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Society of Systematic Biologists

Thirty-One Years of Systematic Zoology


Author(s): David L. Hull
Source: Systematic Zoology, Vol. 32, No. 4 (Dec., 1983), pp. 315-342
Published by: Taylor & Francis, Ltd. for the Society of Systematic Biologists
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Syst.Zool., 32(4):315-342, 1983

THIRTY-ONE YEARS OF SYSTEMATIC ZOOLOGY


DAVID L. HULL
Department
ofPhilosophy,
University
of Wisconsin-Milwaukee,
Milwaukee,Wisconsin53201
Abstract.-The numberand percentageof pages devoted to various topics in Systematic
Zoology
are traced fromthe inception of the journal to the present.The categoriesused include papers
that: (a) are largely descriptive;(b) argue for or against a particulartaxonomicphilosophy; (c)
apply the methods of a particular taxonomic philosophy; (d) are primarilyquantitative; (e)
discuss nomenclaturalproblems,and are eithertheoreticalor applied papers in (f) evolutionary
theory,(g) phylogeny reconstruction,or (h) biogeography. Changes in proportionsof contributions in these categories are mapped onto changes in editorship of the journal to see what
effectssuch changes have had on the sortsof papers published. The changes in Editorof primary
concern involve the move fromthe American Museum of Natural Historyto the Universityof
Kansas in 1963 and the returnto the AmericanMuseum 10 yearslaterin 1973. [Systematic
Zoology;
taxonomicphilosophy; evolutionarysystematics;phenetics; cladistics;numerical taxonomy.]

In 1947 at a meeting of the American SystematicZoology from its inception in


Association for the Advancement of Sci- 1952 to the present. What percentage of
ence, the Society of SystematicZoology papers have been largely descriptive?Of
was formedand had its firstannual busi- those papers thathave actually concerned
ness meetingthe nextyearin Washington, taxonomic philosophy, what percentage
D.C. In 1952 the Society began to publish have favoreda particularphilosophy; how
Systematic
Zoology.R. E. Blackwelder pro- many opposed? How have the ratios of
duced the firstnumber.Thereafter,the ed- positive to negative papers dealing with
itorship passed to John L. Brooks of Yale particular schools changed through the
University,who served as Editot for 61/2 years? What percentage of papers have
years. In 1958 the journal moved to the been largely applications of particular
American Museum of Natural History, taxonomic philosophies, and how many
where Libbie H. Hyman became Editorpro general discussions of quantitative methtemand, a yearlater,Editor.After51/2
years ods? Distinguishing between theoretical
in New York, the journal moved to the and applied papers, what percentagehave
Universityof Kansas, where it was edited concerned the evolutionary process, biosuccessively by George W. Byers (three geography and phylogeny reconstrucyears),RichardF. Johnston(fouryears)and tion? What percentage of papers through
A. J.Rowell (threeyears). In 1973 the jour- the years have dealt with nomenclatural
nal returned to the American Museum, issues? Finally, how have the books rewhere Niles Eldredge and Gareth Nelson viewed in Systematic
Zoologyfared?
edited it for three years, succeeded by
The question of greatest interest is,
Randall T. Schuh foranother three years. however, have changes in Editorbeen acIn 1980 James Dale Smith of the Natural companied by changes in the constitution
History Museum of Los Angeles County of the journal?Sociologistsof science claim
took over fora three-yearterm.
thateditorsand theirpreferencesare very
Everyone who has read Systematic
Zool- significantfactorsin the publishing proogythroughthe years has his or her own cess. For example, W. D. Garvey (1979:84)
ideas about the effectsthat these changes stated that,"like most scientists,most edin editorship have had on the content of itorshave attitudesabout what constitutes
the journal, but such casual, haphazard quality in theirjournal. These attitudesare
observations are notoriously untrustwor- influenced by their theoreticalbias, their
thy. In this paper I trace the history of methodologicalpreferences,etc.,and they
315

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316

SYSTEMATIC ZOOLOGY

tend to select consulting editors and referees who share them." Has this been true
of the Editorsof Systematic
Zoology?I would
be less than candid to not admit that I expected significantchanges when Systematic Zoology moved to Kansas and again
when it leftKansas forthe American Museum, while I did not anticipate any radical changes in editorial policy when the
journal moved across the continent from
New York to Los Angeles. Why is this?
The obvious answer is that the shiftto
Kansas was accompanied by the rise in the
United States of a particularschool of taxonomy centered at Kansas, and the move
to the American Museum happened to coincide with the rise in the United States
of yet another philosophy of systematics,
this time emanating from the American
Museum. Althoughone event cannotcause
another unless the two events are coincident, coincidence does not guarantee
causation. Perhaps these correlationswere
merelyaccidental. Afterall, Allen Press is
in Lawrence, Kansas. Perhaps thatwas actually the most relevant causal factorfor
the move to Kansas. However, I thinkthat
it is verydifficult
to discount the roles that
the early leaders of these two schools of
taxonomy played. Protests to the officers
of the Society of SystematicZoology from
Robert R. Sokal about Hyman's treatment
of papers with numbers in them was instrumentalin transferringthe journal to
Kansas, while Nelson's complaints to
Rowell were an instigatingcause for the
move to New York. Systematic
Zoologyhad
been at Kansas long enough. It was time
fora change.
One chief differencebetween the two
situationsis thatSokal was never Editorof
the journal, and Byers can hardly be
termed a strong advocate for either phenetics or numerical taxonomy. Although
Johnstonand Rowell were somewhatmore
committed to the phenetics movement,
neitherof themwas extremelyvocal in his
position. Yet, my initial suspicion was that
having a journal at one's own institution
is liable to make access to its pages a bit
easier. The situation afterthe move from
Kansas is somewhat clearer. All four sub-

VOL. 32

sequent Editorswere openly committedto


the cause of cladistic analysis, and one of
them was Gareth Nelson.
One minor but irritatingdifficultyin
deciding the effectthatchanges in editorship had on editorial policy in Systematic
Zoologyis thatsome Editorswilled quite a
few papers to their successors,while others cleared out their files before leaving
office.A second complicationis that,at the
end of 1975, Eldredge and Nelson instituteda systemof Associate Editors.As this
systemusually worked, Associate Editors
could rejecta paper on theirown but could
accept one only contingentupon the Managing Editor's approval. On occasion, an
Associate Editor disagreed with the Managing Editor.A few cases of such disagreements are fairly well known. However,
because no figures exist concerning the
number of these disagreements or their
eventual resolution, I have ignored this
complication. I have divided papers according to Managing Editors,not Associate Editors,attributingpossibly too much
influence to the former.I also have not
distinguishedbetween Eldredge and Nelson.
The major omission in this paper is that
I do not discuss the refereeingprocess. In
the early years, rejection rates were quite
low, rarelyexceeding 10%. Under such circumstancesone cannot attributetoo much
influenceto the Editor. But, as the size of
the journal increased,so did rejectionrates.
Under Johnstonthey ranged from20% to
over 30%. Thereafter,they continued to
rise until they reached as high as 50% to
60%. The higher the rejection rates, the
more influence an Editor can have. One
mightalso expect author dissatisfactionto
correlatestronglywith increased rejection
rates.But rejectionratesfromEditorto Editor are not strictlycomparable because of
the different
reportingproceduresused by
differentEditors, some including papers
sent back for revision, others not. I have
omitted any reference to the refereeing
process chieflyfor two reasons. First,it is
confidential,and second the relevant data'
are not available. (The SystematicZoology
archives at the Smithsonian Museum are

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1983

ANALYSIS OF SYSTEMATIC ZOOLOGY

317

very sparse until the tenure of Eldredge My justificationforthisdecision is thatoband Nelson.) A few examples of disputes jections to quantitative methods were
over the handling of papers are legendary, short-lived,while the controversiesover
but I think it would detractfromthe pur- taxonomic philosophy continue. Rather
poses of this paper to discuss them here. quickly the question became not whether
Any impressions I might give would be to use quantitative methods but which
just that-impressions and nothing more. ones. Classing all quantitative papers toAs dissatisfyingas my decision is likely to gether year afteryear would not be very
be to certain readers, in this paper I deal instructive. Papers advocating quantitaZo- tive methods appeared in Systematic
only with the public face of Systematic
Zoolology,the resultsof the refereeingprocess ogylong before the Kansas group got toand not that process itself.
gether,including a posthumous paper by
Categorizing the papers in Systematic Stroud(1953), a close friendof Sokal's from
Zoology for a period of 31 years during college. And many of the quantitativepawhich issues, alliances, interests,and even pers after the advent of Numerical Taxterminologyhave changed drasticallyis far onomy had no apparent connection to the
from easy. I had always been convinced effortsof Sokal and his group. In this pathatthe way one categorizesa subjectmat- per I am not interestedin the frequency
terstronglyinfluencesthe resultsof one's of disembodied ideas but in the influence
investigations,but the currentstudy has of particularresearchprogramson the sysdriven this truthhome to me more con- tematics community.However, ignoring
vincingly than any general arguments all the papers in Systematic
Zoologywhich
could have. For example,in the earlyyears, advocated quantitative methods, but did
the Kansas school was pushing two,at least not happen to mention phenetic philosopartiallyindependent,positions-phenet- phy,would also be misleading. The efforts
ics and numerical taxonomy. Phenetics of the group of workers initially located
concerned a philosophy of classification. at Kansas did give increased impetus to
According to one overly condensed char- the use of quantitativemethodsin systemacterization of this philosophy, classifica- atics. My decision was to score papers first
tions should be constructedon the basis on professed systematicphilosophy and
of numerous, unweighted (or equally then to add later two additional sorts of
Weighted) characters,at least initially,so papers-those applying phenetic methods
that organisms are clustered according to and those that discuss quantitative methoverall similarity. Numerical taxonomy ods. Although this compromiseis farfrom
embodies the conviction that the proce- ideal, it is the best I could do.
dures of classificationshould be as quanCladistics also provided some problems.
titativeas possible. Earlyadvocates of these Cladistics as a taxonomic philosophy had
two views saw a close connectionbetween two independent origins in the United
them. In order to be sufficiently
quantita- States-J. S. Farris,firstat the University
tive and objective, a classificationhad to of Michigan and then at Stony Brook,and
be phenetic-at least initially.
Gareth Nelson at the American Museum.
As it turns out, there is a large overlap Initially, Farris was concerned with debetween papers advocating phenetic phi- veloping quantitative methods for inferlosophy and quantitativemethods,at least ring phylogenies. Later he became inin the early years,but then the two seem creasingly involved in debates over
to have gone theirseparate ways. The con- taxonomicphilosophy,becoming the chief
troversyover pheneticssubsided while the critic of phenetic taxonomy. Of course,
number of papers utilizing or discussing otherworkerswere also writingpapers on
quantitative methods remained high. In numerical cladisticsat the time,including
this paper I treat the controversiesover Sokal himself.The question is how to score
taxonomicphilosophy as primaryand the these papers. To begin with, most of the
use of quantitativemethods as secondary. papers in numerical cladistics do not con-

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318

SYSTEMATIC ZOOLOGY

cern taxonomic philosophy but phylogeny reconstruction(Fitch and Margoliash,


1967; Fitch, 1971). The two, of course, are
not unconnected.Anyone who argues that
phylogenies are impossible to reconstruct
is not likely to argue that classifications
should be based on phylogeny. However,
devising methods forreconstructingphylogeny can be and usually has been distinctfromthe formulationof methods for
translatingphylogeneticinformationinto
classifications.In this paper, I distinguish
between the two. Many numericalcladists
emphasize the numerical side of their
work. I have scored these papers as contributingto "numerical taxonomy"in the
broad sense (quantitativemethods). Other
numerical cladists emphasize the cladistic
side of their work in opposition to phenetics. I have scored these papers as contributions to cladistic philosophy, not
"numerical taxonomy." Subdividing numerical cladistics in this way is likely to
strikesome as unnatural. If my goal were
to show the continuinguse of quantitative
methods in systematics,it would be, but I
see no real point in documenting the obvious. A significantnumber of practicing
systematistsuse quantitative methods. In
this sense, "numerical taxonomy" has
triumphed. The interesting question is,
how about phenetics?
I have made a parallel decision for cladistics and vicariance biogeography.
Whether or not there is a necessary connection between these two sets of ideas, it
just so happens that nearly all the papers
arguing for vicariance biogeography are
by cladists,and these papers have contributed indirectlyto the cladistic movement.
Ignoring them would be misleading. Including them with papers arguing forthe
principles of cladistic systematicswould
be just as misleading. I have opted to add
two additional sortsof papers to the list of
papers classified as cladistics-those applying cladistic methods and those advocating vicariance biogeography. As in the
case of phenetics,I have representedthese
threeclasses of papers so thatanyone who
disagrees with the decisions I have made
can tell which papers are which.

VOL.

32

By now it should be obvious thatI have


not employed a pure strategyin categorizing the papers that have appeared in SystematicZoology.I have not classifiedthem
entirelyin terms of similarityof content
or entirely in terms of the people who
wrote them and their professional allegiances. Instead, I have attemptedto balance these two considerations,paying attention both to the content of the papers
and to their authors. As dissatisfyingas
this compromise may be, the strictutilization of "pure" methods of classification
produced monstrous categories-grouping together papers by deadly enemies.
Scientistslike to thinkof science as an entirelycerebral process, a conflictbetween
disembodied ideas, but scientistsare supposed to be partialto evidence, and all evidence indicates thatthis view of science is
mistaken. At least in the short run, who
presents an idea mattersas well as what
this idea happens to be. Perhaps science
should not be this way, but it is.
Finally, a list of warnings. I have limited myself almost entirely to the controversies that have taken place in the pages
of Systematic
Zoologyand the groups located in the United States. I mention books
and papers thatappeared in otherjournals
only in passing. Phenetics was also an issue in Great Britain,and cladistics originated in Germany.I have largely ignored
these aspects of the controversies.Hence,
when I talk of "cladistics" or "phenetics,"
I always mean to imply "in the United
States" even when I do not mention the
restriction.Many papers also dealt with
more than one topic and were classed in
more than one category(e.g., a given paper might count as a contributionto cladistics, biogeography and phylogeny reconstruction). Hence, most percentages
across categories tend to add up to more
than 100%. Many papers were, as one
might expect, borderline cases and could
have been scored as naturallyin one category as in another. Others might have
preferredothercategoriesand have scored
Because of the
particularpapers differently.
number of decisions that went into the
tabulations I present in this paper, they

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1983

319

ANALYSIS OF SYSTEMATIC ZOOLOGY

Pages

600

I
YaleMuseum

Yale

American

Kansas

KansasMuseum

500

450

40011III
350
3001IIII

550

250

200
150

A1r 1E
1
Brooks

100

501

0
52

I
mi1

lEdredgel
Johnston

-.

Rowell

I POINTS OF VIEW
-

1cu1

j Byers

53 54 55 56 57 58 59 60 61 62

LA

Msu

1
Hyman

American

akNelson

63 64 65 66 67

68 69

I -,J

,,A

\8 REVIEWS

70 71 72 73 74 75 76 77

78 79 80 81 82

Zoology.
FIG. 1. Number of pages per year devoted to articles,points of view, and reviews in Systematic

should be taken with a grain of salt. As


numbers, they may look exact, objective,
etc.,but theyare not. As a result,the only
trends and differencesthat can bear any
weight are those that are extremely
marked. To give the reader a feel forhow
I classified papers, my classification for
1975 is included in the Appendix.
GROWTH OF THE JOURNAL

During the firstdozen years of its existence, Systematic


Zoologyaveraged roughly
200 pages a year. While it was at Kansas,
the journal expanded steadily to its current size of roughly 500 pages a year.
Withina few issues, Systematic
Zoologysettled down to its currentformat-articles,
points of view, book reviews and news.
Initially, one differencebetween articles
and points of view was length. Through
1977, long points of view tended to average about eight pages. In 1978 points of
view increased in length; one by Farrisin
1980 was the longest at 22 pages. A second

differencebetween articles and points of


view is tone. Fromthe start,points of view
have tended to be somewhat more polemical than articlesand theirtone a good deal
sharper.The percentage of pages devoted
to articles climbed steadily in the early
years until the journal moved back to the
AmericanMuseum in 1974,when it began
to decline. In 1980, the number of pages
taken up by points of view and reviews
equaled thatforarticles(see Fig. 1 and Table 1). If increases in the size of the points
of view section are any indication of periods of increased controversy,the peak
yearswere 1969,1975 and 1978-1982.Tone
to one side, articles and points of view
have dealt with exactly the same topics.
Furthermore, the policy regarding responses in Systematic
Zoologyduring most
of its existence was that an author could
respond to an article, the author of the
original articlecould then respond to that
response, and that was that. As a result,
papers that amounted to responses to responses were submitted as independent

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320

SYSTEMATIC ZOOLOGY

VOL.

32

ures forthe journal have also reflectedupsurges in controversy.In 1948, the Society
had 343 chartermembers. By 1952 memPoints
bership had surpassed 1,200, and 371 adReviews
of view
Editor
Articles
vanced subscriptionshad been placed for
8
2
85
Brooks,
the journal. Figure 2 shows the fluctua92
7
1
Hyma'n
tions in circulationand membershipfrom
8
2
87
Byers
1962 to the present. The firstpeak oc12
3
81
Johnston
curred in 1968-1969 at the height of the
7
3
Rowell
86
10
7
79
Eldredge and Nelson
controversyover phenetic taxonomy.Dur10
65
19
Schuh
ing the early 1970s,both membershipand
58
11
29
Smith
circulationbegan to sag, only to pick up
again as the controversyover cladisticsbegan to heat up. In 1982 the figuresbegan
articles, furtherblurring the distinction to drop once again. If these figuresare any
between articlesand points of view. In all indication,controversyhas been good for
subsequent tabulations,I have classed ar- the Society and its journal, at least as far
ticles and points of view together. Book as numbersare concerned. These statistics
reviews are treatedseparately.
notwithstanding,contributorsto the jourBoth membership and circulation fig- nal have frequentlycomplained of what
TABLE 1. Percentageof pages devoted to each subdivision of SystematicZoologyduring the tenure of
each Editor.

Kan sas

2900

American Museum

2800L
2700

Museum

2600
2500
2400
2300
2200

2100
2000
1900
1800
1700
1600
1500 |

/M

EMBERSHIP

65

66

1400
1300

64

67

68

69

70

71

72

73

74

75

76

78

79

80

81

82

FIG.2. Figuresformembershipin the Societyof SystematicZoology and subscriptionsto Systematic


Zoology
between 1964 and 1982.

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1983

ANALYSIS OF SYSTEMATIC ZOOLOGY

they perceived to be excessive polemics.


Such objections surfaced early. For example, beforethe journal was a year old, one
subscriberwrote, "Enclosed please find a
one-dollar bill ($1.00), my dues for 1952;
and, also accept my resignation fromthe
above Society,as its principles do not appeal to me. We need more old-fashioned
constructiveefforts,not the communistic
'tearing down of all old established rules
and methods"' (Syst.Zool., 1:138, 1952). It
is quite easy to determinehow much dues
have increased since 1952, less easy to
gauge the rise and fall of polemics.
PURPOSES OF THE JOURNAL

Accordingto the initialstatementon the


inside back cover of Systematic
Zoology,the
purpose of the journal was to encourage
the preparationof "contributionson basic
aspects of all fields of systematics,principles and problems; to provide a suitable
forum for discussion of the problems of
the systematistand his methods; and to
report as news the other activities of the
Society of Systematic Zoology." Listed
among the topics solicited were "principles and the applications of principles of
wide implication and general interestin
any phase of systematics,such as comparative anatomy, zoogeography, paleontology, taxonomy,classification,evolution,or
genetics."As explicitas thisstatementwas,
Blackwelder (1952:92), as Secretary-Treasurer of the Society, had to publish the
following reminderin the second number
of the journal:

321

paleontology, genetics and classificationthatbear


directlyon systematics,and other papers of general interest to systematistsare particularlydesired. Taxonomic descriptions,revisions,and keys,
or papers on anatomy,physiology, ecology, etc.,
having little or nothing to do with systematics
cannot be accepted [Syst.Zool., 1964].

When Rowell became Editorin 1971, he


dropped the final sentence of the above
description, and finally, when Eldredge
and Nelson assumed the editorship three
years later,they substitutedthe following
brief statementin 1975 fromthe Constitution of the Society:
The objectof thissocietyshall be the advancement
of the science of systematiczoology in all its aspects of theory, principles, methodology, and
practice, for both living and fossil animals with
emphasis on areas of common interestto all taxonomistsregardlessof individual specialization.

Throughoutthe historyof Systematic


Zoology,the intent of the Council and successive Editors has been clear. The main
purpose of the journal was to publish papers on taxonomic theory firstand foremost, and papers on other topics only to
the extentthat theybore directlyon taxonomic theory.Such good intentionswere
easier to proclaim than to practice. No
sooner had Blackwelder reminded the
readers of the journal that it did not publish descriptivepapers than Brookshad to
ask the Council to authorize him to publish just such papers in order to keep the
journal up to size (Blackwelder, 1977:114).
Although such categories are difficultto
define and even more difficultto apply in
particularcases, the only conclusion that
It has perhaps not been sufficiently
emphasized
one can draw after reading through 31
thatSystematic
Zoologyis not intended forthe publication of descriptivepapers. The Council sees its yearsof the journal is thatmorepages have
field as the philosophic aspect of systematics,its been devoted to descriptivework than to
principles and problems, as well as news of sys- any other topic. From its inception until
tematists,their institutions,courses, and publica- Schuh took over as Editorin
1977,roughly
tions. Systematicmaterialis not completelyruled
Zoologyhave
out, but it must be subordinate to the discussion 50% of the pages of Systematic
been taken up with descriptionsof chroof principles.
mosome numbers, geographic distribuA year afterByerstook over the editor- tions, particular phylogenies, classificaship of the journal in 1963, its statement tions and the like (see Fig. 3 and Table 2).
of purpose was made even more pointed:
Given the professedgoals of the journal,
authors
tried to make their papers appear
Contributionsrelating to principles and methodology of systematics,as well as articles in such as theoreticaland philosophical as possifields as evolution, morphology, zoogeography, ble, but in most cases they were not very

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322

SYSTEMATIC ZOOLOGY

100%

Brooks

90%

Byers

Johnston

RoIweIllaNelsoni

80%X0

70

Hymon

VOL. 32

Rowell

Schuh Smt
Smith
Scu

ldredge

Il

0/

I
I

60%

20%
.

50%1

~~~~~I ~

I
II

40%
30%

20

Lii

/IIIIIII

0%

0%

1
52 53 54 55 56 57 58 59 60

61 62 63 64

65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80

81 82

FIG. 3. The percentage of pages per year thatare largely descriptive.

successful.For example, the firstpaper to


appear in the journal, by Alexander Petrunkevitch(1952), the firstPresident of
the Society, was entitled "Principles of
Classificationas Illustrated by Studies of
Arachnida." Although Petrunkevitchbegan and ended his paper with allusions to
"unraveling the mysteryof evolution,"
there is little in his paper that can count
as principles of classification. Similarly,
most papers entitled somethinglike "The
Evolution of the House Mouse" were
straightforwarddescriptionsof particular
taxonomic groups, their characteristics,
distributions,chromosome numbers, etc.

TABLE 2. Percentage of pages during the tenure


of each Editordevoted largely to the presentationof
data and descriptions.
Editor

Percent

Brooks
Hyman
Byers
Johnston
Rowell
Eldredge and Nelson
Schuh
Smith

50
58
44
50
55
62
35
28

They mightconclude with a paragraph or


two on gene flow, but that usually was
about it. Most of the papers listed as descriptiveare justthat,descriptive,but some
also contained enough general discussion
to count as theoreticalas well.
In classifyinga paper as "descriptive,"I
have no intention of denigrating it. Descriptionis a necessarypart of science. In
fact,in a poll of the members of the Society of SystematicZoology published in
1981 (Syst. Zool., 30:224, 1981), 17.8% of
those responding wanted the journal to
publish more "substantive,empirical papers including materials, the use of examples in theoretical papers with application to actual cases." The preferencesof
many readers notwithstanding,Systematic
Zoologywas not founded to publish primarilydescriptivework. Until the editorships of Schuh and Smith, when the figures dropped to 35% and 28%, respectively,
it did (see Fig. 3 and Table 2). Even though
taxonomicphilosophy was to be the main
focus of the journal, papers dealing primarily with taxonomic philosophy averaged only 17% under Brooks, gradually
climbingto a high of 30% under Johnston,
and dropping again to a low of 10% under

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50

323

ANALYSIS OF SYSTEMATIC ZOOLOGY

1983
/0

rooks

Byers

Hyman

BroksH

jNelsoni

mola

400/

35/

Johnston I Rowell I Eldredgel Schuh I Smith

25 %

20 %II
15 %II

ILi

100/

52 53 54 55 56 57 58 59 60 61 62 63 6465 6667 68 69 70 71 72 73 74 75 76 77 7879 80 81 82


FIG.4. The percentage of pages per year discussing taxonomicphilosophy.

Eldredge and Nelson, o'nly to rise again


abruptlyunder Schuh and Smith (see Fig.
4 and Table 3). During the early years of
the journal, the firstreally big debate was
over the subspecies question (see later discussion); the issues turnedas much on differencesin beliefs about the evolutionary
process as about taxonomicphilosophy. If
papers concerning this controversy are
disregarded,the percentages of pages devoted to taxonomic philosophy are even

TABLE 3. Percentage of pages during the tenure


of each Editor devoted to taxonomicphilosophy.
Editor

Percent

Brooks
Hyman
Byers
Johnston
Rowell
Eldredge and Nelson
Schuh
Smith

17
22
25
30
17
10
32
33

lower (e.g., Brooks 9%, Hyman 14% and


Byers23%).
Others might well score individual pathan I have, but it would
pers differently
be difficultto avoid the conclusion that
papers dealing primarilywith taxonomic
philosophy have not been as prominentin
Zoologyas succesthe pages of Systematic
sive statementsof purpose mightlead one
to expect. Looking back over the first25
yearsof thejournal,Blackwelder(1977:207)
raised just this objection, complaining of
so manystudies of "evolution, statistics,or
the revisionary taxonomy of animals."
Even so, Slater (1978) responded that the
journal has been the chief medium
through which the great conceptual debates between various schools of taxonomy have been aired. Slater is surely right,
but this end has been accomplished in
roughly 23% of the pages of the journal.
TAXONOMIC PHILOSOPHIES

The question thatis likelyto be of greatest interestto long-timereaders of System-

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324

VOL. 32

SYSTEMATIC ZOOLOGY

aticZoologyis how many pages of the journal have been devoted to the various
taxonomic philosophies. Once again, definingthe relevant categories is not easy,
especially in the earlyyearswhen the lines
were not clearlydrawn. For example, during the firstdecade of the journal, the prevailing view was, roughly speaking, that
the evolutionary process and phylogeny
were somehow relevant to classification.
Species had to be the units delimited by
reproductive isolation, and higher taxa
should reflectphylogeny. The only real
opponents of this view were Blackwelder,
Alan Boyden, R. S. Bigelow, and Thomas
Borgmeier.According to Blackwelder and
Boyden (1952:31), the "grand object of
classificationeverywhereis the same. It is
to group the objectsof studyin accordance
with their essential natures." Borgmeier
(1957:53) agreed, contending that, as the
science of order,"systematicsis a pure science of relations,unconcerned with time,
space, or cause." These systematistsagreed
that evolution has occurred and that the
distributionof organismsnow apparent is
the result of phylogeny,but they saw no
reason to allow such considerationsto intrude into taxonomy,and numerous reasons to exclude them.Phylogenyis too difficult to reconstruct, the little fossil
evidence available is too spotty,our understandingof the evolutionaryprocess is
highly contentious,and charactersare basic anyway. If the only data thata systematisthas available are characters,if everyone always begins with characters,then
why not just stick with characters and
abandon idle speculation about phylogeny and the evolutionary process, at least
in conjunctionwith classification?
If all I were doing was classifyingtogether papers urging similar views, these
early objections to the prevalent "evolutionary"philosophy mightwell be counted as contributionsto phenetic taxonomy
or even to certain varieties of cladistics.
The criticismssound verysimilar.Butthese
authorswould rightlyobject.Neitherphenetics nor cladistics as scientific movements existed at the time in the United
States and, even after they did emerge,

TABLE 4. Percentage of pages devoted to each


taxonomicphilosophy during the tenureof each Editor.

Editor

Evolutionary Phenetic Cladistic


taxonomy taxonomy taxonomy

Brooks
Hyman
Byers
Johnston
Rowell
Eldredge and Nelson
Schuh
Smith

16
16
20
12
11
5
14
6

11
16
19
9
1
9
15

7
8
9
8
30
30

these early criticsof evolutionarytaxonomy kept their distance. Blackwelder for


one claimed to be urging quite a different
philosophy of classification,the philosophy implicit in the practice of most taxonomists,a philosophy which he termed
"omnispective." In thispaper I ignore omnispectivetaxonomyforthe simple reason
that the number of pages devoted to it in
Systematic
Zoology,both pro and con, is so
small thatincludingitwould not make any
differenceand would complicatediagrams
that are already too difficultto read.
As Table 4 indicates,papers arguing the
virtues and vices of evolutionarysystematics have remained a constant featureof
the journal. The low figure for Eldredge
and Nelson is a reflectionof the few pages
devoted to taxonomic philosophy during
their tenure. The low figureunder Smith
is not. When these papers are divided into
those that defend evolutionary systematics and those that attack it, a pattern
emerges,but it does not coincide with any
putative philosophical preferencesof the
Editors (see Fig. 5). Instead, it exhibits a
lag effectbetween attacks and defenses.
The firstanti-evolutionistonslaughtby the
pheneticiststook place in 1961 during the
tenure of Hyman. The evolutionists responded in 1962 and then,aftercontinued
criticism,launched a massive defense in
1965 during Byers'tenureas Editor.It was
followed by another attack in 1967 met
immediatelyby an evolutionary counterattack in 1968, both while Johnstonwas
Editor.

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1983
PageOs

Brooks

70

60

325

ANALYSIS OF SYSTEMATIC ZOOLOGY

I
Hyman
1~~~II

Pro Evolutionary
Systematics

'Byers 'Johnston 'RowelllEldredgelSchuh 'Smith

laNelson
I

40

30
210
Ic0
0
I0
5

20

Evolutionary
40

50

Systematics

60
52

54

56

58

60

62

64

66

68

70

72

74

76

78

80

82

FIG. 5. The number of pages each year arguing foror against evolutionarysystematics.

The firsthint of the phenetics research


programto appear in the pages of SystematicZoologywas a paper by Michener(1957:
166) in which he wondered "whether or
not it would be desirable to change from
the traditional approaches emphasizing
phylogeny to approaches utilizing only
static relationship." In that same year,
Michener published a paper with Sokal on
a quantitative approach to classification,
and Sneath published two papers urging
similar views (Michener and Sokal, 1957;
Sneath, 1957a, b), but these papers did not
appear in Systematic
Zoology.Most of the
early papers in Systematic
Zoologyadvocating phenetics (as well as quantitative
methods) were measured in their criticisms of traditionalevolutionary systematics and subdued in their polemics (e.g.,
Sneath, 1961; Sokal, 1961; Rohlfand Sokal,
1962; Michener, 1963; Rohlf, 1963). It was

left to Ehrlich (1958, 1961, 1964) to goad


advocates of traditionaltaxonomic methods. At this time, Simpson (1961) published his Principlesof Animal Taxonomy
and, shortlythereafter,Sokal and Sneath
(1963) published theirPrinciplesofNumerIn response to these attacks,
ical Taxonomy.
Mayr (1965) published his major critique
of phenetic taxonomy. Simpson (1964,
1965) published his responses to phenetics
elsewhere. Finally,the pheneticistsjoined
forcesto issue collective responses (Sokal
and Camin, 1965; Sokal et al., 1965).
Strangely enough, one of the earliest
critics of phenetics was Kiriakoff(1962,
1963), a disciple of Hennig who thought
of himself as defending a single alternative taxonomic philosophy subscribed to
equally by Simpson, Mayr and Hennig.
However, Sokal and Sneath (1963) eventually convinced Kiriakoff(1965:63) that

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326

SYSTEMATIC ZOOLOGY

Pages
80

Brooks

70

60

Hyman

VOL.

32

IByers IJohnston I Rowell lEIdredgel Schuh ISmith


I
INelsoniI

laI

Pro PheneticsI

40

II
I

30I

I 0~~~~~~~~~~~~~~~~~~
I

20II
30I

40II
50II

I
I
II

60

An/iPhene/ics

I
IIII

70IIIII
52

54

56

58

60

62

64

66

68

70

72

74

76

78

80

82

FIG. 6. The number of pages each year arguing foror against phenetic taxonomy.

phylogeneticclassificationsare usually not


possible "owing to lack of necessary information"and that only "ersatz" classificationsare possible. Commentsmade by
Mayr (1965) furtherforcedKiriakoff(1966:
93) to distinguishbetween Hennig's views
and those of Simpson and Mayr,realizing
that "cladists are but a rathertare subspecies of taxonomists."That state of affairs
was not to last forlong. Nelson (1972) went
on to subdivide the evolutionistsfurther,
pointing out significantdifferencesin the
philosophies of systematics set out by
Simpson and Mayr.
As Table 4 indicates,papers concerning
phenetics reached theirpeak under Johnston, began to drop under Rowell while

the journal was still at Kansas, dropped


even lower under Eldredge and Nelson,
and then began to climb again under
Schuh and Smith. The same seesawing
patternfound in connection with evolutionarysystematicscan be seen in the data
forphenetics(Fig. 6), in partbecause many
attackson the evolutionistswere also defenses of phenetics, and vice versa. The
firstmajor attacks on phenetics came in
1965, a series of responses appeared in
1967, more attacks in 1968, defenses in
1969, attacks again in 1970, and so on.
Mayr's (1969) Principlesof Systematic
Zoologyalso appeared at this time.The rise and
the fall of papers dealing with phenetics
occurred while SystematicZoologywas at

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1983

ANALYSIS OF SYSTEMATIC ZOOLOGY

327

Kansas, the original home of phenetic tax- (1970). However, the issues were not all
onomy. (Sokal and Rohlf moved to the that clear in the early years. For example,
State University of New York at Stony Kluge and Farris (1969:14) can be found
Brook in 1969.)
saying that "it is quite reasonable to state,
Phenetics was hardly discussed while 'Mammals are derived fromreptiles,' imEldredge and Nelson were Editors, but plying merely that all mammals had a
then neither were any of the other taxo- common ancestor thatwas a reptile,quite
nomic philosophies. If the percentage of independently of the fact that Mammalia
pages devoted to taxonomicphilosophy in and Reptilia are both 'modern' classes."
SystematicZoology during this period is This belief was to be short-lived(Farriset
any indication,taxonomicphilosophy had al., 1970).
fallen on lean times. Sneath and Sokal
As Table 4 indicates,papers dealing with
published their second edition in 1973, cladisticsincreased steadily while Systemwhile the volume by Hecht et al. did not atic Zoology was at Kansas, dropping
appear until 1977. Were theoretical tax- slightlyunder Eldredge and Nelson, and
onomists publishing extensively else- then climbing significantlyunder Schuh
where? Not thatI could discover.The phe- and Smith. The seesawing of attacks and
neticiststurnedtheirattentionaway from defenses is not as apparent with respectto
taxonomic philosophy and toward devel- the cladists as it was forevolutionarysysoping and applying a variety of mathe- tematics and phenetics (see Fig. 7). Almatical techniques. The cladists for some though Crowson's (1970) Classification
and
reason published little on taxonomic phi- Biologyappeared just as the controversyof
losophy during this period. The major pa- cladistics was getting underway, it had
pers on taxonomic philosophy to appear surprisinglylittle impact. A second surge
at this timewere not published in System- of activityoccurred in 1973, but the main
aticZoology(Mayr, 1974; Sokal, 1974). Un- attacks(1977 and 1979) as well as defenses
der Schuh and Smith, things happily (1978-1982) occurred under the editorpicked up again-happily at least fromthe ships of Schuh and Smith.
One surprisingfeatureof Figures 6 and
perspective of those of us who make a
profession of arguing systematicphiloso- 7 is that both of the emerging schools of
phy.
taxonomygot reasonably good startsunThe firstmentionof Hennig in the pages der Editors that one might suspect were
of Systematic
Zoologywas a translationby not especially "sympathetic" to these
Steyskal(1953:41) of Hennig's (1950:4, 10) movements. It is certainly true that Hycontention that the goal of systematicsis man was far from enthusiastic about pato provide a "universal referencesystem" pers with numbersin them,as the followfor biology. With the exception of Kiria- ing comment in her 1961 Editor's report
koff'spapers,Crowson (1965) was the chief indicates:
early defender of Hennig's system.Mayr
One article was rejected because [it was] written
(1965) objected to particular aspects of
in incrediblybad English and another because [it
Hennig's (1950, 1957) system in his criwas] too mathematical.Inquiries among subscribtique of phenetic taxonomy. However, it
ers indicate that the journal has had enough for
was Sokal and Camin (1965) who prethe present of articlesabout numerical taxonomy.
However, furtheropinions on this matterare desented the most sustained criticisms of
sired. An articleof this natureis scheduled forthe
Hennig's system. At the time, Brundin's
March, 1962, issue but thatwill be the last of this
(1966, 1968) exposition of Hennig's prinnature forthe present [SmithsonianArchives].
ciples received littleattention.Even Hennig's (1966) Phylogenetic
seemed
Systematics
However, if one looks at the papers that
not to rouse much immediate response. Hyman published during her final three
Cladistics as a scientificmovement in the years as Editor,neitherquantitativemethUnited States really began with the work ods nor pheneticphilosophyis absentfrom
of Kluge and Farris (1969) and Nelson the pages of Systematic
Zoology.During this

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328

VOL.

ZOOLOGY

SYSTEMATIC

32

Pages

*0 |Brooks
160

I Byers JohnstonlRowelllEldredgelSchuh Smith

Hyman

jaNelson

III

140
120

Pro C/dis tics

F00

III

80I

60III
70IIIII
20

120

Anti Cladistics
80I
100

__

52

54
FIG. 7.

56

58

60

_1

62

64

66

68

70

72

74

76

78

80

82

The number of pages each year arguing foror against cladistic taxonomy.

period, six papers utilizing quantitative


techniquesappeared fora totalof 76 pages.
Three of these papers, however, were not
phenetic. Instead the authors of these papers saw no reason why quantitative
methods could not be used to study evolution and to contributeto evolutionary
systematics.As James(1963) argued, "numerical taxonomy" concerns methodology, not philosophy of systematics.During this same period, nine papers arguing
in favor of phenetics appeared for a total
of 113 pages. As faras percentagesare concerned,these totalsare not high (see Table
4), but if Hyman's percentages indicate
bias, then Rowell's even lower percentage
must indicate even greaterbias. (Perhaps
I can be forgivena personal aside at this
point. One of the papers that Hyman rejected, without review, was submittedby

a philosophy graduate student criticizing


Simpson's definitionof "monophyly." After Simpson's intercessionthe paper was
published [see Hull, 1964].)
As Table 4 also shows, the average percentage of pages arguing the meritsof cladisticshardlychanged forthe 10 yearsbetween 1967 and 1977,hoveringaround 8%.
In general, Figure 7 does not reveal any
majorshiftfromJohnston'slast two years,
throughRowell's tenure,to Eldredge and
Nelson. The major change occurredwhen
Schuh became Editor. During the editorships of Schuh and Smith, cladistics became one of the majorthemesin Systematic
Zoology,surpassing phenetics at its height
under Johnston(see Table 4). The move
fromKansas to the American Museum did
signal a change in the philosophical orientationof the journal's Editors,but it was

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1983

329

ANALYSIS OF SYSTEMATIC ZOOLOGY

Ratios
4.5
4.0
3,5
3.0
2.5

2.0
1.5

1.0
0.5

0
B J R E/NSc Sm
EVOLUTIONARY

SYSTEMATICS

B J R E/NSc Sm
PHENETICS

B J R E/NSc Sm
CLADISTICS

FIG. 8. The ratiosof positive to negative papers foreach of the taxonomicphilosophies under the tenure
as Editor of Byers(B), Johnston(J),Rowell (R), Eldredge and Nelson (E/N), Schuh (Sc) and Smith (Sm).

not accompanied by an immediateshiftin


the distributionsof papers dealing with
the various systematicphilosophies.
Shortof having figuresforrejectionrates
of papers supporting or attacking particular taxonomicphilosophies, the relevant
figuresfor our purposes are the ratios of
positive to negative pages published (see
Fig. 8). Because the issues were not all that
clear in the early years, I have excluded
data for Brooks and Hyman, beginning
with Byers' tenure in office.As Figure 8
shows, more papers supportingevolutionary systematics were published by the
Kansas Editors than opposed. The ratios
were inverted afterthe journal moved to
the American Museum. The one dramatic
featureof the tabulation for phenetics is
the 1.7 ratio under Eldredge and Nelson.
However, the numbers of pages devoted
to phenetics under Eldredge and Nelson
were so low that the ratio does not mean

much (19 pages pro to 11 pages con). Otherwise, the ratios forphenetics were consistentlybelow unity. After a slow start
under Byers,the ratios of positive pages
to n`egativefor the cladists were consistentlyabove one. Once again, the dramatic departure under Eldredge and Nelson
is due in part to small numbers (95 pro to
23 con). The low ratio under Byersis also
not strictlycomparable to the others because the controversyover cladistics was
only just gettingstarted.
Although the preceding figuresdo not
measure directly the matters of greatest
concern,theydo indicate thatthe connections between an Editor's putative biases
and the representationof these views in
the pages of his or her journal are not as
simple as one mightexpect. The ratiosfoi
evolutionary systematicsdropped noticeably when the journal moved to the American Museum, but pheneticists and cla-

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330

SYSTEMATIC

dists alike are opposed to evolutionary


systematics.On the average, the ratios for
phenetics were higherunder the cladists
than they had been when the journal was
at Kansas (0.7 to 0.85)! When allowances
are made for Eldredge and Nelson's artificiallyhigh ratio,the ratio for phenetics
under the cladists is probably somewhat
lower. The ratios for Byers, Rowell and
Smith are not appreciably different.The
ratios for cladists were highest under
Johnston,Eldredgeand Nelson, and Smith;
they dropped a bit under Rowell and
Schuh. Although Schuh was predisposed
to the principles of cladistic analysis and
Rowell was not, the ratios of positive to
negative pages under the tenure of these
two Editorsare roughlythe same. Similarly, although Johnstonand Smith hardly
held the same philosophical preferences,
their ratios for cladistics turned out to be
approximately alike. The philosophical
preferences of Editors may well make a
differenceto a journal, but other factors
are also clearly at work. But one thing is
clear; cladists have had their say in the
pages of SystematicZoology.The average
cumulative ratio forthe last six Editorsof
the journal forthe threepredominantphilosophies of systematicsare evolutionary
systematics (1.03), phenetics (0.78), and
cladistics (1.77).
Anothermeasure of the success of a particularschool of systematicsis the number
of papers that appear which simply take
these principles for granted and apply
themwithoutespecially arguing forthem.
In Figure 9, I have added "applied" papers
to theoretical papers for both phenetics
and cladistics. I have also included for
phenetics all the quantitativepapers that
were not explicitlyhostileto phenetics.For
cladistics,I have added papers arguing in
favor of vicariance biogeography as well
as those applying methods of cladistic
analysis. In each case, I have diagrammed
these data so that the various categories
can be distinguished. The reader should
be warned, however, thatthese figuresare
the most impressionisticof any presented
in this paper.
Given the data recorded in Figure 9,

ZOOLOGY

VOL.

32

pheneticswas at its heightunder Johnston


and Rowell. The drop in theoretical papers thatoccurredunder Rowell was compensated forby papers in applied phenetics as well as papers discussingquantitative
techniques. Included in the latterare most
papers on numerical cladistics. Papers on
applied pheneticsall but disappeared from
Zoologyin the latter
the pages of Systematic
part of Schuh's tenure and early in thatof
Smith. Applied and quantitative papers
reappeared during Smith'sfinaltwo years.
Papers on applied cladisticsand vicariance
biogeographygraduallyincreased through
the years, dropping only in 1981. When
papers that give indirect support to the
pheneticistsand cladists are included, the
representationof the cladistic viewpoint
in recent years becomes even more
marked.
EVOLUTION, PHYLOGENY, BIOGEOGRAPHY
AND NOMENCLATURE

Taxonomy has hardly been the only


Zoology.Just
topic of interestin Systematic
as much space has been devoted to discussing evolutionary theory, phylogeny
reconstruction,biogeographyand nomenclature. In this section I present data for
each of these subjects,distinguishingbetween those papers thatwere largely general discussions of the relevanttheoretical
and methodological issues and those that
were primarilyapplications. Needless to
say, the borderline between "theoretical"
and "applied" in each of these categories
is farfromsharp. In each case, I have also
presented one example of the sort of dispute that took place under each of these
rubrics.
As I understand it, evolutionarytheory
concerns the evolutionary process-how
does evolution occur? A theoreticalpaper
might address such issues as the effectof
gene flowbetween populations on the genetic differencesbetween them. An applied paper mightactuallysample the gene
flowbetween populations, possibly to test
a particularhypothesisabout the effectsof
gene flow. As indicated earlier, the first
majorcontroversyto breakout in the pages
of Systematic
Zoologywas the greatsubspe-

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1983

331

ANALYSIS OF SYSTEMATIC ZOOLOGY

Pages

I
i Pro Clodistics
App//edC/ad/stics I
V/car/nc Biogeography

L5
125
00

l
I

50
25
r-

----1

'-

25I

Rowell

Johnston

Byers

Pages
50

Smith

Schuh

Eldredge a
Nelson

L_

25
0
25
50
70

100II
125II
150

175
200
225

64

65

66

67

ro Phenetics I
App//edPhenetf/cs
uant/tative Methods

.,,

III

I
68

69

70

71

72

73

74

75

76

77

78

79

80

81

82

FIG. 9. Three categoriesof papers are presentedforcladisticsand phenetics: the numberof pages arguing
in favorof cladistics(white above the abscissa), applying cladistic methods (shaded), and arguing forvicariance biogeography(white below the abscissa); and the number of pages arguing forphenetics (white above
the abscissa), applying phenetic methods (shaded), and discussing quantitativemethods (white below the
abscissa).

cies debate. It began with a paper by Wilson and Brown (1953) condemning the
subspecies concept. Although admitting
some problemsexistedwith respectto discerning species, especially in establishing
lower limits and estimatingthe conspecificity of allopatric populations, they
neverthelessconcluded thatMayr'sspecies
conceptis theoreticallywell-groundedand
sufficiently
operational to deserve the centralrole it plays in both taxonomicphilosophy and evolutionarytheory.Wilson and
Brown (1953:100) came to very different
conclusions about the subspecies concept
and its nomenclatural correlate, the trinomen. "From our experience in the literaturewe are convinced that the subspecies concept is the most critical and
disorderlyarea of modern systematicthe-

ory-more so than taxonomistshave realized or theoristshave admitted."


Wilson and Brown(1953) continuedthat,
contraryto the view then current,populations are not characterized by a single
predominantgenome, forminga coadapted system.In point of fact,they claimed,
gene distributionsare not always concordant with character distributions. How
many and which groups are recognized as
subspecies depends on how many and
they
which traitsare studied. Furthermore,
argued, various rules about the frequency
with which traitsmust be correlated(e.g.,
the 75% rule) are arbitraryand subjective,
as they must be because the boundaries
between infraspecificgroups are themselves arbitraryand subjective. However,
even when infraspecificgroups are dis-

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332

SYSTEMATIC ZOOLOGY

crete and homogeneous, Wilson and


Brown (1953) saw no reason to burden the
literature with officialLatin names. Instead,significantinfraspecific
groupscould
be designated by their specific name followed by ecological informationand their
locale. That would designate them sufficiently.
The biologistswho responded to Wilson
and Brown's paper tended to agree that
the nomenclaturalcode was at best a necessary evil and that subspecies posed
problemsdifferent
in kind fromthose presented by species. The few biologists who
defended the use of trinomials did so in
terms of the importance of infraspecific
variation, the need for identifying this
variation, and particular weaknesses of
Wilson and Brown'salternative.Both sides
of the dispute agreed that the biological
phenomena are what really count and not
the names. Burt (1954:99), for example,
noted thatthus far150 subspecies of pocket gophers had been named by energetic
mammalogists."What does all this show
fromthe biological point of view? It shows
that Thomomysbottae is highly variable
throughoutits range. But is it necessaryto
have 150 or more names in the literature
to show this? I think not."
The problem was how to indicate the
sortof statisticalcovariationof traitscharacteristicof both intra- and interspecific
variation. Giving a name to a taxon seems
to imply that the group is homogeneous
and discrete. Hence, naming subspecies
tends to obscure the variation that occurs
within and between subspecies, implying
that each subspecies is homogeneous;
while naming only species tends to obscure the infraspecificvariation. It seems
as ifyou are damned ifyou do and damned
if you don't. However, as the controversy
progressed, systematistspointed out that
the same problemsexistat the species level
as well. If one were to stop naming subspecies simply because of the lack of perfect concordance in defining traits,then
one would also have to stop naming
species. Fox (1955), for instance, claimed
that the organisms that he studied (Ithomiidae) could be divided neatlyinto sub-

VOL. 32

species with no discordance of characters.


Gillham (1956), to the contrary,found the
discordanceclaimed by Wilson and Brown
in thissame group. The differencewas that
Fox had studied museumcollections,while
Gillham's data resulted from extensive
samplings of naturalpopulations. Without
ever being resolved,littleby littlethe subspecies controversy dropped from the
pages of Systematic
Zoology;for a review,
see Pimentel (1959).
However, without the subspecies controversy,discussions of the evolutionary
process (not to mentiontaxonomicphilosophy) would have played even less of a
role in the early years of the journal than
it did. As Figure 10 and Table 5 show, general discussions of the evolutionary process did especially well under Brooks,
Byersand Smithand especially poorly under Johnston,Rowell, and Eldredge and
Nelson. When figuresfor theoretical papers are combined with more applied
work, only Hyman comes out scoring noticeablybelow average.
Discussions of phylogeny reconstruction are frequentlyconfused with discussions of "evolutionarytheory"on the one
side, and with systematicphilosophy on
the other. Without the evolutionary process, therewould be no product-phylogeny-but one can reconstructphylogeny
without opting on most of the issues that
exercise evolutionarybiologists. Is reproductive isolation an adaptation or merely
an effect?How can the 50% cost of meiosis
be made up in order to explain the prevalence of sexual reproduction? The answers to questions such as these are
fundamentallyirrelevant to the paleontologist's task of reconstructingphylogeny. Some questions, however, are relevant
(e.g., punctuational versus more gradualistic modes of speciation). Similarly,classificationis necessary for phylogeny reconstructionbecause paleontologistsmust
have their groups, but they can reconstruct phylogeny without presenting
higher classifications.Differentpaleontologists can agree on a particular reconstruction and yet disagree violently on
how to classifyit. Thus, I think thatthere

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1983

333

ANALYSIS OF SYSTEMATIC ZOOLOGY

Pages

Brooks

160

1610

Hyman

I
I Byers I JohnstonI Rowell aNelson
Nelsong Schuh I Smith

120
120

100 Evolutionary Theory Iheor


I
II
~~~~~~~~~I
80IIIII

IIIII

40
20
0

5 5

20

60II

80IIII
120

Applied EvolutionaryTheoryII

120IIIIIII

160

52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82

FIG. 10. The number of pages each year discussing evolutionarytheory(white above the abscissa) and
applying evolutionarytheory(shaded below the abscissa).

are excellent reasons forcounting phylogeny reconstructionas an independent category.


The chief question with respectto phylogeny reconstructionhas always been its
feasibility.Throughoutthe historyof SystematicZoology,a series of objections have
been raised to it,startingwith Blackwelder, Bigelow, Boyden and Borgmeier,continuing with the pheneticists,and culminating with at least some cladists. In this
connection, the distinction between

cladograms and trees is useful. Cladograms purportto depict only cladistic relations(sister-grouprelations).They do not
indicate which species are actually descended from which, how much divergence in characters has taken place between speciation events, and a host of
other evolutionary relationships. If one
wishes to depict these relationships,the
resultsare trees.Both cladogramsand trees
are branching diagrams, but their implications and the evidence necessary for

TABLE 5. Percentage of pages during the tenure of each Editordevoted to evolutionarytheory,phylogeny


reconstruction,biogeography,and nomenclature.
Evolution
Editor

Brooks
Hyman
Byers
Johnston
Rowell
Eldredge and Nelson
Schuh
Smith

Phylogeny

Biogeography

Nomen-

Theory

Applied

Theory

Applied

Theory

Applied

clature

17
7
12
2
4
3
6
18

12
2
7
15
15
18
11
5

6
10
10
12
10
8
17
15

18
9
8
6
13
17
7
10

0
1
5
2
2
7
6
5

9
18
19
12
8
11
7
5

10
7
4
4
2
1
0
0

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334

SYSTEMATIC ZOOLOGY

Pages

Brooks

140

Hyman

Byers I JohnstonNelsonI
IRowell Ielson:

Theory of Phy/ogeny
I
Reconstruction
100IIII

I
I

120

80

VOL. 32

III
I
I

Schuh I Smith

I
I
II

60

40

20II
0
20

60
80

App/liedPhylaogenyI
I
Reconstruct/on

10o
10

Ii

140
160

52

54

56

58

60

62

64

66

68

70

72

74

76

78

80

82

FIG. 11. The number of pages each year discussing methods of phylogeny reconstruction(white above
the abscissa) and presentingactual reconstructions(shaded below the abscissa).

theirconstructionare quite different.Nelson coined the term "arboristics"for the


science of tree construction, retaining
"cladistics" for the constructionof cladograms(Nelson and Platnick,1981:265).The
termis not intended to be complimentary.
As Figure 11 and Table 5 indicate, papers dealing with phylogeny reconstruction, both theoretical and applied, have
remained fairly constant, occupying between 20% and 25% of the pages of the
journal. The significantamount of work
on the theory of phylogeny reconstruction under Schuh and Smith was primarily numerical cladistics. The only surprising featureof the figures,for me at least,
is the high percentageof pages devoted to
setting out particular phylogenies while

Eldredge and Nelson were Editors.At the


time, both men were extremelyskeptical
of our abilityto inferanythingabout phylogeny save sister-grouprelations.For example, prior to becoming Editor, Nelson
(1972:368) can be found concluding that
"Hennigian relationships embody the
principles that all common ancestral
species are necessarily hypothetical,and
that ancestral species, although they may
be reconstructed(e.g., Fitch, 1971), will
forever remain unknown and unknowable in a directlyempirical sense (e.g., in
the sense that species are 'known' by way
of inference from observation of study
material)."
Eldredge (1979:169), several years later
when commenting on his own earlier

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1983
Pages

Brooks

80 _
60

40
20
20

335

ANALYSIS OF SYSTEMATIC ZOOLOGY

I Byers IJohnston IRowellIEldredgelSchuh ISmith

Hyman

Biogeogrophic

h
er
7z

nI

Nelsoni

20

40
60
80

App/lied I
BiogeographyII

I
I

100IIII
120

52

54

56

Ij

58

60

62

64

66

68

70

72

74

76

78

80

82

FIG. 12. The number of pages each year discussing biogeographic theory(white above the abscissa) and
presentingactual biogeographic distributions(shaded below the abscissa).

views, stated,"I no longer oppose the constructionof phylogenetic trees out-right,


or for that matter,scenarios (which are,
afterall, the most fun), but merely point
out that, in moving through the more
complex levels, we inevitablybecome further removed fromthe original data base
in adding assumptions and ad hoc (and
largelyuntestable)hypotheses."Partof the
explanation forthe extensive data on particular phylogenies published while Nelson and Eldredge were Editorsis thattheir
final number in 1976 was given over entirely to a symposium on marine mammals. Even so, it says somethingabout the
effectof an Editor-s own views on a topic
that Eldredge and Nelson published so
many phylogenetic reconstructionswhen
they themselves were so skeptical concerning the general validity of the activity.
Under "applied biogeography," I in-

cluded all papers thatdealt with particular


distributionsof organismsacross space regardlessof the extentof the territory.
Some
papers concerned single states; others entirecontinentsor seas. One thingthatboth
Table 5 and Figure 12 amply demonstrate
is that,formost of its history,most of the
papers in SystematicZoologyon biogeography were applied. Not until Eldredge
and Nelson took over did the theory of
biogeography receive much attention.As
I mentioned earlier, most of this theoretical discussionconcernedthe strengthsand
weaknesses of vicariance biogeography
and its connection to cladistics. Early on,
advocates of vicariance biogeography introducedit as originatingwith Croizat and
as being very closely connected to the
principlesof cladisticanalysis.For his part,
Hennig never cited Croizat,while Croizat
(1982) publicly rejected cladistic analysis
and complained that vicariance biogeog-

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336

SYSTEMATIC ZOOLOGY

raphy differedsignificantlyfromhis own


panbiogeography.
Although issues of nomenclaturewere
not included explicitlyin any of the statements of purpose of SystematicZoology,
theyhave figuredin the pages of the journal throughout most of its existence. By
"issues of nomenclature,"I do not mean
the continuing debates about appropriate
definitionsfor such terms as "monophyly," "homology" and "phylogenetic," but
disputes over words like "paratype" and
the correct names of taxonomic groups.
The "Bryozoan Affair"is typical of these
disputes. Fromthe middle of the 19thcentury onward, a "battle raged," chiefly
among British systematists, whether a
group of coelomates should be termed
"Polyzoa" or "Bryozoa." When Ehrenberg
established the phylum Bryozoa in 1831,
he included only fourgenera. Later three
of these genera were removed to RotatoMuch later,a
ria, leaving only Zoobotryon.
pseudo-coelomategroup of entoproctswas
included in the phylum and then removed. Upon removal of the entoprocts,
Hyman (1959) abandoned the name
"Bryozoa" and replaced it with "Ectoprocta," termingthe removed entoprocts"Entoprocta."
In a short point of view, Schopf (1967)
recommended the adoption of Hyman's
terminology.If anyone insisted on lumping the ectoproctsand entoproctstogether,the term"Bryozoa" should apply to this
combined group only and not to the ectoproctsalone. Mayr (1968) objected that
the term"Bryozoa" should be retained in
its original usage because Schopf's suggestion violated a very basic principle of nomenclature (e.g., the name of a taxon
should not be changed when an alien elementis removed). Schopf (1968) remained
unconvinced by Mayr's arguments.Soule
and Soule (1969) in their response to
Schopf remarked that they hoped that a
new "Hundred Years War" over "Bryozoa"fversus "Ectoprocta"was not about to
replace the old one over "Polyzoa" versus
"Bryozoa." Cuffey(1969) entered into the
frayto support Schopf, while Beattyand
Blackwelder (1974) estimated that if
Schopf's conventionwere adopted, rough-

VOL.

32

ly one-third of the names of widely accepted phyla would have to be changed.


Finally, Ghiselin (1977) used this controversy to set out a tentativelist of principles to govern changing the names of
higher taxa. From what I can tell, the Hyman-Schopf suggestion has not been
widely adopted. The one featureof the totals forpages devoted to nomenclaturalisZoologyworth noting is
sues in Systematic
theirgradual elimination (see Table 5). In
the past five years, only one short paper
on nomenclature has appeared (Gloyd,
1982).
BOOK REVIEWS

ZoDuring the firstdecade of Systematic


ology,book reviews were not especially
significant.They were short, infrequent
and either positive or gently critical.Under Brooks and Hyman, 16 book reviews
were published totallingonly 35 pages. In
his review of Mayr et al. (1953), Hubbs
(1953:93) complained that the "use of calculatingmachines,standardin some fields,
is not mentioned." Blackwelder's (1961)
review of Simpson (1961) was critical,but
also very respectful.The number of reviews began to increase slowly afterthe
journal moved to the Universityof Kansas-79 reviews totalling131 pages during
a 10-yearperiod. Even so, the pages devoted to reviews never rose above 3% (see
Fig. 1). The mix of positive to negative reviews remained largely unchanged during this period, but several of the negative
reviewsbecame somewhatsharperin tone.
For example,Ross (1964:108) concluded his
review of Sokal and Sneath (1963) with his
"considered opinion that numerical taxonomy is an excursioninto futility,"while
I (Hull, 1969:240) ended my review of a
book on philosophy of biology by a Jesuit
priest,Giovanni Blandino (1969), with the
remarkthatthe Catholic Churchwas about
to reopen the Galileo affair."Anyone who
thinksthat the outcome of these proceedings has any relevance to contemporar.y
science will probably find Theorieson the
NatureofLifestimulatingand timely."
When Eldredge and Nelson took over
editorship in 1974, they added a Review

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1983

ANALYSIS OF SYSTEMATIC ZOOLOGY

Editor, and thereafterthe number and


length of reviews began to climb, taking
up 7% of the pages of the journal under
Eldredge and Nelson, 10% under Schuh,
and 11% under Smith (see Table 1). In
point of fact,this increase in the review
section was greater than it appears because Eldredge and Nelson decreased the
size of type used for reviews beginning
with the second issue in 1974, increasing
the average number of words per page by
50%. Michael Ghiselin was the firstReview Editor, followed by Norman Platnick. As in the case of "Editors-in-Chief,"
the borderlinesbetween Review Editorsis
not as sharp in practiceas the masthead of
the journal might imply. Ghiselin inherited several reviews from Rowell, and
Platnick's name appeared as the official
Review Editor two issues prior to his actually taking over. That way publishers
would send books to the incomingReview
Editor. Ghiselin was Review Editor for
three years between 1974 and 1976, while
Platnick served in the post for six years
between 1977 and 1982.
Prior to the introductionof a Review
Editor,the Editor of the journal authored
an occasional review: Brooks (2), Hyman
(0), Byers(1), Johnston(2), and Rowell (1).
In each case, these reviews were positive.
While Nelson was Editor,he published 28
reviews, of which 11 were basically de.scriptive,8 were largely favorable, and 9
were critical.During this same period, Eldredge authored two reviews, one critical
and one mixed. While Schuh was Editor,
he authoredthreereviews,two criticaland
one mixed. During his stint as Editor,
Smithcontributedno book reviews. While
Ghiselin was Review Editor,he reviewed
seven books, all favorably.During his six
years as Review Editor,Platnick reviewed
17 books, 10 of themwith Eugene Gaffney
on the philosophy of Popper. Numerous
Zoology
papers also appeared in Systematic
during this period that argued the relevance of Popper's philosophical views to
taxonomy. Of the reviews that Platnick
authored by himself, three were favorable, threeunfavorable,and one verycritical.
As Johnston(1969:361) statedduring his

337

tenure as Editor, book reviews are handled differentlyfrom articles and points
of view, noting that"within the generally
accepted limitsof good taste,a review is a
privileged communication. Reviews are
exempt from editorial tampering,and in
Systematic
Zoologya reviewer is essentially
at libertyto say what he wishes." The occasion for these comments was a critical
review by Rising (1969) of Blackwelder's
(1967) text. Three readers had writtento
the Editor defending alpha taxonomy of
the sortadvocated by Blackwelder against
Rising's "one-sided" review. Johnston
published the lettersbecause he had been
well aware of the sort of review that Rising was likely to give Blackwelder's book
when he asked him to review it. Although
Editors cannot "tamper" with reviews,
they are in a position to select reviewers
whose past record indicates the sortof review thattheyare likelyto write.One way
around this difficulty
is to have a book reviewed by two authors,one who is likely
to be favorablypredisposed to the book,
one who is not. This practice was begun
under the editorshipof Eldredge and Nelson for Sneath and Sokal (1973). At first
Ghiselin had attemptedto get one person
representingeach of the threemostprominent taxonomic philosophies to review
the book but eventually settled fortwoW. W. Moss and DieterSchlee. Otherbooks
given dual reviews were Lovtrup 1977),
Boudreaux (1979), Eldredge and Cracraft
(1979), Wiley (1981), and Nelson and Platnick (1981). The only book forwhich this
treatmentdid not have the intended effect
was Boudreaux (1979). Neither reviewer
liked the book.
As Table 6 indicates, the book review
section of Systematic
Zoologyhas not only
increased in size but also become more polemical. Prior to Platnick taking over as
Review Editor, I judged only three reviews to be extremely"harsh"-12 thereafter.However, as Table 6 indicates,these
12 verynegative reviews make up only 8%
of the totalnumber of reviews under Platnick. Opinions as to the value of polemics
and the propriety of highly critical reviews vary. Justas children are very difficultto raise (even badly), writinga book,

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338

VOL.

SYSTEMATIC ZOOLOGY

32

TABLE 6. Number of book reviews and percentage of pages during the tenure of each Editor or Review
Editor categorized according to the characterof the review.
Editor
(years)

Brooks(6)
Hyman (6)
Byers(3)
Johnston(4)
Rowell (3)
Ghiselin (3)
Platnick (6)

No.
reviews

8
8
12
42
25
70
142

Positive

_ _ _ _ _ _ _ _ _ _

Mixed or neutral

Negative

Very negative

No.

Percent

No.

Percent

No.

Percent

No.

Percent

5
4
9
26
17
33
69

63
50
75
62
68
47
49

1
1
0
10
5
19
22

13
13
0
24
20
27
16

2
3
1
5
3
18
39

25
38
8
12
12
26
27

0
0
2
1
0
0
12

0
0
17
2
0
0
8

no matterhow poor others might find it,


is a lot of work. Aftersuch labors, an authoris not likelyto take kindlyto negative
evaluations of his or her brainchild. For
my own part,I findoverlydeferent,largely descriptive reviews not very useful. If
the reviewerfindsa book eitherverygood
or verybad, he or she should say so. However, the fact that book reviews do not
undergo the same refereeingprocess that
other publications do should inhibit gratuitous remarksat the author's expense.
Unfortunately,I am in no position to wax
too moralisticallyon this point because 2
of the 15 reviews that I judged very negative were writtenby me. During thissame
period, several authors began to use reviews as an opportunityto write an essay
of their own choosing in the manner of
the New YorkReviewof Books.The continued popularity of this latter publication
indicates that opinions as to the value of
such "book reviews" also vary.
CONCLUSION

In the preceding pages I have attempted


to characterizethe contents of Systematic
Zoologyas this journal has been presented
to its readers. Whether warranted or not,
many readers have gathered the impression that the pheneticists "owned" the
journal while it was at Kansas, while the
cladistswere in controlwhile it was at the
American Museum and then the Los Angeles County Museum. The data do not
supportthis easy conclusion. The increase
in the number of pages devoted to dis-

cussing phenetics began under Hyman,


reached its peak under Johnston,and then
began to fall under Rowell-while the
journal was still at Kansas. The decline
continued under Eldredge and Nelson but
then was reversedunder Schuh and Smith.
While the journal was at Kansas, more papers critical of phenetics were published
than in support,and these figuresdid not
change all thatmuch once the journal left.
If "cladist editors" were harder on papers
supporting phenetics, the effectsof this
putative bias are not apparent. The only
significantdecrease under the cladistswas
in pages devoted to applications of phenetic methods.
Cladistics got a good start while the
journal was at Kansas. None of the relevant figureschange significantlyafterthe
journal moved to the American Museum
while Eldredge and Nelson were Editors.
FromJohnstonuntil the present,more papers in favor of cladistics have been published than those opposed. Changes in the
taxonomic allegiances of the Editors
seemed to have made no difference.However, under Schuh and Smith, the percentage of pages devoted to cladisticstook
a sharp jump.
One peculiar feature of the data presented in this paper concerns evolutionary systematics.While the journal was at
Kansas, the ratio of positive to negative
papers was quite high. When the journal
moved to the American Museum, this ratio dropped sharplyand remainedlow. But
these figuresare quite puzzling, since the

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1983

339

ANALYSIS OF SYSTEMATIC ZOOLOGY

principles of phenetic taxonomyand cladistic analysis are equally incompatible


with those of evolutionary systematics.
Editorial bias, like God, seems to move in
strangeways.
My own impression is that convictions
about bias stem from the treatmentof a
relatively few papers, so few that the effectsof this putative bias are not likely to
appear in summaryfigures.However, in
suggesting that editorial bias may well
have been less common than it may seem,
I do not intend to diminish its importance
in individual cases. A victim of a miscarriage of justice is not in the least impressed by the observationthat such miscarriages are probably quite rare, and
justifiablyso. If the controversiesin SystematicZoologyare any indication, scientific disputes seem to have a rhythmof
their own-attacks elicit defenses and
counterattacks, which produce another
round, all with lag times of from one to
two years.
In short, casual impressions gathered
over many years can be deceptive. For example, I had not noticed the drop in discussions of taxonomic theory under Eldredge and Nelson, nor the decrease in
applied phenetics under Schuh, nor the
near total elimination of papers on nomenclature after the journal returned to
the American Museum. These were the
markedchanges. Othersmay have noticed
them; I did not. I am also struckby how
informallychanges of Editors occurred in
the early years of the journal. The officers
got togetheror talked over the telephone,
a few inquiries were made, and a new Editor was appointed. This informalsystem
ceased with Smith. Both Smith and the
current Editor, Gary D. Schnell, became
Editorunder verydifferentcircumstances.
Finally,there is some evidence as to the
impression that the readers have of SystematicZoology,at least for 1980. In the
previous year, a committee was formed
consistingof Richard F. Johnston,Randall
T. Schuh, James Dale Smith, and Everett
C. Olson to constructa questionnaire to
send to the membersof the Society to see

how well the Society and its journal were


fulfillingtheirneeds; 418 replies were received. The important question for our
purposes concerned the aims of the journal. "Systematic Zoology is devoted to
publication of resultsof researchprojects,
serves as an open forumforpoints of view
on systematics,is a medium forreview of
pertinentbooks. Do you agree that these
prescribedaims are appropriate?"Slightly
over 90% of those who responded marked
"Yes." From writtencomments,however,
the committeeconcluded thatsome of the
respondents interpretedthe question literally-do you agree with these aims?
Others were responding to how well they
thought the journal was fulfillingthese
aims. In general, 61% of the answers were
favorableand 39% unfavorable(Syst.Zool.,
30:224, 1981).
ACKNOWLEDGMENTS
I am grateful to the following people for commenting on an earlier version of this paper: George
Byers,Joel Cracraft,Steve Farris,Vickie Funk, Mike
Ghiselin, Richard Johnston,ErnstMayr, Norm Platnick, JimRohlf, Bert Rowell, Toby Schuh and Bob
Sokal. The paper is much betterfor their criticisms
and suggestions. I would also like to thank Berndt
Ehman forproducing the diagrams forthis paper.
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Received5 May 1983; accepted24 June1983.

341

ZOOLOGY

APPENDIX

Categorization
ofPapersin Vol. 24
(1975) ofSystematic
Zoology
Number 1.-Maxson and Wilson(15 pages)-largely descriptive;test of concordance between organismal and proteinevolution; phylogenyreconstruction
and the evolutionary process; some quantification;
historical biogeography. Fouquette(8)-descriptive;
isolating mechanisms in a group of frogs; applied
evolutionary theory; local geographic distribution.
Waage (13)-descriptive; reproductiveisolation in a
group of damselflies; applied evolutionary theory;
some geographic distribution.Dodson (18)-descriptive; the classification of some hadrosaurs using
biometryof skulls; applied phylogeny. Coombs(8)descriptive; classification of Chalicotheres using
biometryof radius and tibia lengths. Zimmerman
and
Ludwig(9)-descriptive; quantitative; biogeography
of the Dytisidae; applied biogeography;applied evolutionary theory. Por (7)-descriptive; applied biogeography; applied phylogeny reconstruction.Robinsonand Hoffman
(10)-descriptive; numerical study
of some ground squirrels; applied evolutionarytheory;local distributions.Caireand Zimmerman
(7)-descriptive;chromosomalvariation in the deer mouse;
applied evolutionarytheory;local distributions.Lovtrup (13)-phylogeny reconstructionand implicationsforclassification;
applied cladistics.Sabrosky
(2)nomenclature. Robinson(1)-nomenclature. Packard
and Stiverson
(2)-descriptive; geographicvariationin
a species of spider. Bretsky
(6)-evolutionary theory;
phylogeny reconstruction; anti-cladistic analysis.
Cracraft(1)-phylogeny reconstruction;pro-cladistic
analysis.
Number 2.-Baker, Bleierand Atchley(10)-descriptive;chromosomestudyof contactzones between taxa
of Urodema;applied evolutionary theory. Kilpatrick
and Zimmerman
(20)-descriptive; chromosomestudy
of four species of mouse; applied evolutionarytheory. Rauschand Rausch(8)-descriptive; chromosome
study of the red-backed vole; applied evolutionary
theory.Jannett
(5)-descriptive; hip glands in voles;
applied phylogeny reconstruction.Leamy (15)-descriptive;biometry;osteometryof mice; applied evoand Kistner(8)-numerical
lutionarytheory.Jacobson
analysis of a group; applied phenetics.Moss and Power (10)-pro-phenetics; using machines to record
morphometricdata. Read (13)-theory and applied;
quantitative;comparisonofphylogeniesinferredfrom
fossil data and albumin. Van Valen (1)-problems in
publication; issues relevant to the profession. Matthews(2)-issues relevant to the profession. Schuh
(2)-response to Matthews. Corruccini
(7)-phylogeny reconstruction.Wiley (11)-pro-cladistics; antievolutionary taxonomy; Popper. Hennig (13)-procladistics; anti-evolutionarytaxonomy. Sokal (6)mixed defense of evolutionary taxonomy; anti-cladistics;implicitdefense of phenetics.

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342

SYSTEMATIC ZOOLOGY

Number 3.-Case, Hanelineand Smith(15)-descriptive; protein variation in several species of Hyla; applied evolutionary theory. Patton,Yang and Myers
(15)-descriptive; divergence of rats on the Galapagos; applied evolutionarytheory. Sneath,Sackinand
Ambler(22)-descriptive; applied evolutionary theory;quantitative;applied phenetics.Huffman
andaBullock(13)-descriptive; application of a graph method
to Acanthocephala. Kohn and Riggs(14)-morphometry of shells; descriptive and quantitative. Sneath
(8)-detection and representationof reticulaterela-

VOL. 32

tions; anti-cladistics.Gormanand Kim (4)-descriptive; genetic variation of some lizards; applied evolutionarytheory. Sharrockand Felsenstein(5)-quantitative methods for finding monothetic sets;
quantitative.Stunkard(8)-descriptive; systematicsof
some flatworms.
Number 4.-Ball (24)-biogeographic theory; applied biogeography. Rosen (34)-biogeographic theory; applied biogeography; pro-vicariancebiogeography. Vagvolgyi(24)-applied biogeography with
some theory.

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