and domestic swine with 38. The 36 X 36 

cross produced only pigs with 36; the 36 X 37 and 37 X 38 crosses t 

yielded the paren numbers in about equal numbers of pigs. All pigs 

resulting from the 36 X 38 cross had 37, while crossing 37 X 37 gave a 

progeny with 36, 37 or 38 in about 1:Z:l ratio. It is surmised that 

the three unpaired members in the 37-chromosome animal act as a 

trivalent during meiosis with two telocentric chromosomes behavinq as a 

unit. No firm evidence indicated reduced fertility in any of the 

animals nor were any physical changes evident which could be associated 

with different chromosome forms. [Authors' summar.vv'] 

kFee, A. F., M. W. Banner, and J. M. Rary 1966. Variation in 

chromosome number among European wild pigs. Cytogenetics 5:75-81. 

Peripheral leukocyte cultures from 36 European wild pigs have 

shown that 73% of the animals possessed 36 chromosomes while 27% had 

37. Karyotypes of 36-chromosome individuals differed from those of 

domestic swine in that they possessed one pair of submetacentric 

chromosomes not found in domestic pigs and lacked two oairs of the 

domestic's telocentrics. The 37-chromosome animals had one chromosome 

of each of these three pairs but lacked their homologous members. 

Matings between animals with 36 and 38 chromosomes have produced 

fertile offsprinq with 37. A sinqle litter from a 36 X 37 mating 

consisted of three pigs with 36 and three with 37 chromosomes, while 

one from a 37 X 37 mating contained proqeny with 36, 37, or 38 

chromosomes. The odd number of chromosomes in some wild pigs is 

postulated to have resulted from the of domestic breedinq into 

the wild herd. [Authors' abstract] 

Meier. M. N. 1968. ComDlex taxonom 

entry 

c ana 

07

ysis of the species based on 07

the study of cornon voles (genus - Microtus). The Zoo1 ZH 47(6): 

850-859. 

The taxonomic status of 3 forms of comnon voles: Microtus fortis 

Buchner, - M. ungurensis, Kostschenko and M. Mongolicus Radde was 

studied. The species independence of alT the 3 forms is established 

after comol ex methods of invest i qat i on (mor phol oq i c a1 , hybrid i zat i on 

and karyoloqical) . Stable morpholoqical differences, as well as larqe 

differences in the number and form of chromosomes were found between M. 

fortis and M. ungurensis. These forms do not cross due to the above 

citeddifferences. No morphological differences are observed between 

- M. monqolicus and M. arvalis (formerly thought to be subspecies of the 

same speciesj, nevFrtETET-both forms are strictly isolated 

genetically. [Abstract from Bioloqical Abstracts] 

- 
Meier, M N., V. N. Orlov, and E. D. Skhool' 1969. Use of data on 

karyological, phsyiological and cytophysiological analyses for 

isolating a new species of rcdent (Rodentia, Mammalia). Dokl Akad Nauk 

- 

SSSR 188( 6) : 1411-1414. 

The isolation of a new species of vole of the genus Microtus by 

means of karyological, physiological, and cytophysiological criteria 

was established. The form proposed as an independent species is 

Microtus arvalis Pallas. A study of the karotype of this vole showed 

that in the USSR there are 2 distinctly different forms. which are 

designated as forms A and B. 

to the conclusion that forms A and B of the common vole, having 

different karyotypes, differing in heat resistance of the cells, and 

reproductively isolated, can be regarded as independent species desp 

the absence of differences in the morphological characters studied. 

The name M. arvalis Pallas can be left for form A, since its origina 

descriptizn sooner pertain to the 46-chromosome form, and the 

54-chromosome form B can be given the name of some subspecies, for 

example Microtus arvalis Caspicus, if it turns out that this subspec 

belongs wholly to form B or only B is found in the locale from where 

is described. [Abstract from Biological Abstracts] 

A comparison of all the data obtained ed 

te 

es 

it 

07

Nadler, C. F., R. S. Hoffmann, and A. Woolf 1973. G-band patterns as 07

chromosomal markers, and the interpretation of chromosomal evolution in 

wild sheep (Ovis). Experientia 29:117-119. 

Results of G-banding in wild sheep suggest the following 

hypotheses may apply: Chromosomal convergence among populations of 

European and Asiatic sheep implies that they have been derived from a 

common ancestor, possibly in Asia Minor or Europe, in which centric 

fusions have resulted in the present 2n=54 populations. Independently 

and earlier, another ancestral population was isolated somewhere to the 

east, perhaps in middle or central Asia. Centric fusion in that 

population resulted in sheep with a diploid number of 56, eventually 

giving rise to modern arkhar/argali, which in turn was the source for 

amphi-Beringian populations. Ancestral 2n=56 sheep migrated across the 

Bering land bridge in the Pliestocene and became isolated from their 

ancestral populations. Through additional centric fusion, they 

developed into modern Dall and Bighorn sheep. Fusions between specific 

acrocentric chromosomes is suggested as a means by which North American 

2n=54 populations evolved. [Abstract by GLW] 
f 

.- 

Nadler, C. F., R. S. Hoffmann, and A. Woolf 1974. G-band patterns, 07

chromosomal homologies, and evolutionary relationships among wild 

sheep, goats, and auodads (Mammalia: Artiodactyla). Experientia 

30:744-746. 

The tribe Caprini (family Bovidae) contains five genera, among 

Other members of the tribe, particularly the genus 

Analyses suggest that the diploid 

which the wild sheep (Ovis) and goats (Capra) are most closely related. 

Chromosome analyses omee genera demonstrated a comnon fundamental 

nunber (NF=60). 

Ovis, have divergent 2n numbers. 

Wer of 60 is ancestral, having evolved by Robertsonian translocation 

into forms with lower diploid numbers. [Abstract by GLW] 

Nadler, C. F., K. V. Korobitsyna, R. S. Hoffmann, and N. N. Vorontsov 07

1973. Cytogenetic differentiation, geographic distribution, and 

domestication of palearctic sheep. (Ovis) Z. Sauqetierkunde 

38 ~109-125. 

Old World sheep may be divided into four groups, with different 

diploid chromosome numbers and morpholoqical characters. There are, 

from west to east, mouflon (2n=54) in the Mediterranean reqion and the 

Near East; urial (2n=58) in Middle Asia from eastern Iran to 

Tadzhikistan and probably West Pakistan; arkhar/argal i (2n=56) in 

Middle and Central Asia; and snow sheep (2n=?) (subgenus Pachyceros) in 

eastern Siberia. The systematic status of sheep from Eurooe to Central 

Asia (subgenus Ovis) remains uncertain; in this broad area wild sheep 

are d istr ibuteflinterconnected ri nqs many.of which have been 

repeatedly broken and rejoined during the Pleistocene, leading to 

differentiation of isolates and subsequent introqression. Domestic 

breeds of sheep so far studied all appear to be derived from a mouflon 

(2n=54) ancestor in the Near East. [Authors' summary] 

,ill 

4- 

pl 

Nadler, C. F., D. M. Lay, and J. D. Hassinqer 1971. Cytogenetic 0

analyses of wild sheep populations in Northern Iran. Cytogenetics 

10~137-152. 

Seven populations of wild sheep (Ovis ammon) were studied in 

northern Iran. Analysis of 34 individx -strated that the 

eastern populations possess a 2n=58, including 1 pair of metacentric 

and 27 pairs of acrocentric autosomes. The 2n for the three western 

localities is 54, characterized by 3 pairs of metacentric and 23 

pairs of acrocentric autosomes. In two intermediate localities 

animals with the following diploid nunbers were collected:' 2n=54, 

identical to western local ity animals; 2n=55, containing five 

metacentric autosomes; 2n=56, containinq four metacentric autosomes; 

2n=57, with three metacentric autosomes; 2n=58, identical to the 

eastern two localities. 

acrocentric and a small biarmed chromosome, respectively, in every 

specimen. The taxonomy of Iranian wild sheep and several 

evolutionary factors are considered in the liqht of these chromosomal 

data. The nonconformity of currently accepted theories on the 

origins of domestic sheep with the karyoloqical data now available 

for wild palaearctic and domestic sheep is discussed. [Authors' 

ab s t r act ] 

The X and Y chromosomes are a large 
Nelson-Rees, W. A., A. J. Kniazeff, R. J. Baker, and J. L. Patton 

1968. Intraspecific chromosome variation in the bat, Macrotus 

waterhousi i Gray. J. Mamalogy 49(4) :706-712. 

07

The karyotypes of two subspecies of bat, Macrotus waterhousii Gray 

(Phyllostomatidae), have been studied and compared. Specimens of M. 

waterhousii mexicanus from Morelos and Guerrero, Mexico, as well as 

a karyotype with a diploid number of 46 including a telocentric 

Y-chromosome in the male complement. Another sample of M. waterhousii 

California, had a diploid number of 40. Of these samples, the Arizona 

specimens also revealed a telocentric Y-chromosome and in addition a 

secondary constriction on the supposed X-chromosome, whereas the 

California sample was characterized by a pronounced achromatic region 

near the centromere on one of the large pairs of telocentric 

chromosomes and the existance of a meta or sub-metacentric Y-chromosome 

in the male. This appears to be the first major karyot.ypic variation 

of bats reported at the subspecific leve?. [Authors' abstract] 

specimens in one sample of M. waterhousii californicus from Sonora, had 

californicus from Sonora, as well as samples from Arizona and 

Nevo, E. and H. Cleve 1978. Genetic differentiation during 

speciation. Nature 275:125-126. 

Karyotypes of the subterranean mama1 Spa1 ax ehrenberqi were 

studied in liqht of genetic differentiation. Results support the 

hypothesis that habitat specialists (such as the mole rat, S. 

ehrenbergi) display significantly lower qenetic variation tEan do 

habitat generalists. Concerning speciation theory, the S. ehrenberqi 

complex represents a remarkable case of speciation with relatively 

few genomic changes. [Abstract by GLW] 

07

Pathak, S., T. C. Hsu, and F. E. Arrighi 1973. Chromosomes of O

Peromyscus (Rodentia, Cricetidae) IV. The role of heterochromatin in 

karyotypic evolution. Cytogenet. Cell Genet. 12:315-326. 

All species in the genus Permyscus possess a diploid number of 

48, but the number of total chromosome arms varies from 56 (e.g., P. 

crinitus) to 96 (e.g.,I. eremicus). Data are presented, usinq these 

two extreme cases, to illustrate that all short arms of biarmed 

chromosomes are made of constitutive heterochromatin. 

preparations revealed that the long arms (euchromatin) of the two 

species are essentially the same. Since constitutive heterochromatin 

contains few, if any, stuctural genes, the two species, and presumably 

other species as well, do not differ drastically in their information 

content, but differ in the amount of chromatin material, including DNA. 

The results indicate that in studying karyotypic evolution, C-banding 

and G-banding (or Q-bandinq) are essential tools without which 

erroneous conclusions may be reached. [Authors' abstract] 

G-banding 
Patton, J. L. 1972. Patterns of qeographic variation in karyotype 

in the pocket gopher, Thomomys bottae (Eydoux and Gervais). 

Evolution 26:574-586. 

_L 

Variation in the mitotic chromosome complement is greater within 

the pocket gopher, Thomomys bottae, than any other recorded species 

of mammal. This variation involves almost exclusively 

populations are polytypic with over 40 different population 

karyotypes having been recorded, but a few cases of intrapopulation 

polymorphism are known. The variation is most extensive within and 

between populations in the Basin and Range Province of the Southwest 

whereas most populations in coastal and central Cal ifornia are 

identical, or nearly so. The extensive karyotypic variability is not 

a function of single causative aqents, but rather appears to be 

dependent upon a suite of highly interrelated selective factors. 

Historical biogeographic events which provided the means for 

population isolation under which chromosomal divergence can occur are 

detailed with selected examples. These events in combination with 

biological properties of gophers, particularly those centered around 

their fossorial habitus, are considered largely responsible for the 

karyotypic diversity. 

- non-Robertsonian morphological changes and not diploid number. Most 

II 

There is no evidence that this diversity has contributed to 

reproductive isolation between populations of T. bottae. Rather, it 

appears to have led to increased population adTptation by modifying 

linkage qroups via redistribution of the genome. The chiasmas 

localization found in the species aids in tightening linkage groups 

and reduces genetic Variability resulting from segregation and 

recombination, while at the same time preserves qenic heterozyqosity. 

Hence, optimal gene combinations can evolve to meet the highly 

localized selective parameters under which each population exists. 

This can be done without sacrificing genetic heterozygosity or 

forcinq reproductive isolation. [Author's summary] 

Patton, J.L. and R. E. Dingman 1968. Chromosome studies of pocket 

gophers, genus Thomomys. I. The specific status of Thomomys umbrinus 

(Richardson) in Arizona. J. Mammalogy 49(1) :1-13. 

The complexities of morphology and, hence, taxonomy of pocket 

qophers (genus Thomomys) in southern Arizona are reflected by extreme 

interpopulation chromosomal variation in both T. bottae (2n=76) and 

- T. umbrinus (2n=78). 

morphological types of chromosomes for nearly each population 

karyotype. The known range of variation in either species is less 

than the amount of difference between the two. A somewhat strict 

ecological separation exists between T. bottae and T. umbrinus in 

areas of sympatry or near sympatry, withtheormerpreferrinq the 

more friable soils of the valley floors and mountain tops and the 

latter confined to the indurate soils of the oak zones at 

intermediate elevations. Chromosomal and ecoloqical concordance 

support the interpretation that T. bottae and T. umbrinus are 

distinct species. 

one localty of sympatric contact, however, is known. [Authors' 

abstract) 

The variation consists of dmng numbers of 

Limited hybrixization between the two species at 

08 1 

082 
Raman, R. and T. Sharma 1972. Similarity in karyotypes of Rattus 083 

rattus with 38 chromosomes from India and other parts of the world. 

Experl’entia 28(11) :1375-1377. 

The chromosomes of 12 male and 12 female specimens of R. rattus, 

collected from various locations in India, possessed a diploid 

complement of 38. 

others; however, their karyotypes did not differ in diploid number. 

The Indian populations of R. rattus supDort a concept of chromosomal 

convergence as a means of evom. The Indian rats are probably 

derived from a European ancestor having a diploid number of 42. 

[Abstract by GLW] 

Two of the rats had darker underparts than the 

Rausch, R. L. and V. R. Rausch 1972. Observations on chromosomes of 084 

Dicrostonyx torquatus stevensoni Nelson, and chromosomal diversity in 

varying leminqs. 2. Sauqetierkunde 37:372-384. 

The cytogenetic characteristics of the varying lemning, 

Dicrostonyx torquatus stevensoni, (2n=34), were investigated, and 

diploid chromosomal numbers were reported for four other nominal 

subspecies (exsul, nelsoni, richardsoni, and rubricatus) of the 

torquatus-group in North America. The diploid complements ranqed from 

30 to 44 chromosomes, and the fundamental number from 50 to 55. 

Chromosomal ploynorDhism was observed i n a l l forms. In cross-breeding 

experiments, the matinq of F1 progeny was not productive. The 

findings support the zoogeographic concept that populations of 

Dicrostonyx became fraqmented or displaced southward during Wurm time, 

with re1 ict stocks persisting in unglaciated refugia or periglacial 

tundra. Speciation in the isolates led to chromosomal evolution, with 

the result that populations spreading from refugia in post-glacial time 

are reproductively isolated. The torquatus-group in North Pmerican 

appears to be a super-species. [Authors’ summary] 
1 Rausch, R. L. and V. R. Rausch 1975. Relationships of the red-backed 085 

vole, Clethrionomys rutilus (Pallas), in North America: Karyotypes of 

the subspecies dawsomalbiventer. Syst. Zoo1 . 24(2) :163-170. 

Cytogenetic comparisons were made of northern red-backed voles of 

two subspecies, Clethrionom s rutilus dawsoni Merriam and C. r. 

albiventer to the F18 generation. 

karyotypes were indistinquishable for all animals. with the exceotion 

albiventer Hall + and Gi more, asas of intergrades of JawToni X 

The diploid nmber was 56, and 

of some individuals of Cl r. albiventer (with 2n=55) which were 

heterozygous for a chroZos?imal polynorphism evidently resulting from a 

Robertsonian rearranqement. All species of Clethrionomys for which 

karyotypes are known have 2n=56, but two groups are distinguishable by 

the form of the Y-chromosome: in C. rufocanus (Sundevall), C. gapperi 

(Viqors), and C. occidental is (MeFri'vY is acrocentr%,whereas 

in C. qlareolu? (Schreber) and C. rutilus, it is metacentric. Thus, 

the-karyotypes of dawsoni and aTbiventer are indistinquishable from 

that of the Eurasim. mikado (Thomas), and the different forms of 

the Y-chromosome separatz C.rutl'lus and C. gapperi in North Prnerica. 

These findings further supoormoncepT tr rutilus is a 

hol arct ic species. (Red-backed vole; karyotypes.) ms' abstract] 

Reig, 0. A., J. V. Pincheira, A. 0. Spotorno, and P. Waller 1972. 

New evidence of a 38-chromosome karyotype in South American 

populations of roof rat Rattus rattus L.(Rodentia, Muridae). 

Experientia 28(2) :225-227. 

In two populations of Rattus rattus, one from Chile and the 

other from Venezuela, karyotypes with chromosome complements of 2n=38 

were observed in all of the animals studied. Rattus rattus is shown 

to be highly variable in chromosome nunber and structure, showing 

both geoqraphic, and possibly subspecific, multiformity and 

polynorphism in karyotypes. 

The analyses of chromosomes of the South American R. rattus 

populations suggests that they were derived by means of-Robertsoni an 

translocation from a European ancestor having a diploid number of 

42. [Abstract by GLW] 

Robinson, T. J. and J. D. Skinner 1976. A karyological survey of 

springbok subspecies. S. African J. Science 72:147-148. 

Studies of subspecies of springbok (A. marsupial is) indicate 

that the subspecies are characterized by a monomorpic karyotype 

throughout their distribution in southern Africa, and their diploid 

chromosome nunber 2n=56 (54 autosomes and 2 sex chromosomes). 

[Abstract by GLW] 

086 

087 
Schmidly, D. J. and G. L. Schroeter 1974. Karyotypic variation in 088 

Peromyscus boy1 ii (Rodentia: Cricetidae) from Mexico and corresponding 

taxonomic implications. Syst. Zool. 23:333-342. 

Nine different karyotypes are evident amonq samples of six 

subspecies of Permyscus bo lii (P. b. rowleyi, P. b. spicilegus, -- P. b. 

simulus, P. b. levipes, P. + ambTquTs,-ZdT bT bFatae). All 

spec imens-have mid-nunber of 48; the p'Fincipmferences are in 

the nunbers of large and medium-sized biarmed autosomes which range 

from 2-10. Three chromosome races are recognizable in P. boylii; they 

are distributed parapatrical ly with respect to one anotFer, and present 

evidence suggests that limited gene exchanqe occurs among them. The 

karyotype for topotypes of P. b. simulus from Nayarit, Mexico, is 

markedly different from all-otTierwof P. boylii, sugqestinq that 

this taxon is a distinct species. 

non-Robertsonian type is seen in certain populations of P. boylii and 

may be the result of intergradation between the differenT chromosome 

races. The polymorphism involves a variable number of large biarmed 

autosomes with certain individuals in a population possessins a 

heteromorphic pair of autosomes. - P. evides, both members of the boylii species group, are also described 

for the first time. [Authors' abstract] 

- 

Chromosomal-orphism of a 

The karyot-ypes of - P. oaxacensis and 

Schroder, J., J. Antoni, and W. Van der Loo 1978. Comparison of the 08

karyotypes in the jack rabbit (Lepus californicus deserticola) and 

the European hare (Lepus europam Hereditas 89:134-135. 

The karyotypes of Lepus europeaus (European Hare) and L. 

cal ifornicus deserticolxsubspecies of herican jack rab6it) were 

compared to determine correlation between qeoqraphic isolation and 

karyotypical divergence. Both species possessed a diploid number of 

48, and differed mainly in chromosome morphology. The study supports 

the argument that chromosomal speciation is encouraqed by qeographic 

proximity, and that geographically isolated species have no need to 

develop divergent karyotypes. [Abstract by GLW] 
Schroder, J:, H. Suomalainen, W. Van Der Loo, and E. Schroder 1978. 

Karyotypes in lymphocytes of two strains of rabbit, and two species 

of hare. Hereditas 88:183-188. 

G-banding of metaphase chromosomes in cultured lymphocytes was 

used to compare the karyotypes of two strains of domestic rabbit. 

cuniculus (NZW and Dutch rabbit) and two species of hare 

and Lepus t imidus) . 

The NZW and the Dutch rabbit had identical karyotypes with 44 

chromosomes, as did Lepus europaeus and Lepus timidus with 48 

chromosomes. 

However, a1 1 individual rabbits and hares showed chromosomal 

rearrangments (e.g., deletions and duplications) in 30-50s of all 

lymphocytes studied. 

discussed. [Authors' abstract] 

The significance of the findings are 

Schroder, 3. and W. Van Der Loo 1979. Comparison of karyotypes in 

three species of rabbit: Or ctolagus cuniculus, Sylvilagus 

nuttallii, and - S. Idahoensis + Here itas 91:27-30. 

The karyotypes in three species of rabbit (Oryctola us 

cuniculus, Sylvilagus nuttallii and S. idahoensis T--e were compared. 

The karyotype of Oryctolagus cuniculus was used as a reference. The 

chromosome nunber in Oryctola us cuniculus and Sylvilagus idahoensis 

is 44, and in S. nuttd Four chromosome pairs in Sylvilagus 

idahoensis and-three in S. nuttallii are clearly different from those 

in Oryctolaqus cuniculus: but minor differences can also be found in 

other chromosomes in these species. A very striking similarity in 

the karyotypes of Oryctolagus cuniculus and Sylvil aqus nuttall i i is 

that they have identical metacentric chromosomes 1, whereas in S. 

idahoensis this chromosome is represented by two separate chromosome 

pairs. In spite of this, the karyotypes in the three species have 

diverged from each other about equally. [Authors' abstract] 

090 

091 

Selander, R. K.. 0. W. Kaufman. R. J. Baker. and S. L. Williams 1974. 092 

Genic and chromosomal differentiation in pocket gophers of the Geomys 

bursarius group. Evolution 28:557-564. 

The processes of karyotypic and genic differentiation have 

proceeded independently in the evolution of pocket gophers of the 

Geomys bursarius group. In the peripheral relict G. tropicalis, a 

reduction in 2N chromosome number from-70 to 38 was not accompanied 

by an unusual degree of allelic substitution at structural gene loci. 

The possibility that a reduction in number of linkage groups and an 

apparent loss of variability at structural gene loci in G. tropicalis 

represent adaptation to an unusally uniform environment Ts 

discussed. [Authors' summary] 
Sen, S. and T. Sharma 1979. Sparse distribution of constitutive 

heterochromatin and its variation in two species of monqooses 

(Carnivora) with exact G-band homology. Genet ica 50( 3) :221-226. 

The diploid chromosome number in two species of mongooses viz., 

Herpestes auropunctatus and H. edwardsi is 35 in males and 36 in 

females, since the Y chromosome in the males of these species is 

translocated onto an autosome. C-banding of these karyotypes reveals 

that (1) they contain an extremely low amount of C-band positive 

constitutive heterochromatin (C-heterochromatin); (2) H. edwardsi 

possesses an increased amount of C-heterochromatin comDared to H. 

auropunctatus which is localized as prominent arms on two pairs-of 

chromosomes; (3) the translocated Y chromosome in both species is not 

stained differentially by this technique. Hoechst 33258' flourescence 

does not distinquish any differentially briqht heterochromatic reqion 

in either of the two genomes. G-bandinq patterns in the chromosomes of 

these two species show remarkable similarity. [Authors' abstract1 

Sharma, T. and I. K. Gadi 1977. Constitutive heterochromatin 

variation in two species of rattus with apparently similar karyotypes. 

Genetica 47(1) 77-80. 

Rattus blanfordi and R. cutchicus medius both have a chromosome 

complement of 2n = 36 and all chromosomes except the submetacentric Y 

of R. blanfordi are acrocentric. The apparently similar karyotypes of 

the-two species, however, show variations in the nature and quantity of 

C-band-posi t ive constitutive heterochromatin (C-heterochromat in) as 

revealed by C- and G-banding and Hoechst 33258 fluorescence. R. 

bl anfordi with larqe-sized X and Y chromosomes and consDicuousTy larqer 

centromeric heterochromatin in all the autosomes as compared to that of 

- R. cutchicus medius has much more C-heterochromatin in its qenome than 

the latter. The variation in the quantity of C-heterochromatin has 

been accomplished without alterinq the morphology of the acrocentric 

chromosomes unlike other mammals in which variations have been reported 

to result qenerally in the addition or deletion of a totally 

heterochrornat ic second arm. [Authors' abstract] 

093 

094 

Sharma, T. and G. S. Gara 1975. Constitutive heterochromatin and 095 

karyotype variation in Indian pygmy mouse, -- Mus dunni. Genet. Res. 25: 

189-191. 

The Indian pyyny mouse, Mus dunni, exhibits qreat variation 

[between separate geographic regimin the number of chromosome arms 

while its diploid number of chromosomes remains constant. The 

variation seems to be due to addition or deletion of C-band Dositive 

constitutive heterochromatin in the short arms of autosomes. [Authors' 

summary] 

- 
Sharma, T. and R. Raman 1973. Variation of constitutive 

heterochromatin in the sex chromosomes of the rodent Randicota 

benqalensis bengalensis (Gray). Chromosoma 41:75-84. 

Bandicota benqalensis ben alensis (Gray) trapped from different 

and morphology of sex chromosomes. Three types of X's were found; A) 

simple acrocentric, B) composite subtelocentric and C) composite 

submetacentric X with their relative sizes 5.9%, 7.5% and 9.6% of the 

genome respectively . The autosomes remained una1 tered . It was shown 

that this variation in the size of sex chromosomes was caused by 

deletion of constitutive heterochromatin. The Y chromosome was also 

found to be variable. Usually a large X was combined with a large Y. 

The preponderance of homozygotes for each type of X chromosome in 

populations, suggested that probable role of sex chromosomes 

heterochromatin in spec i at ion. [Authors abstract] 

localities of India and Nepa + ex 1 ited a marked variation in the size 

096 

Sinqh, R. P. and D. B. McMillan 1966. Karyotypes of three subspecies 097 

of Peromyscus. J. Mammalogy 47(2) :261-266. 

Karyot-ypes of Peromyscus maniculatus gracilis Le Conte, Peromyscus 

maniculatus bairdi i Hoy and Kemd Peromyscus leucopus 

noveboracens-cher) from southern Ontario were studied to 

determine whether or not identification of these animals can be made on 

a cytological basis. 

plates of bone marrow and newborn splenic tissue confirm the diploid 

nunber of 48 in each case as given by Makino (1951) and are 

sufficiently distinct to permit positive identification of each 

subspecies. [Authors' abstract] 

The karyotypes as determined from metaphase 

Sokolov, V. E., V. N. Orlov, G. A. Chudinovskaya, and A. A. Danilkin 098 

1978. 

Capreolus capreolus capreolus L. and C. c. pygargus Pall. Zool. ZH 

Chromosomal differences between two subspecies of roe deer, 

-- 

39(7)09 - 1 1 1 2. 

Differences in chromosomes are described between roe deer from 2 

populations, C. c. capreolus from the Bryansk District and C. c. 

pyqarctus from-thT South Ural [USSR]. The diploid number is-70-for C. - c. capreolus and 74 for C. c. Dygarqus. Unlike the former the 

karm the latter xas-2 pairs of point-like chromosomes. These 

differences may be used in studying the evolution of C. capreolus. 

[Abstract from Biological Abstracts] 

- 

- 
Soldatovic, B., R. D.julic, I. Savic, and D. Rimsa 1969. Chromosomes 09

of two species of the genus Apodemus (A. aqrarius and A. mystacinus -- 

Mammalia, Rodentia) from Yuqoslavia. Arhiv. Riol. Sci. 21(1-4) :27-32. 

The karyotypes of Apodemus agrarius kahmanni and A. mystacinus 

epimelas are described. The phenomenon of two different karyotvpes in 

%he species A. aqrarius from Yugoslavia (form from Bosnia with N. F. = 

54, while a17 other forms from Yugoslavia have N.F. = 56) indicates 

that this is also a case of chromosome polymorphism, probably a 

consequence of Dericentric inversion. It may be hypothesized that 

pericentric inversion of a metacentric chromosome gave rise to a pair 

of small acrocentric chromosomes. This would mean that the Bosnian 

type had the transformed karyotype. 

transformation is possible, so it is difficult to say which is the 

primary form. More detailed phenotype analysis will show whether the 

- A.aqrarius from Bosnia represents a separate species or subspecies or 

whem chromosome polymorphism should be ascribed only to lability 

of the karyotype resulting in a balanced translocation. [Abstract by 

GLW] 

On the other hand, the converse 

Sparkes, R. S. and D. T. Arakaki 1966. Intrasubspecific and 10

intersubspecific chromosomal polymorphism in Peromyscus maniculatus 

(deer mouse). Cytogenetics 5:411-418. 

Peromyscus, maniculatus rubidus, P. m. gracilis and P. m. bairdii have 

demonstrated an intra- and intersGbspecif ic chromoso?iialpomism in 

which the karyotypic patterns vary amonq individuals of the same and 

different subspecies while maintaining a diploid number of 48 and a 

constant karyotype in each animal. Present information suggests that 

pericentric inversion seems the best explanation for this 

polymorphism. [Authors’ abstract] 

Cytogenetic studies of three subspecies of the deer mouse, 

Sparkes, R. S. and D. T. Arakaki 1971. Chromosome polymorphism in 10

interbred subspecies of Peromyscus maniculatus (deer mouse). Can. J. 

Genet. Cytol . 13:277-282. 

Karyotype analyses of eight animals from an interbreeding colony 

of three subspecies of Peromyscus maniculatus (P. m. gambelli, P. m. 

rabidus, and P. m. sonoriensis) demonstrated a chr%mosomal 

pblphism,prZbably due to pericentric inversions involvinq at 

least seven chromosomes. 

in the cytogenetic taxonomy of these animals, and may be related to 

the wideswead distribution of deer mice in North Pmerica and their 

apDarent adaptabil ity to many different environments. 

abstract] 

-- 

This polymorphism may require consideration 

[Authors’ 
w 

I- 

- 

Stock, A. D. 1975. Chromosome banding pattern homoloqy and its 102 

phylogenetic imp1 ications in the bat genera Carol 1 ia and Choeroniscus. 

Cytogenetics and Cell Genetics, 14:34-41. 

The chromosome banding patterns of the mitotic chromosomes of 

The G-band patterns of the Carollia species were similar but the 

The 

three species of Carol1 ia and Choeroniscus intermedius was compared. 

C-band (heterochromatin) pattern of C. Castanea (Peru) differed. 

X-autosome translocation in C. perspycillata and C. brevicauda was 

compared to the untranslocatFd homologues in C. c'stanea (Peru). 

from that of the Carollia species and the placement of these qenera 

into different subfamil ies was supported. 

- 

The G- and C-banding patterns of Choeroniscus intermedius differed 

[Author's abstract] 

Thaeler, C. S. Jr. 1968. Karyotypes of sixteen populations of the 10 3 + Thomom s tal poides complex of pocket gophers (Rodentia - Geomyidae) . hromosoma 25:172-183. 

Chromosome material of mitotic bone marrow cells from 59 

individuals representing 16 populations of Thomomvs talpoides was 

studied. 

northern New Mexico and northern Arizona. A limited amount of 

intrapopulation variation in form but not number of chromosomes was 

found. Eight distinct karyotypes were observed. These can be 

characterized by the following diploid numbers and fundamental numbers 

of chromosomes: 40( 70), 44( 70), 56(70), 60( 70), 58( 76), 48( 78), 56( 78) 

and 46(82). 

interpotwlation variation was noted. In several instances karyotype 

differences coincided with populations that co-exist without 

interbreeding. [Author's abstract] 

These populations were found in southernWyoming, Colorado, 

Among the six populations with 48 chromosomes some 

Thaeler, C. S. Jr. 1974. Four contacts betwen ranges of different 104 

chromosome forms of the Thomomys talpoides complex (Rodentia: 

Geomyidae) . System. Zoom) :343-354. 

Contacts between the ranges of four pairs of different 

chromosome forms of pocket gophers of the Thomomys talpoides complex 

were studied in an effort to gain insight into the relationship 

between karyotype differences and level of evolutionary diverqence. 

All contacts are located in the mountainous region of western 

Colorado. Three different results were found. At one contact no 

evidence of hybridization between the two chromosome forms was 

detected by an examination of karyotypes. At two other contacts 

limited hybridization occurs between the chromosome forms. The 

fourth contact studied is a broad zone of intergradation (20 miles or 

mor,e in width) in which more or less unrestricted interbreeding 

occurs. A discussion of each contact area is presented and I 

conclude that frequently, though not always, karyotype differences in 

pocket gophers are indicative of species-level differences. 

ab s t r act] 

[Author's 
Tikhonov, V. N. and A. I. Troshina 1974. Identification of 105 

chromosomes and their aberrations in karyotypes of subspecies of Sus 

scrofa L. by differential staininq. Doklady Biol. Science 214:4-7. 

Investigations of the karyotype of four widely separated 

populations of wild boars showed that the diploid set of chromosomes 

varies from 36 to 38. Many boars from a central Asian population (24 

animals) had 36 chromosomes whereas 2n=37 was found in smaller numbers 

in that population. No individuals having 2n=36 were found among boars 

of Far-Eastern, Transcaucasian, and western European populations. All 

homologous chromosomes of the karyotype could be identified by 

differential staininq. [Abstract by GLW] 

Valdez, R., C. F. Nadler, and T. 0. Bunch 1978. Evolution of wild 106 

sheep in Iran. Evolution 32:56-72. 

Eight populations of wild sheep (Ovis) throughout Iran were 

sampled and analyzed for chromosome pattern, horn configuration, pelage 

characteristics, fetal rates, and serum transferrin t.ypes. Individual 

populations from northwestern Iran, the Zagros Mountains, and Laristan 

uniformly exhibited a 2" of 54, and sheep from northeastern Iran 

displayed a 2N of 58. 

ranginq from 54-58 and were considered hybrids, as were sheep from the 

Kavir desert and the Kerman region, which had 2"s of 54 and 55. 

- 

Sheep of the Central Alborz possessed 2"s 

Examination of preqnant ewes from the eastern reqion of the 

Central Alborz hybrid revealed 2"s of 56, 57 and 58. Fetal rates were 

similar to parental types with 2"s of 54 and 58. 

chromosomal, and transferrin data suggest that Iranian sheep are 

classified most correctly as semispecies, or alternatively as 

subspecies, because they readily hybridize. 

Morphological, 

The Kavir population probably is derived from Central Alborz 

individuals which became isolated in the Kavir desert. The Esfahan and 

Laristan populations also may have been derived from isolated hybrid 

populations although a pure Armenian or urial origin is possible also. 

AI 1 these popul at ions are characterized by homonynous horns, black 

ruffs and white saddle patches. 

A tall vegetation barrier is h-ypothesized to have oriqinallv 

separated Armenian and urial populations in northern Iran. When this 

barrier disappeared due to drier climatic conditions, these populations 

met and hybridized. Populations in southern Iran also became isolated 

by vegetational barriers. The diverse phenotypic variability of 

Iranian wild sheep probably evolved within the last 15,000 years 

through hybridization coupled with founder events. [Authors' summary] 
Venegas, W. 1974. Karyotypic variation in Ph,yllotis micropus 

Waterhouse (Rodentia, Cricetidae). Bol. SOC. Biol. Concepcion 

48 ~69-76 

micropus 107 

The karyologic results obtained from the study of 5 populations 

of P. micropus micropus are presented. 

wit5 Sherman traps in the provinces of Nuble, Malleco and Aysen 

[Chile]. Females and males (14) were analyzed cytogenetically using 

routine techniques and bone marrow cultures. A diploid number of 2n = 

32 chromosomes with a N.F. [chromosome arm number] = 34 was found in 

the specimens of 2 populations from the province of Nuble, while a 

diploid nunber of 2n = 34 chromosomes with a N.F. = 36 was found in 

specimens of one population from the province of Malleco and 2 

populations from the province of Aysen. 

karyotypic variability found may indicate a speciation process. 

[Abstract from Biological Abstracts] 

The specimens were captured 

The intrasubspecif ic 

Warner, J. W. 1976. Chromosomal variation in the plains woodrat: 

Geographical distribution of three chromosomal morphs. Evolution 

30( 3) : 592-598. 

Four hundred and twenty-eight specimens of Neotoma micropus from 

throughout the species ranqe were karyotypical lymed. There was 

noticeable variation in the occurrence of the three morphs within 

various populations. 

in a state of equilibrium in each population. Statistical analyses of 

cranial morphometrics revealed little significant difference amonq the 

three morphs based on the variables used. There is evidence that this 

system conforms to White's model for stasipatric speciation. Also one 

can speculate that it may eventually fit Rrown's model for centrifugal 

speciation. [Author's smary] 

However, chi-square tests indicate that they were 

Wentworth, F. A., D. A. Sutton 1969. Chromosomes of the townsend 

pocket gopher, Thomomys townsendi i. Southwestern Natural ist 

Animals were collected, subjected to in vivo colchicine 

14( 2) ~157-161. 

treatment and red bone marrow cells were stained with giemsa. 

Photomicrographs and karyoqrams were prepared for a1 1 seven 

subspecies of the Townsend pocket gopher, Thomomys townsendii. 

diploid chromosome number of 76 was found for six subspecies. T. t. 

A 

-- 

bachmani, T. t. elkoensis, T. t. nevadensis, T. t. ow-yhensis, T. t., 

relictus azd T. t. townsendii. 

similis, has a 2s chromosome number of 40 which, alonq wTth-other 

unique morphological traits, indicates either confusion with the 

Thomomys talooides complex or consideration of taxonomic revision to 

Thomomys similis. [Authors' abstract] 

T. - -- The seventh s<bsFecies, T. t. 

108 

109 
White, 1. J. D. 1978. Chain processes in chromosomal speciation. 110 

Syst. 2001. 27( 3) ~285-298. 

In many qroups of animals of restricted vagility very closely 

related species not only differ in karyotype, but exhibit very 

extensive differences, due to the establishment of "chains" of 

several or many structural chromosomal changes. These cases which 

until recently appeared anomalous or inexplicable, can now be used to 

interpret a basic feature of the "stasipatric" mode of speciation 

(parapatric speciation of some authors). It is suggested that an 

important reason for the establishment of these chromosomal 

rearranqements is their role in protectinq coadapted gene complexes 

("area effects") from disruption by introgression from neighboring 

populations. A model, based on the mouse (Mus musculus) populations 

of Italy and Switzerland studied by E. CapaG, A. Gropp, and their 

collaborators, is proposed, it re1 ies on sequential establ ishment of 

chromosomal rearrangements, each within the ranqe of the previous one. 

By this means the genetic isolation of the area population is 

progressively reinforced. [Author's abstract] 

Wroblewski, R. and D. Dziekanowska 1968. Karyology of primates. 11

Prezegl . Zoo1 . 12( 2) :127-140. 

A review of chromosome numbers and kar.vot.ype data is qiven for 116 

spp. and subspecies of Primates. Analysis of chromosome complements 

indicate that polyploidy did not play any role in the karyological 

differentiation of this group. 

fusions and other chromosomal arranqements are considered as possible 

factors in evolution of Primate karyotypes. A significance of 

karyoloqical studies for taxonomic and phyloqenetic cosiderations is 

discussed too. [Abstract from Biological Abstracts] 

The Robertsonian type of centric 

Wurster, D.H. and N.B. Atkin 1972. Muntjac chromosomes: A new 11

karyotype for Muntiacus muntjak. Experientia 28(8) :972-973. 

There are eight listed subspecies of the Indian muntjac (Muntiacus 

The diploid number of chromosomes in M. muntjak vaqinalis is 

Tissue culture preparations of a female M. muntjak 

A diploid number of 9 would be expected in the male of 

muntjak). 

7 in the male and 6 in the female; both sexes carry an X autosome 

translocation. 

muntjak show a diploid number of 8 with an X to autosometranslbcation. 

this subwecies. 

The kar-votype differences between theses subspecies are pronounced 

enough that one would predict synaptic incompatibility in meiotic 

division of a h-hrid offsorinq, thus conferrinq sterility on this 

individual. It is possible that a taxonomic revision should be 

considered and that either subspecies muntjak or subspecies vaqinal is 

should receive full species status. Possible evolutionary siqnificance 

with other species in the same genus is also discussed. [Abstract by 

GLW) 
<- 

IWurster, 0. H. and K. Benirschke, 1968. Chromosome studies in the 

superfamily Bovoidea. Chromosoma 25:152-171. 

113 

The chromosome morpholoqy of about 50 species of Bovoidea has 

now been investigated. 

60 amonq these species, the fundamental number(NF) varies only (with 

but three exceptions) from 58 to 62. This indicates an almost 

exclusive use of the Robertsonian fusion mechanism of karyot.ype 

evolution in this qrouo of species which represent 30 different 

genera. All known cytogenetic information on the Bovoidea has been 

sunmarized and a complete bibliography is presented for each species. 

Karyotypes and data on a number of previously unstudied sDecies are 

presented. [Authors' abstract] 

Although the diploid number varies from 30 to 

Yosida, T. H. 1977. Frequencies of chromosome pol.ymorphism in pairs 114 

no. 1, 9, and 13 in three geographical variants of black rats, Rattus 

rattus. Chromosoma 60:391-398. 

Frequencies of the acrocentric and subtelocentric polymorphism 

in pairs no. 1, 9 and 13 chromosomes have been examined in 358 black 

rats, Rattus rattus, distributed over several countries of Asia, 

Austramd United States. The black rats are divided into three 

geographical types by the different chromosome numbers, such as Asian 

(2n=42), Ceylon (2n=40) and Oceanian types (2n=38). Pairs no. 13 

polymorphism was found widely in these all types, but the pair no. 1 

and 9 polymorphisms were found in on1.y Asian type black rats. 

Asian type rats, however, those distributed in northern India and 

Pakistan showed always the subtelocentric pairs no. 1 and 9 like as 

those in Ceylon and Oceanian type black rats. This finding supports 

that the Ceylon and Oceanian type rats have developed in India or 

Pakistan from the Asian t-rpe. The present study also suggests that 

inversion of the pair no. 13 could have occurred in earlier period 

than those of the pairs no. 1 and 9. 

In the 

[Author's abstract] 
Yosida, T. H. and K. Pmano 1965. Autosomal polynorohism in laboratory 115 

bred and wild Norway rats, Rattus norveqicus, found in Misima. 

Chromosoma 16:658-667. 

Polymorphism in chromosome pair no. 3 of Rattus norveqicus was 

'found in laboratory strains and wild rats. Sowthe animals had a 

subtelocentric pair no. 3, while others had a telocentric pair no. 3. 

14 inbred strains were classified into two types concerning the pair 

no. 3. 

ACI-, Albany-, Buffalo-, CW-, Fisher-, Long-Evans-, NIG-IV-, Wayne 

pinkeyed hooded-, MA- and YOS-strains belong to this type. 

Donryu-, NIG-111-, W/T- and WIS-strains are included in this type. 

heteromorphic pair no.3 consisting of a telocentric and a 

subtelocentric chromosomes. 

1. YOS-type (characterized by a subtelocentric pair no.3) : 

2. WIS-type (characterized by a telocentric pair no.3) : 

F1 hybrids between YOS- and WIS-strain rats had a 

Polymorphism in pair no. 3 was also found in 43 wild rats 

collected in Misima. They were classified into three types 

concerning pair no. 3. Among 43 rats, 17 were WIS-type, 3 were 

YOS-type and the remaining 23 were of hybrid-type. [Authors' sumary] 

Yosida, T. H., H. Kato, K. Tsuchiya and K. Moriwaki 1971. Karyot-vpes 11

and serun transferrin patterns of hybrids between Asian and Oceanian 

black rats, Rattus rattus. Chromosoma 34:40-50. 

KarvotvDes and serm transferrin Datterns were examined in Asian " ... and Oceanian black rats (R. rattus). Japanese R. r. tanezumi and 

Malayan R. r. diardii had-2nmut Australian-an7 New Guinea R. - r. 

rattus sKowed 2n=38hromosomes. F, hvbrids between JaDanese zd 

Australian rats and Malayan and New'Gu:nea rats had 2n=40 chromosomes 

which consists of the two genomes of both parents. Although various 

matings betwen the F1 h.ybrids were made, only one F2 male rat with 

2n=39 chromosomes was obtained. 

semisterile. Parental transferrin phenotypes were TfR in Japanese rats 

and TfCD in Oceanian rats. F1 hybrids examined showed TfRD in both 

male and female and F2.hybrid had TfR type transferrin. 

above investigations, it is suggested that Asian and Oceanian black 

rats are qeographical ly is01 ated and evolved different chromosomal and 

serm transferrin characteristics, but the sexual isolation of the two 

groups is incomplete at the present time. [Authors' abstract] 

The F1 hybrids seem to be 

Based on the 
Yosida, T. H., H. Kato, K. Tsuchiya, T. Sagai, and K. Moriwaki 1972 117 

Ceylon Population of black rats with 40 diploid chromosomes. 

Genet. 47( 6) :451-454. 

Jap. J. 

F1 hybrids between Asian and Oceanian black rats had 40 

chromosomes in diploid nunber, but the karyotype was usually a 

composite consistinq of two genomes of Asian and Oceanian black rats. 

F rats obtained from F1 hybrids in the laboratory and in natural 

pgpulation in Eniwetok island had 2n=39. 

a17 Ceylon black rats was 40 and the karyotype was well balanced 

consisting of all homozygous pairs. The karyotype of the Ceylon 

black rats with one large metacentric pair developed by Robertsonian 

fusion of pairs No. 11 and 12 should be a transient type from Asian 

to Oceanian black rats and by the following Robertsonian fusion of 

pairs No. 

somewhere in the Southwest Asia, and then they migrated to Europe as 

we already sugqested (Yosida -- et al. 1972). 

The chromosome number of 

4 and 7 the Oceanian type black rat with 2n=38 could occur 

[Authors' sumnary] 

Yosida, T. H., H. Kato, K. Tsuchiya, T. Sagai, and K. Moriwaki 

Cytogenetical Survey of black rats, Rattus rattus, in southwest 

central Asia, with special regard to the evmnal relationsh 

between three geoqraphical types. Chromosoma 45:99-109. 

1974. 118 

and 

P 

A chromosome survey of the black rat, Rattus rattus, was m de from 

animals collected at different localities inhwest and Central ' 

Asia. Asian t-vpe black rats (2n=42) were distributed in northern 

India, northern Pakistan, while the Oceanian type rats (2n=38) were 

found in southern India, southern Pakistan and Central Asia. A border 

line--of distribution of rats with Asian and Oceanian types can be drawn 

dividinq India and Pakistan into northern and southern parts. A hybrid 

type betwen Asian and Oceanian types was found in Karachi, Pakistan. 

Rats with 40 chromosomes, probably a transient type from Asian to 

Oceanian type, were found in Sri Lanka (Ceylon). 

these three geographic variants have developed via sequential events of 

Robertsonian fusion of acrocentric chromosomes in Asian type black 

rats. This fusion probably took place somewhere in southern India. 

The Oceanian type black rats that thus developed in southern India 

migrated widely to the rest of the world through Central Asia and 

Europe accompanying the movement of mankind. [Authors' abstract] 

It is suggested that 
Yosida, T. H.! A. Nakamura, and T. Fukaya 1965. Chromosomal 119 

polynorphism in Rattus rattus (L.) collected in Kusudomari and Misima. 

Chromosoma 16:70-78. 

Chromosomes of Rattus rattus (L.), collected in Kusudomari 

The homozygotic, 

(Nagasaki) and Misimmumere examined. 

a remarkable heteromorphism in chromosome no. 1. 

i.e., standard type, was characterized by 13 pairs of telocentric and 7 

pairs of metacentric chromosomes. Chromosome pair no. 1 was 

telocentric. X and Y chromosomes were also telocentrics. 18.4 per 

cent of rats from Kusudomari and 40 per cent from Misima showed 

heteromorphic Dair in chromosome no. 1. One chromosome of the 

heteromorphic pair is conspicuous by the subtelocentric centromere. 

Total length of the telocentric chromosome of no. 1 is almost the same 

as of .its subtelocentric partner. These facts indicate that the 

subtelocentric no. 1 chromosome might have arisen by a centromeric 

inversion of the telocentric chromosome. 

the subtelocentric no.1 chromosome could not be found in either 

population. 

collected in Kusudomari and Misima was statistically significant. 

Possible causes of the difference are discussed. [Authors' sumnary] 

The karyotype revealed 

Individuals homozygous for 

The difference in the frequency of the dimorphics 

Yosida, T. H. and T. Sagai 1972. Randinq pattern analysis of 120 

pol.porphic karyotypes in the black rat by a new differential staining 

technique. Chromosoma 37:387-394. 

Polymorphic karyotypes of black rats (Rattus rattus) collected in 

Japan, Australia and India were analysed by a new differential staininq 

technique by which banding patterns in the metaphase chromosomes are 

revealed. The technique consists in two steps: immersion of slides in 

a mixture of 2 X SSC and 0.1% (w/v) SDS (sodium dodecyl sulfate) for a 

few seconds at room temperature, and staining in Giemsa. 

treatment characteristic banding patterns were obtained in each 

chromosome pair. From the banding pattern analysis, subtelocentric 

pairs No. 1 and 9, which are polymorphic in respect to the acrocentrics 

and the subtelocentrics, were proven to have originated by pericentric 

inversion in the acrocentrics. The origin of two large metacentrics 

observed in Australian and Indian black rats was confirmed to have been 

developed by Robertsonian fusion of the acrocentrics No. 4 and 7 and 

No. 11 and 12 present in the Asian type black rat. 

abstract] 

By this 

[Authors' 
Yosida, T. H. and T. Saqai 1973. Similarity of Geimsa banding 

patterns of chromosomes in several species of the genus Rattus. 

Chromosoma 41:93-101. 

Geimsa banding patterns of chromosomes in seven Rattus species 

were compared. Four species (R. rattus tanezumi, R. norveqicus, R. 

exulans and R. meulleri) had aT1 =and their karyotypes and bznding 

patterns were similay, although slight differences were observed. 

Another subspecies (R. rattus rattus) and two other species (R. 

fuscipes and R. conaTus')hadfewerromosomes than the above-species 

by having 1 arse biarmed chromosomes developed probably by Robertsonian 

fusion. The origin of the arms of biarmed chromosomes was recognized 

by their characteristic banding patterns. The remaining species, R. 

sabanus, had a karyotype markedly different from the other species-by 

havinq two small metacentrics althouqh in the others their number was 

7. 

also very similar to those of the other, and therefore the 7 small 

metacentrics seemed to have orginated by pericentric inversion of small 

acrocentrics. [Authors' abstract] 

Banding patterns of the chromosomes in this species, however, were 

Yosida, T. H. and T. Sagai 1975. Variation of C-bands in the 

chromosomes of several subspecies of Rattus rattus. Chromosoma 

50 : 283-300. 

121 

122 

All subspecies of black rats (Rattus rattus) used in the present 

study are characterized by having large anmr C-bands at the 

centromeric region. The appearance of the bands, however, is different 

in the subspecies. Chromosome pair No. 1 in Asian type black rats 

(2n=42), which are characterized by an acrocentric and subtelocentric 

polynorphism, showed C-band polynorphism. 

rattus mindanensis) the pair was subtelocentric with C-bands, bGt in 

Malayan black rats (R. rattus diardii) it was usually acrocentric with 

C-bands. In Hong-Kong (R.tmvipectus) and Japanese black rats (E. rattus tanezumi) it kasmorphic with respect to the presence of 

acrocentrics with C-bands or subtelocentrics without C-bands. The 

other chromosome pairs showed clear C-bands, but in Hong-Kong black 

rats the pairs bo. 2 and 5 were polymorphic with and without C-bands. 

In Japanese black rats, 6 chromosome pairs (No. 3, 4, 7, 9, 11 and 13) 

were polymorphic in reqard to presence and absence of C-bands, but the 

other 5 chromosome pairs (No. 2, 5, 6, 8 and 10) showed always absence 

of C-bands. Only pair No. 12 usually showed C-bands. C-bands in small 

metacentric pairs (No. 14 to 20) in Asian type black rats were 

generally large in size, but those in the Oceanian (2n=38) and Ceylon 

type black rats (2n=40) were small. In the hybrids between Asian and 

Oceanian type rats, heteromorphic C-bands, one large and the other 

small, were observed. 

evolution in the black rats, the C-band is suqgested to have a tendency 

toward the diminution as far as the related species are concerned. 

[Authors ' abstract 3 

In Philippine rats (R. 

Based on the consideration of karyotype 
, y, , 

chromosome types of the black rat, Rattus rattus, collected in Oceania 

and F1 hybrids between Japanese and Austramats. Jap. J. Genet. 

44(2) :89-91. 

A new chromosome type of the black rat, Rattus rattus, was found 

in an Oceanian population. 

large metacentric chromosomes were noted. 

rats, Rattus rattus, coming from Japan as well as from many other 

countries in South East Asia showed 43 chromosomes (Yosida et al. 

unpubl ished) . obtained in the F2 progenies, it might be assumed that these two 

qroups in Japan and Oceania are derived from a comnon ancestor and the 

process is attributable to a polymorphic chromosome a1 teration. 

They showed 38 chromosomes, among which two 

The chromosomes of black 

If normal segregation of the chromosome typercx be 

Meiotic behaviour of chromosomes in F1 animals and test crosses 

are under investigation. [Authors I summary] 

Yosida, T. H., K. Tsuchiya, and K. Moriwaki 1971. Karyotypic 

differences of black rats, Rattus rattus, collected in various 

localities of east and southeast Asia and Oceania. Chromosoma 

33( 3) 952-267. 

Karyotypes of several subspecies of black rats, Rattus rattus, 

collected in different localities of Asia and Oceania were examined 

with special emphasis on the relationship between the chromosome 

polynorphism and differentiation of the subspecies. 

black rats (R. - rattus) collected were as follows: tanezurni jalorensis 

from Japan; flavipectus and sladeni from Hong Konq; diardii from Kuala 

Lumpur, Malaysia; argentivenmm Kuala Lumpur, am and 

Celebes, Indonesia; mindanensis from Luzon and Mindanao, Phillipines; 

and rattus from Australia, New Zealand, and New quinea. Subspecies in 

Formosa, Korea and Thailand were not determined. All black rats 

collected in the above Asian districts had 42 diploid chromosomes, 

while those in Oceania had 38. The rats collected in Japan (tanezumi), 

Korea, Formosa, Thailand and Malaysia (diardii) had A/A No. 1 pair or 

polynorohic No. 1 (A/A, A/S and S/S) pamile those collected in 

Java and Celebes (argentiventer), Luzon and Mindanao (mindanensis) 

showed a higher frequency of S/S No 1. pair. 

occurrence of No.1 A/A pair of black rats in the Asian continent where 

the black rats oriqinated, it is suqqested that the original type of 

No. 1 chromosome pair of the black rats is A/A, and a pericentric 

inversion occurred in the acrocentric No. 1 chromosome and thus rats 

with subtelocentric No. 1 pair formed.--Black rats with 38 chromosomes 

were observed in Australia, New Guinea and New Zealand. These 

karyotypes seem to have developed by Robertsonian fusion of 4 

acrocentric pairs (No. 4 and 7, and No. 11 and 12) in black rats of the 

Asian t.vpe. A relationship between body size and chromosome 

constitution was observed in subspecies of the black rats. [Authors' 

abstract 3 

Subspecies of 

From the higher 

12
)IZartman, 0. L. and A. N. Rruere 1974. Giemsa banding of the 125 

chromosomes of the domestic sheep (Ovis -- aries). Can. 3. Genet. Cytol. 

16:555-564. 

A Giemsa bandinp procedure was used to construct a basic G-band 

idioqram for the domestic sheep. 

systematic manner accordinq to the routine recomnended for human 

chromosomes. 

of comparison for further studies on intra- and interspecific 

chromosome homologies in addition to identification of chromosomal 

abnormalities. 

The idiogram is labelled in a 

This pattern based on NaOH treatment, provides a standard 

Late replicating regions of chromosomal DNA were detected with 

tritiated thynidine. Partial homologies between G-bands and these late 

rep1 ic.ating areas were found. 

secondary constrictions were seen to coincide with late replicating, 

G-positive regions on the metacentric and X chromosomes. [Authors’ 

abstract 3 

Previously reported areas of prevalent 

Zimnerman, E. G. and M. R. Lee 1968. Variation in chromosomes of the 126 

cotton rat, Sigmodon hispidus. Chromosoma 24:243-250. 

Chromosomes were analyzed from 38 hispid cotton rats, currently 

assigned to the species Sigmodon hispidus, from populations in 

southeastern and western United States. Cotton rats from southeastern 

United States had a 2N of 52 and an F.N. which varied from 52 to 54. 

Specimens from Obion County, Tennessee, and Highlands County, Florida, 

were found to be polynorphic with a varying nunber of arms on the 

largest pair of autosomes. Cotton rats form Arizona had a 2N of 22 and 

an F.N. of 38;.each pair of chromosomes is distinguishable, and a 

nunbering system is proposed. The cytological data suqgest that cotton 

rats from the southeastern populations and those from the Arizona 

populations belong to separate species, though morphological characters 

do not indicate such a difference. [Authors’ abstract] 

Zivkovic. S.. B. Soldatovic, M. Milosevic. and I. Savic 1968. 127 

Analysis-of chromosomes of three populations of Citellus citellus from 

Serbia. Zool. Anz. 181(3/4) :181-185. 

The chromosomes of 3 populations of European suslik from Serbia 

were studied. The diploid nunber of chromosomes in somatic cells of 

described populations is 2n = 40. The morphologically identical 

chromosome groups were found in a1 1 analyzed populations. Morphologic 

analysis made it possible to differentiate 4 qrouDs of chromosomes by 

similarity. This classifications [sic.] could be helpful in further 

morphometric and comparative caryotypic investiqation in the genus 

Citellus and the species Citellus citellus itself. [Abstract from 

Bi o 1 og i c a 1 Abstracts] 

- 
APPENDIX A 

Articles Relating to Cytogenetics 

of Subspecific Taxa 
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Articles Relating to Cytogenetic Analysis 

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Articles Relating to Cytogenetics 

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Articles Relating to Cytogenetics of Taxa 

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