Title Page
 Table of Contents
 Scientists and social progress
 Masculinity-femininity responses...
 A study of the dragonflies of the...
 Outline of the geological history...
 Addenda to the list of birds of...
 Notes on the breeding habits of...
 A diagnostic taxonomic constant...
 The vapor phase oxidation of alpha...
 Catalytic hydrogenation of oleoresin...
 Dehydroabietic acid and certain...
 Florida pocket-gopher burrows and...
 A brief history of tomato production...
 A marine biological laboratory...
 Biological effects of radiations...
 The scientific status of fish culture...
 The effects of immersion on the...
 The shark fishing industry...
 A preliminary note on the Vitamin...
 The reaction of magnolia, scrub...
 The development of the social sciences...
 The flat-tailed water snake
 Preliminary list of myxomycetes...
 Geography and the social scien...
 Studies of foraminifera from seven...
 Tests for determining prime...
 What science should be taught children...
 The monarch butterfly (danaus menippe...
 Cooperation of work projects administration...
 Mechanics and sponsorship of W.P.A....
 The future of Florida archeological...
 Academy business and personnel

Title: Proceedings of the Florida Academy of Sciences
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00001490/00004
 Material Information
Title: Proceedings of the Florida Academy of Sciences
Abbreviated Title: Proc. Fla. Acad. Sci.
Physical Description: 7 v. : ; 23 cm.
Language: English
Creator: Florida Academy of Sciences
Publisher: Rose Printing Co., etc.
Place of Publication: Tallahassee
Frequency: annual
Subject: Science -- Periodicals   ( lcsh )
Genre: periodical   ( marcgt )
Dates or Sequential Designation: v. 1-7; 1936-44.
 Record Information
Bibliographic ID: UF00001490
Volume ID: VID00004
Source Institution: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: aleph - 001745383
oclc - 01385276
notis - AJF8161
lccn - sn 85003387
issn - 0097-0581
 Related Items
Succeeded by: Quarterly journal of the Florida Academy of Sciences

Table of Contents
    Title Page
        Page i
        Page ii
    Table of Contents
        Page iii
        Page iv
    Scientists and social progress
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
    Masculinity-femininity responses of Florida state college for women and University of Florida students
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
    A study of the dragonflies of the genus progomphus (gomphoides) with a description of a new species
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
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        Page 80
        Page 81
        Page 82
        Page 83
        Page 84
        Page 85
        Page 86
    Outline of the geological history of Peninsular Florida
        Page 87
        Page 88
        Page 89
        Page 90
        Page 91
        Page 92
        Page 93
        Page 94
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        Page 98
        Page 99
        Page 100
        Page 101
        Page 102
        Page 103
        Page 104
        Page 105
    Addenda to the list of birds of Alachua County, Florida
        Page 106
        Page 107
    Notes on the breeding habits of the warmouth bass
        Page 108
        Page 109
        Page 110
        Page 111
        Page 112
    A diagnostic taxonomic constant for separating slash and longleaf pines
        Page 113
        Page 114
        Page 115
    The vapor phase oxidation of alpha pinene
        Page 116
        Page 117
        Page 118
        Page 119
    Catalytic hydrogenation of oleoresin from the slash pine (pinus caribaea)
        Page 120
        Page 121
        Page 122
        Page 123
    Dehydroabietic acid and certain of its derivatives
        Page 124
        Page 125
        Page 126
    Florida pocket-gopher burrows and their arthropod inhabitants
        Page 127
        Page 128
        Page 129
        Page 130
        Page 131
        Page 132
        Page 133
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        Page 164
        Page 165
        Page 166
    A brief history of tomato production in Florida
        Page 167
        Page 168
        Page 169
        Page 170
        Page 171
        Page 172
        Page 173
        Page 174
    A marine biological laboratory on the Gulf Coast of Florida
        Page 175
        Page 176
        Page 176a
        Page 177
        Page 178
    Biological effects of radiations of different wave-lengths
        Page 179
        Page 180
        Page 181
    The scientific status of fish culture and lake management in Florida
        Page 182
        Page 183
        Page 184
        Page 185
    The effects of immersion on the hemograms of swimmers
        Page 186
        Page 187
        Page 188
    The shark fishing industry of Florida
        Page 189
        Page 190
        Page 191
        Page 192
    A preliminary note on the Vitamin A content of the liver oil of the Florida lemon shark (hypoprion brevirostris poey)
        Page 193
        Page 194
    The reaction of magnolia, scrub live-oak, slash-pine, palmetto and other plants to dune activity on the Western Florida Coast
        Page 195
        Page 196
        Page 197
        Page 198
        Page 199
        Page 200
        Page 201
        Page 202
        Page 203
    The development of the social sciences in the Southeast
        Page 204
        Page 205
        Page 206
        Page 207
        Page 208
        Page 209
    The flat-tailed water snake
        Page 210
        Page 211
    Preliminary list of myxomycetes from Alachua County
        Page 212
        Page 213
        Page 214
        Page 215
        Page 216
        Page 217
    Geography and the social sciences
        Page 218
        Page 219
        Page 220
        Page 221
        Page 222
        Page 223
        Page 224
    Studies of foraminifera from seven stations in the vicinity of Biscayne Bay
        Page 225
        Page 226
        Page 227
        Page 228
        Page 229
        Page 230
    Tests for determining prime factors
        Page 231
        Page 232
        Page 233
        Page 234
        Page 235
        Page 236
        Page 237
        Page 238
        Page 239
        Page 240
        Page 241
        Page 242
        Page 243
        Page 244
        Page 245
    What science should be taught children of Florida? Methods of investigating this problem
        Page 246
        Page 247
        Page 248
        Page 249
        Page 250
        Page 251
    The monarch butterfly (danaus menippe hub.) in Florida
        Page 252
        Page 253
        Page 254
    Cooperation of work projects administration with university and college research programs
        Page 255
        Page 256
        Page 257
        Page 258
        Page 259
        Page 260
        Page 261
        Page 262
    Mechanics and sponsorship of W.P.A. projects
        Page 263
        Page 264
        Page 265
    The future of Florida archeological research
        Page 266
        Page 267
        Page 268
        Page 269
        Page 270
        Page 271
        Page 272
        Page 273
        Page 274
        Page 275
        Page 276
        Page 277
        Page 278
        Page 279
        Page 280
        Page 281
        Page 282
        Page 283
        Page 284
        Page 285
        Page 286
    Academy business and personnel
        Page 287
        Page 288
        Page 289
        Page 290
        Page 291
        Page 292
        Page 293
        Page 294
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Full Text


of the

Florida Academy of Sciences




Published by the Academy, Gainesville, Florida
August, 1940


Published Annually by the Academy

Editor: L. Y. DYRENFORTH, St. Luke's Hospital, Jacksonville
Managing Editor: J. H. KUSNER, University of Florida
Editorial Committee:
R. S. ATwOOD, University of Florida
A. F. CAnR, University of Florida
EZDA DEVINEY, Florida State College for Women
R. S. JOHNSOn, University of Florida
E. M. MILLER, University of Miami
MAURICE MULVANIA, Florida Southern College
P. W. PENNINGROTH, St. Petersburg Junior College
B. P. REINSCH, Florida Southern College
S. A. STUBBS. Florida Geological Survey
FRANCES WEST, St. Petersburg Junior College
R. C. WIULIAMSON, University of Florida
Editorial assistance in connection with this volume was also rendered by:
Lillian Arnold, University of Florida; Shirley J. Case, Florida Southern Col-
lege; M. J. Dauer, University of Florida; U. P. Davis, University of Florida; B. F.
Dostal, University of Florida; J. G. Eldridge, University of Florida; Ralph Greene,
Coral Gables; T. H. Hubbell, University of Florida; J. M. Leake, University of
Florida; J. F. W. Pearson, University of Miami.
The Academy makes grateful acknowledgement of the cooperation of the
Florida Writers' Project of the Work Projects Administration in making available
the services of Vera Louise Pritchett as assistant to the Managing Editor.
A paper-bound copy of the Proceedings is sent to each member of
the Academy, without charge. A cloth-bound copy may be obtained,
instead, upon payment of $1.00.
The sale price of the Proceedings is:
Paper-bound-$2.50 per copy
Cloth-bound-$3.50 per copy
The Academy will be pleased to enter into exchange arrangements
with other scientific societies in any field and in any country.
Orders for copies of the Proceedings, subscriptions, exchange pub-
lications, inquiries concerning exchange, and general correspondence
concerning Academy matters should be addressed to:
J. H. KUSNER, Secretary


Scientists and Social Progress.-B. P. REINSCH................. .............................. 1
Masculinity-Feminity Responses of Florida State College for Women and
University of Florida students.-DoROTHY ROSE DISHER........................... 11
A Study of the Dragonflies of the Genus Progomphus (Gomphoides) with
a Description of a New Species.-C. FRANCIS BYERS................................ '.. 19
Outline of the Geological History of Peninsular Florida.-ROBERT B.
CAM PBELL ...................................................................... .................... ......... 87
Addenda to the List of Birds of Alachua County, Florida.-J. C. DICKIN-
SON JR. ........................................................................................................ 106
Notes on the Breeding Habits of the Warmouth Bass.-A. F. CARR, J ............. 108
A Diagnostic Taxonomic Constant for Separating Slash and Longleaf Pines.
W ILBUR B. DE VALL ................................................................................ 113
The Vaper Phase Oxidation of Alpha Pinene.-CHARLEs K. CLARK and
J. ERSKINE HAWKINS .............................. ........................................................ 116
Catalytic Hydrogenation of Oleoresin from the Slash Pine (Pinus Caribaea).
-CHARLES R. STEARNS, JR. and J. ERSKINE HAWKINS .............................. 120
Dehydroabietic Acid and Certain of its Derivatives.-W. DAVID STArLLCP and
J. ERSKINE H AW KINS .................................................. ..................................... 124
Florida Pocket-Gopher Burrows and their Arthropod Inhabitants.-T. H.
H UBBELL and C. C. GOFF ........................................................ ........................ 127
A Brief History of Tomato Production in Florida.-GEoRGE F. WEBER............ 167
A Marine Biological Laboratory on the Gulf Coast of Florida.-A. E.
H OP IN S .................................................................................................................... 175
Biological Effects of Radiation of Different Wave-Lengths.-A. A. BLESS........ 179
The Scientific Status of Fish Culture and Lake Management in Florida.
-0. LLOYD MEEHEAN 2........................................ 182
The Effects of Immersion on the Hemograms of Swimmers.-LucEN Y.
D YRENFORTH ........................................................................................... 186
The Shark Fishing Industry in Florida.-L. L. Ruso ........................................ 189
A Preliminary Note on the Vitamin A Content of the Liver Oil of the
Florida Lemon Shark (Hypoprion Brevirostris Poey).-L. L. RusorT........ 193
The Reaction of Magnolia, Scrub Live-Oak, Slash-Pine, Palmetto and Other
Plants to Dune Activity on the Western Florida Coast.-HIRMAN KURE 195'
The Development of the Social Sciences in the Southeast.-JoHN M. MAC-
LACH LAN .................................................................................................................... 204
The Flat-Tailed Water Snake.-CRAIG PHILLIPS ................................................. 210
Preliminary List of Myxomycetes from Alachua County.-ERDMAN WEST ........ 212
Geography and the Social Sciences.-ROLLIN S. ATWOOD ................................... 218
Studies of Foraminifera from Seven Stations in the Vicinity of Biscayne Bay.
SIDNEY A. STUBBS ................................ ......................................................... 225'
Tests for Determining Prime Factors.-GuY G. BECKNELL .............................. 231
What Science Should be Taught Children of Florida? Methods of Investi-
gating This Problem.-LEo L. BOLES ........................................................... 246
The Monarch Butterfly (Danaus Menippe Hub.) in Florida.-H. T. FERNALD 252



Cooperation of Work Projects Administration with University and College
Research Programs.-MILToN C. FORSTER ..................................................... 255
Mechanics and Sponsorship of W. P. A. Projects.-FRANKLIN E. ALBERT............ 263
The Future of Florida Archeological Research.-SIDNEY A. STUBBS 266
Some Aspects of the Naval Stores Problem.-J. E. HAWKINS, C. K. CLARK,
W D STALLCUP and C. R. STEARNS .................................................................. 271
Ovum and Spermatozoon Age and the Course of Development and Gesta-
tion in the Guinea Pig.-WILAM C. YOUNG ............................................. 271
Superstition and Science in the Field of Mental Illness.-PHILIP WORCHELL.... 273
Raman Spectra of Dimethylenechlorhydrin and Nonamethylenechlorhydrin.
R C. W L IIAM SON .................................................................. 273
The Pteridophytes of Florida.-MARY B. DIDDELL ........................................... 275
Certification Requirements for Teaching Science in the High Schools.-
R OBERT I. A LLEN .............................................................. .............................. 276
The Effect of Cold Storage on Certain Native American Perennial Herbs.
H ERM AN K URZ ................................................................ ............. ................ 276
The Historical Method in the Social Studies.-KATHRYN T. ABBEY.................. 277
Sample Applications of Rectilinear Correlational Techniques and Psycho-
physical Methods of Concomitant Variation to the Concepts of Incidental
Communality and Causal Efficacy.-CHRISTIAN PAUL HEINLEIN............ 279
A Cubic Foot of Evidence.-RAYMOND F. BELLAMY .......................................... 279
Ovoviviparous Mayflies in Florida.-LEwIS BERNER .......................................... 280
Cyclo-Geometric Series.-GuY G. BECKNELL .............................. ........................ 280
Observations on Captive Porpoises.-ARTHUR F. MCBRIDE .............................. 282
Word Dangers in Scientific Thinking.-EARL L. VANCE ...................................... 283
Forest Land Income from Naval Stores in Alachua County, Florida, in 1937.
E. A ZIEGLER .................................................................................................... 283
The Sun-A Component of a Binary System.-J. FRANCES KNORR, JR., and
ARNOLD EDM UND HAYES ..................................................................................... 285
The National Survey of Science Teaching.-W. LEROY MACGOWAN ................ 285
R report of the Secretary ....................................................... ................................ 287
Report of the Treasurer ;........................ ...............288
The Achievement Medal for 1939 ....................................... ..................... 289
Resolution Concerning the Social Sciences ............................................................ 290
Program of the Fourth Annual Meeting ................................................................. 291
Program of the Organization Meeting Junior Academy of Sciences of Florida.... 295
Summary of Attendance at Organization Meeting of Junior Academy.............. 296
Officers of the Junior Academy for 1940 ...................... .................... 296
Officers of the Academy for 1939 .................................................. 297
Officers of the Academy for 1940 ......................... ............. ............. 297
Alphabetical List of M em bers for 1939 .............................................................. 298
Institutional Sustaining Members .................................................. .... 304
Geographical Distribution of Members for 1939 ....................... ................... 305
Charter of the Academy ..................... ................... ..... ................... 307
By-Law s of the Academ y .................................................................. ......................... 308


VOLUME 4 1939

Florida Southern College

In these recent years of economic difficulties and social injustices,
it has become the fashion to direct varied accusations and criticisms
against science. The many applications of scientific knowledge to hu-
man affairs, admittedly valuable in themselves, and tending to give man
control over his environment, are being held responsible for many of
our present-day evils. Here are some of the things they say: Tech-
nological advances produce unemployment. Mechanization of agricul-
ture leads to overproduction. Science and invention produce more and
more ingenious machines of destruction so that a future war will very
likely destroy our civilization. The widespread fruits of science encour-
age an excessive worship of the material and tend to make mankind
spiritually bankrupt. Science should take a ten-year holiday, give
man time to catch up morally and ethically. Science has failed to
solve the pressing problems of human needs and human relationships.
These criticisms have all been aptly summarized by Harold G.
Moulton in his address on "Science and Society" to the American As-
sociation for the Advancement of Science in Indianapolis:1
In what concrete ways is science held to be a menace-a threat to the future of
civilization? At its door various commentators, reflecting upon the undisciplined
progress of the past 100 years, have placed responsibility: for developing an in-
dustrial organization of such vast complexity as to baffle human control; for
creating an international economic structure in a world of political nationalism;
for building implements of warfare which threaten the very extinction of peoples;
for so mechanizing work processes as to dull the qualities of human intelligence;
for changing the relative rates of population growth in the upper and lower strata
of society; for bringing into existence new forms of goods and services in such
rapid succession and in such profusion as to make it difficult for slowly changing
human beings to assimilate them; for giving us leisure that we do not know how
*Address of the retiring president.
1Science, Vol. 87 (Feb. 25, 1938), p. 174.


to use; for producing chronic unemployment and the grave social problems
which it entails; for building up a capacity for production beyond our pow-
ers of consumption; for creating an artificial way of life in place of the old
simplicity; and for distorting ethical values and undermining religion and morals.
Meanwhile, the ambitions of science are not being realized; indeed, there is
a deep feeling of frustration. Applications of new knowledge and inventions to
productive processes are delayed by restrictive business practices and govern-
mental regulations and especially by great economic dislocations which thwart
the profit incentive and at the same time diminish the financial resources required
for continuing research.

To meet these accusations, scientists are naturally becoming quite
vociferous in defense of science. They point out that "Science Makes
Jobs"--witness the automobile, the telephone, the chemical and the
electrical industries. They list the many beneficial effects of scientific
discoveries upon society. They explain that the machines and products
of science can be used either for noble or for nefarious purposes. For
example, the same materials are used to make both gunpowder and fer-
tilizers; even high explosives have many peace-time uses. Science is
knowledge only. It is not concerned with the uses that are made of
this knowledge. The right use is a matter of morality and religion.
The scientist is not responsible for the misuse of his gifts; let society
take care of that.
Much of this defense has really proven to be quite incomplete and
unsatisfying. For example, the argument that science makes jobs isn't
very helpful when we consider that there are still about 10,000,000
unemployed. Science has helped make this a land of plenty; yet, par-
adoxically, millions are in dire need of the bare necessities of life.

Men of science are beginning to bestir themselves and coming to
realize their special responsibilities in the solution of social problems.
For example: The International Council of Scientific Unions in May,
1937, instituted a Committee on Science and its Social Relations
with Prof. J. M. Burgers of Delft, Holland, as Secretary. They are
planning extensive world questionnaires to investigate among other
things: "the part played by scientific thought in the outlook of
various social groups; the forms in which scientific workers and their
work are involved in the various struggles and conflicts of human so-
ciety; the forms in which the consciousness of a social responsibility of
science and of scientific workers is taking shape."2

2Science News Letter, Vol. 34 (Oct. 15, 1938), p. 248.


In December, 1938, scientists in Boston and Cambridge, Mass., or-
ganized3 another branch of the American Association of Scientific
Workers with the purpose of bringing scientific workers together to
promote an understanding of the relationship between science and
social problems.
The aims of this American Association parallel closely those of the
Association of Scientific Workers, organized in Great Britain in 1918.
Their president, Sir F. Gowland Hopkins, past president of the British
Association for the Advancement of Science and of the Royal Society,
wrote a congratulatory letter to the American Association of Scientific
Workers saying: "In these days when science plays so great a part
in every field of modern life it is essential for scientific workers to
organize, both to protect their own economic and professional status
and to work for the better organization and application of science for
the benefit of the community."4
Our American Association for the Advancement of Science has be-
gun with the Indianapolis meeting5 in 1937 the practice of scheduling
many addresses and symposia on this vital problem of the interrela-
tionships of science and society.
The actions of these groups testify to a growing interest in social
problems. Yet I think we will agree that the great majority of our
scientific workers have assumed little responsibility or shown little
interest thus far.
On what grounds should we expect scientists to participate more
actively in the solution of our many human problems: social, eco-
nomic, and political? Scientists have with their discoveries and de-
velopments made the greatest contributions toward creation of the
modern order with its problems. To refuse to play an active part in
the solution of these problems would be tantamount to a betrayal of
the scientific movement. In sheer self-defense scientists must become
The fundamental reason for the scientist's social responsibility lies
in the fact that he lives in a democracy. He is a vital part of it. As a
citizen, with an intelligence high above the average, he has greater
responsibilities than merely voting, accepting majority rule, and under-
standing governmental processes. His own welfare is intimately wrapped
up in the welfare of his country. He needs to keep informed regarding

3Science, Vol. 88 (Dec. 16, 1938), p. 562.
4Science, Vol. 89 (Jan. 20, 1939), p. 58.
5Science, Vol. 87 (Feb. 25, 1938), pp. 173-179.


public issues in local, state, and national affairs, to arrive at intelligent
convictions, and to give vigorous expression to them. He must give
guiding principles to the masses, help them to understand the complex
and confusing issues, and enable them to distinguish between the gen-
uine and the counterfeit. He will need to cooperate actively with
other citizens in carrying on the necessary work of the community and
There is another reason why the scientist has a special responsibility.
That is his peculiar intellectual equipment. His habits of careful and
impartial investigation, his conviction that nature is orderly and un-
derstandable, his habit of subjecting hypotheses to the test of irre-
ducible and stubborn facts, his passion for truth, his unwillingness
to manipulate the truth for the sake of doctrine, his knowledge that
it is impossible to get something for nothing-all these qualities make
him peculiarly valuable in social and economic affairs.
Our country needs the services of that type of man. This thought
was already voiced in a prophetic statement of James Madison: "the
task of adapting institutions and laws to the intellectual, moral, and
social changes to come in 1930 will require for the task all the wisdom
of the wisest patriots."
We should stop for a moment to say what we mean by social
progress. It does not merely denote a change to a new order. Change is
commonly mistaken for progress. Progress requires the new order to be
superior to the old. We might say that progress is that which secures
an increase in human happiness, that which advances toward a more
desirable type of living. We shall now discuss a few specific ways in
which the individual scientist may become a more helpful factor in
social progress.
1. Scientists need to educate the people as to the true character of
their scientific work. Most ordinary persons take the many wonderful
gifts of science for granted. All too few have a realization of the funda-
mental way in which science is responsible for the innumerable good
things in man's life and in man's way of thinking. Think of the many
immediate social benefits of researches in soil chemistry, food chemistry,
horticulture, entomology, bacteriology, metallurgy and the like.
The scientist should pass along to the people information concern-
ing the present-day attempts of scientists to solve certain social prob-
lems. For example, how many people know that a Science Committee
of the National Resources Committee is making studies of technological
trends and recommending national policies for meeting the problems of
technological unemployment?


How many know of the present attempts of certain technical schools
to give their students not only work in their special sciences but also
a substantial amount of work in the social sciences (economics, so-
ciology, history, etc.) in order to produce men of social vision and
sound training-men who will have a definite understanding of the
problems awaiting them in industry and society?
Then, the common man should be made to realize that there can
be no stopping of scientific research-that not only is it a normal hu-
man activity in response to natural curiosity, but also it is being urgent-
ly demanded by society to give more knowledge to doctor, engineer,
manufacturer, farmer, economist, psychologist, etc. Most scientists be-
lieve, not that there is today too much science, but rather that the scien-
tific attitude-the essence of science-is far too little understood and
too inadequately applied to human problems. In the words of Albert
R. Mann in his Report on the Program in Southern Education: "Most
obviously the sciences are destined to play an increasing part in the
progress of the South as the years pass."6
In this connection, warning should be given that our social problems
are not going to be solved by the development and extension of sci-
entific knowledge alone, but that a decided change in mental attitude,
a spirit of brotherhood, and a desire for social justice are also needed.
2. Scientists need to become more interested in social problems. They
must demonstrate that they have a vital interest in, and a human con-
cern for, the problems of society. Many will need a radical reconstruc-
tion of their attitudes-their desires to be left alone in their scientific
cloisters, out of touch with the world about them, completely absorbed
in the advancing of the frontiers of scientific knowledge. They will
need to come out occasionally-to look about them-to make note of
various problems of the community, state, and nation. Is the govern-
ment efficient and honest? If not, why not? What about improving
housing and employment conditions? Is improvement needed in the local
schools, local health conditions, opportunities for recreation, and so
,on without end? The reason why certain political subdivisions often
elect to do the foolish and unwise thing is because the simple everyday
issues in economic and political questions are often too complex and
involved to be understood by the untrained and uninformed minds.
What can be expected if the cultivated minds of the community do not
attempt to clarify issues or call attention to advantages, disadvantages,
or dangers?
3. To help solve social problems, the scientist must study them care-
Jully. He must avoid approaching the problem with a superior air,
6General Education Board, Annual Report, 1938 (New York), pp. 5-63.


thinking the answers will be immediately forthcoming because of his
scientific methods of attack. He will find the social problems much.
more difficult than the problems in the physical and biological sciences
because of the numerous contributing factors and the unpredictable
human element. As an illustration, consider the problem of old age se-
curity and pensions-a challenge to the best minds of the nation. I
doubt whether anyone has the final solution to that one.
4. More scientists should select research problems more closely re-
lated to present social problems. There is no intention here to detract
from the many important basic scientific research projects under way.
They are necessary even though no immediate applications to in-
dustry and every day problems are apparent just now. However, if
scientists for a time gave more attention to research that finds immedi-
ate social applications, they would surely be performing a greater serv-
ice to our present society. It may very well mean the difference between.
the formation of a dictatorship and the preservation of our democracy.
So says John Studebaker,7 United States Commissioner of Education:
"We must use our scientific knowledge to benefit the common man by
creating a widespread and stable prosperity, or we may lose our
democracy, the very mother of science." We must recognize this to be-
a time of crisis in which extraordinary measures are demanded. As an
illustration, recall how during the great war it was necessary for a
multitude of our chemists to give up their pet researches for the prob-
lems of manufacturing dyes and war gases. Particularly valuable today
would be researches leading to new industries or new uses for farm
Now let us see how an organized group of scientists like the Florida
Academy may be a factor working towards social progress.
1. By encouraging research in social problems-putting more em-
phasis on science problems in relation to society. The formation of our
Social Sciences Section with this meeting is significant. This should
mark the beginning of increased attention to social, economic, and
political problems, not alone by the regular workers in the social
sciences but also by others.
2. By concerning itself with the conservation of natural resources.
It is natural and appropriate that the Florida Academy of Sciences
should act as guardian of the natural resources of the state. The welfare
and happiness of our people depend much on the wise use and the pro-
tection of our water, soil, and minerals; on the preservation of our useful
and interesting plant and animal life; and on the destruction of harmful.
7Vital Speeches, Vol. 5 (Sept. 15, 1939), p. 727.


organisms. Other state academies of science have already done valuable
work in this regard through their special committees on conservation.
You will be pleased to know that during this year your Council or-
ganized a Conservation Committee with Sidney A. Stubbs, Assistant
State Geologist, as chairman. This committee has already acted in re-
gard to the conservation of Florida's artesian water supply, and has
by resolutions urging appropriate control measures brought the mat-
ter to the attention of the legislators and others vitally concerned.
The other members of this committee are R. S. Atwood, H. L. Baker,
G. G. Becknell, R. F. Bellamy, A. F. Carr, Jr., Herman Gunter, H.
J. Gut, H. H. Hume, Edward T. Keenan, B. M. Kinser, Herman
Kurz, H. S. Newins, W. S. Phillips, and G. G. Scott, representing all
parts of the state and various phases of conservation. This work on
conservation will undoubtedly become increasingly important in Flor-
ida. Every member of the Academy should become conservation mind-
ed, and should help to create a public interest in conservation, and
to give valuable information and suggestions to this committee.
3. By encouraging and suggesting research. The Academy can well
become a major force toward social progress in Florida if it serves
to encourage research in all possible fields. At the first four annual
meetings of our Academy the numbers of papers given were respectively
28, 28, 33, and 46. Next year we ought to have at least 60 papers
presented. It is safe to say that much of this work would not have
been done without the Academy.
There is no better way to get inspiration for one's work and ideas
for one's research problems than by attending all the sessions of the
Academy. There is a particular value in listening to papers in fields
other than one's own-it keeps one from becoming too special-too
provincial, if you please. The mathematician can get ideas from the
psychologist, the chemist can learn from the biologist, etc. We need
more opportunity to look around to see what the other fellow is doing.
For this reason we are fortunate in not having too many different sec-
tions-some academies have as many as ten or fourteen.
For this very purpose of giving the academy members ideas and
suggestions for research problems, your Council brings to you in to-
morrow morning's session, Mr. M. C. Forster, Consultant on Re-
search Projects for the Work Projects Administration, from Wash-
ington, D. C. Don't miss that special opportunity.
4. By promoting and encouraging more effective science teaching
in our secondary schools and colleges. The Academy, representing all
the sciences, is the logical group to be interested in seeing that our
young folks receive the proper instruction in the sciences. As Doctor
Allen pointed out this afternoon, and as many of us know, there is


considerable room for improvement in our science teaching in the
high schools. To enable the academies to exert a helpful influence
in this regard, some of them have instituted sections given over en-
tirely to papers and round-table discussions on science teaching. Then,
too, some of the academies have made, and are still making, thorough
studies of the status and trend of science instruction in the high
schools. The American Association for the Advancement of Science
has also acknowledged the need for improvement of teaching of sci-
ence, and has organized a Committee on the Improvement of Science
in General Education. This committee has begun its work by studying
the teaching of science in the colleges9. Mr. W. L. MacGowan is giving
a brief report at this meeting on the formation of a National Committee
on Science Teaching by the National Education Association which has
thus shown its awareness of the problems in this field.
It is of fundamental importance to the various professions, to in-
dustry, and to society how science is taught. We have too many people
graduating from high school without having acquired the ability to do
elementary scientific thinking. Our Academy should likewise consider
both the encouragement of the presentation of many papers on science
teaching and the making of a study of the present status and trends in
science instruction in the high schools.
5. By promoting a general interest in science and scientific methods.
Let me read from the constitution of our Academy. "The purposes of
the Academy shall be to promote scientific research, to stimulate
interest in the sciences, to further the diffusion of scientific knowledge,
to unify the scientific interests of the state The Academy can
stimulate general interest by encouraging the widest possible member-
ship-including all high school science teachers and others who have
a genuine interest in science. To attract and satisfy this element your
Councils have welcomed the presentation of a fair number of papers of
general interest, in addition to the more technical papers.
Another important way to stimulate a wide interest in science is
through the creation of Junior Academies of Science-composed of
the science clubs in the high schools. Eleven of the thirty state
academies have already done so. Your Council also has during these
past two years worked toward the creation of a Junior Academy. Last
year our Vice-President, Miss Charlotte Buckland, did valuable ground
work; and this year our Vice-President, W. L. MacGowan, with
a great deal of effort, brought the work to fruition. You see by your
program that the organization meeting of the Junior Academy of Sci-
ences of Florida is scheduled for tomorrow morning.
8Science, Vol. 87 (May 20, 1938), p. 454.
9Science, Vol. 88 (Jan. 13, 1939, p. 34.


Every member of the Academy should be vitally interested in these
high school science clubs in his community and should give them every
possible assistance and encouragement. It should be remembered that
the work is being sponsored by the Academy. It is one of our best
methods of furthering the diffusion of scientific knowledge.
To illustrate further the possibilities of promoting a general interest
in science, it is well to note what the Kansas Academy of Science is
doing. It is undertaking to arouse the layman's interest in the out-
doors and the physical world by writing handbooks on the botany,
entomology, zoology, archaeology and geology of the state. Florida
would offer excellent material in this regard.
6. By giving proper publicity to scientific matters. The Academy
can let the public know in various ways just what its scientists are
doing. Newspaper publicity at the times of the annual meetings usually
doesn't do more than give the names of the officers and the general
character of the meeting. We need some sort of permanent publicity
work active throughout the year. From the material in the third volume
of our annual Proceedings a skillful writer could make a dozen science
news features, highly interesting and informative to the public. The
papers in our present program would provide at least twenty feature
stories. The intelligent public would like to hear what their nutrition-
ists, entomologists, botanists, geologists, psychologists, and economists
have worked out.
Scientists may now be assured of increased interest and cooperation
on the part of the newspapers. More and more of them are adding
science editors to their staffs. An increasing number of newspaper re-
porters and writers are devoting their major efforts to the field of sci-
ence. They have organized a National Association of Science Writers.
In their constitution they state their one and only purpose: "To foster
the dissemination of accurate scientific knowledge by the press of the
nation in cooperation with scientific organizations and individual sci-
entists." Mr. David Dietz, Science Editor of the Scripps-Howard News-
papers, ably described this increased newspaper cooperation in his
address 10 to the American Association for the Advancement of Science.
He says: "I do not believe that any scientist may feel that he has
completed his work when he has finished a piece of research in the
laboratory. It is likewise his duty to disseminate the new knowledge
which he has uncovered. And today, publication in a journal read by
a circle of his colleagues is not sufficient. The welfare of society
demands that the general public be made aware of scientific progress.
This can be done only through the newspapers, and hence the scientist
today must be willing to cooperate with the newspaperman."
10Science, Vol. 85 (Jan. 29, 1937), pp. 107-112.


7. Finally, the Academy may perform an important service to the
state by always keeping a close contact with the state legislature on
matters in which the scientists can give competent assistance.
In reviewing all these different ways in which scientists may become
helpful and significant factors in social progress, one cannot help be-
ing impressed with the fact that we have a great responsibility, re-
quiring mature thought and judgment, and determined action. We
must cease drifting along. We dare not fail society. The Florida
Academy of Sciences has a great work to do!

Florida State College for Women

The belief that men and women differ in some characteristic way
temperamentally as well as physiologically, commonly accepted in the
lay mind as well as in song and story, has only recently been sub-
jected to objective analysis. This analysis, developing primarily as
a sub-problem in connection with some other major issue, has culmin-
ated in the carefully developed Attitude Interest Analysis Test of
Lewis M. Terman and Catherine Cox Miles, the main object of which
is the discovery and description of sex differences in attitude and
interest.1 While this test, built upon the earlier studies analyzed
for sex differences, is the result of ten years of investigation, both
extensive and intensive, the authors consider it by no means a fin-
ished product. When an instrument of this sort appears it challenges
the rest of us to investigate the uses to which it can be put, to
discover the strength and weakness of it. This paper reports one
step in the study to discover to what extent the test describes our
That sex differences in attitude and interests do exist in our pres-
ent civilization seems to have been clearly demonstrated,2 although
whether the exact extent or nature of these differences has been stated
mathematically is open to question. Such sex differences as Mr. Ter-
man and Mrs. Miles report might be a reflection of the population
from whose responses the test was constructed and upon which the test
was standardized. In short, it is conceivable that Mr. Terman and
Mrs. Miles might find differences of some magnitude within a single
sex group providing the representatives in the group were reared in
strikingly divergent cultures.
An application of the Attitude Interest Analysis Test to diverse
populations should throw further light upon the reliability of the
description of sex temperament as made by the test.3 With this in
1Terman, L. M. and Miles, C. C., Sex and Personality (New York: McGraw-
Hill Book Company, Inc., 1936.)
2Ibid., p. 472.
3Masculinity and femininity are to be interpreted in terms of the sex difference
disclosed by the test.


view, the test (Form A) was administered to 556 students at Florida
State College for Women during the winter of 1938. The test had
not yet been extensively publicized so it is fairly safe to say that
we were working with naive subjects. In the late spring of 1939 the
test was given to 490 University of Florida students. By that time
a number of popular descriptions of the test had appeared in daily
newspapers and in popular magazines, so in all probability our mas-
culine population was not entirely naive.
The question at once arises as to why the application of the
test to this particular group would be in any way crucial. Our first
thought was that these are southern colleges and would therefore
reflect the southern tradition, a tradition of sheltered and protected
womanhood, of chivalrous gentlemen and great statesmen. Actually
83% of our female population and 73% of our male population were
born south of the Mason-Dixon line, 59% and 54% respectively
having been born in Florida. In Florida itself there are three cultural
influences, to some extent contradictory in nature: (1) in some cases
the southern tradition undoubtedly operates, families are proud of their
connection with the Old South and endeavor to rear their children
in its traditions; (2) there is a large pioneer group which has come
to a comfortable climate, perhaps to a region which seems to offer
an easier living to a less aggressive people or a more propitious climate
to a physically weaker people than a more rigorous climate would;
and (3) there is a large, wealthy, pleasure seeking class.
We have the opportunity of comparing such a group with a group
consisting of sophomore women and men who were attending Stan-
ford University, Oregon State, Washington State, and the University
of Utah for whom Terman and Miles give M-F distributions. Data
from these colleges are combined into one distribution. For our
purpose it would be desirable to treat these groups separately, although
it is likely that there were no outstanding differences in the group
M-F responses since this was not done. The Stanford population,
at least, must represent both a socio-economic level and an intelligence
level different from that of the state universities since it has higher
admission requirements. The western state universities, in contrast
to Florida State College for Women draw their populations from
sections of rigorous winters with industries appropriate to that climate,
a country settled by a people who expected to find a hard life as
well as a good one.
A final factor must be mentioned. The Florida State College for
Women population is entirely female and the University of Florida
is almost completely male, whereas the western group is co-educa-
tional. Possibly such an environment would tend to make students
at Florida State College for Women more introverted, less likely


to exaggerate their emotional or other traditionally feminine reactions.
Where such an interesting environmental factor as men is lacking
one may have to look inward instead of outward for interests; and
where there is no appreciative audience it really does not pay to act.
Similarly, the University of Florida is a man's university. Again, the
male in the absence of the female does not have to act protective or
superior or any one of the dozens of things a man is popularly sup-
posed to be.
Such an analysis of our population together with an analysis of the
sex differences revealed by the test provides grounds for a surmise
that the northwestern and the southeastern populations might be dif-
ferent in attitudes and interests. The test is made up of seven exercises;
two association tests, an information test, a test of emotional and
ethical attitudes, one of opinions, and one of introvertive response.
In general it may be said, on the basis of the Terman-Miles investi-
gation, that:
the males included in the standardization groups evinced a distinctive in-
terest in exploit and adventure, in outdoor and physically strenuous occu-
pations, in machinery and tools, in science, physical phenomena, and in-
ventions; and, from rather occasional evidence, in business and commerce. On
the other hand the females of our group have evinced a distinctive interest
in domestic affairs and in aesthetic objects and occupations; they have dis-
tinctively preferred more sedentary and indoor occupations, and occupations
more directly ministrative, particularly to the young, the helpless, the distressed.
Supporting and supplementing these are the more subjective differences-those
in emotion, disposition and direction. The males directly or indirectly manifest
the greater self-assertion and aggressiveness; they express more hardihood and
fearlessness, and more roughness of manners, language and sentiment. The
females express themselves as more compassionate and sympathetic, more timid,
more fastidious and aesthetically sensitive, more emotional in general (or at
least more expressive of the four emotions considered), more severe moralists,
yet admit in themselves more weakness in emotional control, and (less notice-
ably) in physique.4
The southern woman, according to tradition, has centered her
life around her home and has left economic and political problems
to her husband; whereas the Northern and Western women have
been, supposedly, more militant suffragists, demanding release from
home duties and a voice in political and economic circles.
Terman and Miles find that as men and women become more
educated the M-F differences become less marked, the men becoming
more feminine in interests and attitudes and the women becoming
more masculine. Possibly high interest in education tends to select

4Terman and Miles, op. cit., p. 447.


an intermediate physiological type; surely the educated woman and
man are exposed to a broader common environment than are the
uneducated. Be that as it may, one might expect to find a less
masculine population at the Florida State College for Women and
a more masculine population at the University of Florida, since: (1)
the students at both colleges come from an environment in which
emphasis upon education is of relatively recent origin, (2) in both
colleges, classroom examples and applications of learning probably tend
to be exclusively in either the feminine or the masculine field of interest,
depending upon the sex of the group addressed, and (3) in both
institutions there are fewer outside interests-notably in the one case
lack of men and in the other lack of women.
The climate might be expected to influence both groups in the
direction of femininity.
On the other hand, absence of men or women-little emphasis
on the traditional differentiation of the sexes socially-and the pioneer
conditions in some parts of the state might tend to produce more
masculine men and more masculine women.
Turning now to the analysis of our data, the comparison of Florida
and Western distributions reveals two points of difference in both
male and female distributions (See Table 1). In both cases the Florida
group provides more extremely feminine scores and in both cases the
median score for the Florida groups is more feminine, the difference
being greater for the male than for the female distributions. These
differences in median are very slight, however, when the difference
between male and female medians is noted.
SCORE Western I U. of Fla. Western F. S. C. W.
100 +200 +187 + 41 + 80
95 +139 +140 + 8 + 18
90 +125 +124 4 10
80 +106 + 99 22 30
70 + 95 + 87 40 46
60 + 86 + 73 53 60
50 + 77 + 59 65 73
40 + 66 + 47 76 83
30 + 51 + 32 85 96
20 + 32 +17 95 -110
10 + 7 3 -107 -126
5 17 23 -117 -143
0 20 -116 -140 -194
N __492 __556
slbid., p. 472.


Study of the various sub-tests in two cases reveals interesting dif-
ferences between Florida and western populations, particularly in the
male distributions. These differences occur in test 1, a word asso-
ciation test, and in test 4, a test of emotional and ethical response (See
Table 2.) The Terman-Miles interpretation of test 1 is:
Females picking more often than the males upon terms for domestic things or
happenings, for kindly and sympathetic activities, and for articles or tokens of
adornment; males more often than females upon words relating to physical science,
machinery, outdoor pursuits, and terms suggestive of excitement and adventure,
and rather less predominantly upon political, business, and commercial words.6


CENTILE] Male | Female I Male Female
SCORE I West. I Fla. | West. I Fla. J West. Fla. West. I Fla.
100 +15 +14 +70 +60
95 +12 + 8 + 1 + 1 +58 +50 +40 +50
90 +10 + 5 2 5 +49 +44 +33 +34
so + 7 0 -5 -9 +40 +35 +25 +26
70 + 4 3 8 -11 +34 +32 +19 +17
60 +2 5 -11 -14 +29 +22 +13 +12
50 0 -7 -14 -16 +25 +17 + 6 +4
40 2 9 -16 -18 +20 +12 0 0
30 4 -11 -18 -20 +16 + 6 6 6
20 7 -13 -20 -23 +10 + 2 -12 -13
10 -10 -17 -22 -26 +1 8 -20 -24
5 -13 -19 -23 -28 8 -14 -26 -34
0 -25 --35 -__ -40 --54
'Ibid., p. 476.

In this case the University of Florida population falls about midway
between the western male and female groups, and the Florida State
College for Women tends to move in the direction of greater femin-
inity. Turning now to test four one finds a similar state of affairs
with respect to the male populations. Again borrowing the Terman-
Miles description of the sex reaction:
The female tends much more than the male to extreme expression in all four
emotions and in moral censure, particularly in disgust, rather less in pity, less
still in fear, and least in anger and censure; though her excess, if less marked
for censure than for pity, fear and anger, is more ubiquitous.8
On this test also the University of Florida population falls between
the western male and female populations.

6lbid., p. 380.
8Ibid., p. 415.


On the basis of these last two observations one is tempted to say
that our populations are somewhat different from the western popula-
tions; that, given more closely defined cultures than those represented
at our College or University, one might find greater or other differ-
ences. Or, differences disclosed may not be significant since the western
distributions were made on the basis of two forms of the test whereas
our study is based on Form A.
Accordingly the data were analyzed from another angle; both
Florida State College for Women and University of Florida groups
were divided into three sub-groups: (1) those subjects born in Florida;
(2) those born in the South; and (3) those born in the North. In-
terestingly enough this classification of subjects serves to separate the
distributions further (See Table 3). The subjects who were born in
the North, as a whole, rate as masculine as do those of the western
group, whereas those born in Florida rank still more feminine than
the western group. Not only do our data from the northern group cor-
respond rather closely to the western group, but they also agree, as far
as data are given, with a study reported from Minnesota by Gilkinson9
in which the median score for a group of Minnesota college men was

SCORES I North Florida North Florida
100 +179 +187 + 80 + 60
95 +147 +149 + 25 + 1
90 +119 +126 + 11 18
80 + 99 +99 17 32
70 + 94 +84 38 52
60 + 86 +69 51 68
50 + 74 +56 59 79
40 + 54 +43 71 89
30 + 47 +29 78 -102
20 + 27 +14 91 -115
10 + 12 5 -100 -133
5 5 32 -118 -150
0 52 -112 -149 -194
N 1 106 1 267 1 71 [ 240

Pursuing the same type of analysis further, the subjects born in
the North of pure northern-born parentage were compared with those

9Gilkinson, Howard, "Masculine Temperament and Secondary Sex Character-
istics: A Study of the Relationship between Psychological and Physical Measures
of Masculinity," Genet. Psychol. Monog., Vol. 19 (1937), pp. 105-154.


born in Florida of pure Florida-born parentage. In the case of both
men and women this selection of data serves to separate northern and
Florida groups still more (See Table 4), the Florida groups, both
male and female, tending to greater femininity in attitude and interest.

SCORE North Florida North [ Florida
100 +177 +187 + 75 + 2
95 +160 +140 + 25 22
90 +144 + 99 + 5 29
80 +104 + 81 33 52
70 + 96 + 71 40 63
60 + 87 + 61 53 72
50 + 82 + 50 60 90
40 + 72 + 34 73 -100
30 + 53 + 21 79 -110
20 + 40 + 6 97 -120
10 + 19 7 -110 -130
5 + 12 32 -130 -142
0 52 47 -144 -149
N 50 1 80 44 | 65

These findings suggest strongly that some selective factor related to
regional differences is operating here. The analysis suggests that as
groups become more homogeneous within themselves with respect to
cultural influences they tend to draw apart from each other in mas-
culinity-femininity response as described by this test. It is possible,
however, that this drawing apart of groups is due to some other factor;
possibly climate, race, or nutritional influences. This study does not
segregate the particular factors but merely suggests that there is
either a single factor or a group of factors inclining the Florida pop-
ulation to femininity of response.
In conclusion (1) our results support the Terman-Miles finding
that this test describes male and female populations differently; (2)
they disclose a greater femininity on the part of Florida born as com-
pared with the northern born; and (3) this greater femininity may
reflect different cultural influences or different physiological conditions
due to climatic, racial, or nutritional differences.


I wish to express my appreciation of the aid rendered by the National
Youth Administration through the assignment of students to this


project and of the support, both moral and financial, of the Social
Science Research Council which made possible the analysis of data from
the University of Florida. I also desire to thank Dr. Elmer HIinckley
of the Psychology Department of the University of Florida for collect-
ing the University data for this study. Similarly I wish to thank the
students who served as subjects in this experiment or otherwise con-
tributed to its progress. Finally, I am especially grateful for the interest
in and critical analysis of my work on the part of my colleagues in the
Psychology Department at Florida State College for Women.

University of Florida

Introduction ............................................................... ....................... 20
2. The Genus Progomphus....................................................... 21
3. N om enclature ............................................................ ........................ 22
4. G generic D definition .................................................. ............. ........... 24
5. The Species of the Genus ...................................... ...................... 26
6. Key to the Species of Progomphus.......................... ....................... 27
1. Historical Survey .............................................. 30
2. Geeographic Distribution ........................... ... ......................... 35
P. obscurus
P. borealis
P. alachuensis
Flying seasons
3. Progom phus obscurus ............................................... ............................ 40
Description and Figure references
Diagnostic characters
Variation study
Ecological Notes
4. Progomphus borealis ............................................. 46
Description and Figure references
Diagnostic characters
Variation study
Ecological Notes
5. Progomphus alachuensis, n. sp. ........................ .................... 0

1Contribution from the Department of Biology, University of Florida. The
author wishes to express his appreciation to the Museum of Zoology, Univer-
sity of Michigan, for materials and aid in the preparation of this paper. Also,
to Drs. Philip P. Calvert and James G. Needham for materials and advice.


Original description of male and female
Diagnostic characters
Variation study
Ecological Notes
1. Description of Progomphus Nymph ..................................... 56
2. H historical Survey ........................................................ ............................ 57
3. K ey to K now n N ym phs .......................................... ........................... 58
4. N ym ph of P obscurus ............................................. ............................ 60
Description and Figure references
Ecological Notes
5. Nym ph of P. borealis .............................................. .. .................. 63
Description and Figure references
Ecological Notes
6. Nymph of P. alachuensis .................... ... ................... 64
Ecology and Evolution notes
IV M material Studied .................................................................................................... 66
B bibliography ........................................................... ............ .... .............. 68
P la tes .................................................................................... ............................ 74
For the past ten years, the author has been interested sporadically
in a study of the species of dragonflies belonging to a genus best known
under the name Progomphus. Nearly one hundred years ago, 1842, the
first species of the genus, as it is now constituted, was described. Since
then twenty-four species have been named and at least two unnamed
species are known-one of which is described in this paper.2 The ma-
jority of these species occur in South America and Central America;
only three are to be found in North America north of Mexico.
Originally the plan was to write a formal monograph of the genus.
However, my experience with the North American species made it
conclusive that the genus could not be so treated until adequate series
of the South and Central American species were available. Intraspecific
variation is such that only after studying large numbers of individuals
can a comprehensive picture of the species be obtained.
Rather than delay publication until such a time as series are avail-
able, the author has decided to publish his study of the North Amer-
2The other unnamed species was described by E. B. Williamson (1920a: 16-17)
from Santa Marta but was simply designated as "Progomphus species". In the
same paper (1920a: 11), Williamson indicates the probability of other unnamed


ican species, where large numbers of individuals are obtainable, with
-as a background for future work-some information about the
remainder of the genus gathered from literature and a study of the rel-
atively few specimens in the Odonata collection at the University of
Michigan (see page 66 for a list of material studied).

In the Monographie des Gomphines (1858), De Selys presented a
comprehensive system for the classification of the Gomphus-like drag-
onflies. In this work, following his earlier system (Synopsis des
Gomphines 1854), De Selys recognized two divisions of the "Subfamily
Gomphinae"; one, the "Division of Integrilabiees", with the labium en-
tire; the second, "Division Fissilabiees", with the labium divided by a
median cleft.
The grand genus Progomphus was placed in the second legion
(Legion II. Gomphoides) of the second subdivision of the integrilabiees.
As related grand genera, De Selys listed: Gomphoides subgeneraa:
Gomphoides, Cyclophylla, Aphylla); Zonophora subgeneraa: Zono-
phora, Diaphlebia); Hagenius subgeneraa: Hagenius, Sieboldus). The
first legion was reserved for the grand genus Gomphus with fifteen
subgenera (most of which are considered good genera today); the
third legion included the grand genera Diastatomma and Lindenia,
the latter with four subgenera. These three legions constituted the
two divisions of the integrilabiees. The fissilabiees included Chloro-
gomphus, Cordulegaster, Petalia, Petalura, Uropetalia and Phenes.
Williamson (1920b:7) writes, "Since de Selys' Monographie des
Gomphines was published our definition of Gomphinae has grown by
reduction ... The Gomphinae of recent authors is the "Division Integri-
labiees" of de Selys, containing over fifty genera and three hundred
and ninety species."
The dragonflies of the "Division Fissilabiees" have been revised
by Fraser (1929). Those of the integrilabiees remain as a group to be
revised. However, Williamson (1920b: 6-11) offers a discussion of this
complex. On the basis of wing venation, Williamson divides these
Gomphines into six series: (1) The Epigomphus series to include eight
of the genera from De Selys, first legion. (2) The Progomphus series
to include the grand genera Progomphus and Gomphoides of De Selys
and Diaphlebia and Desmogomphus. (3) The Gomphus series to in-
clude the grand genus Gomphus of De Selys less the Epigomphus series.
"It is the dominant gomphine type in the world at the present time."
(4) The Zonophora series with the single genus Zonophora. (5) The
Hagenius series to include Hagenius and Sieboldius. (6) The Diasta-
tomma series-Legion III of De Selys.


Thus the Progomphus series includes four genera: Progomphus,
Gomphoides, Diaphlebia and Desmogomphus. The genus Gomphoides
has three distinct subgroups-Gomphoides, Cyclophylla and Aphylla-
which may very well be good genera, bringing the number of genera
of the series to six. This complex consists of about 60 species, mostly
Neotropical, a few of which have penetrated the Nearctic region (Pro-
gomphus obscurus being the most successful). Williamson (1920b)
characterizes the Progomphus series thus: "Genera of this series may
be recognized by having the triangles, supratriangles, and subtriangles
more or less cross-veined, by dissimilar triangles and subtriangles in fore
and hind wings, the triangles of the fore wings being relatively shorter
and of the hind wings relatively longer; by the absence of sectors of
Rs and M4 and of strongly developed trigonal supplements, and by
the outer side of the triangles not distinctly concave." Williamson
makes his genus Desmogomphus the most primitive of the series and
places the densely veined Gomphoides, such as G. stigmata, at the op-
posite end of the series.
Nomenclatural difficulties have beset the Progomphus series since
the middle of the last century when the genera were being established
by De Selys and Hagen (De Selys 1850, 1854, 1858). As regards the
genera "Progomphus" and "Gomphoides" these difficulties may be
stated as follows:
Under the generic name Diastatomma, Rambur (1842) described
two American species of the gomphines which he named D. obscurus
and D. infumata. In 1843 under the generic name Aeshna there was
described the fossil gomphine A. brodiei Buckman. In 1850, while
working on their famous monographs, Hagen placed brodiei in the genus
Gomphus; De Selys disagreed and said (1850:360 footnote), "I think
this belongs to my new American genus Gomphoides, the actual type
of which is Diastatomma obscura Rambur." He thus, according to
modern rules of nomenclature, established a valid genus with a desig-
nated type. In 1854, evidently forgetting the above quoted note, De
Selys reestablished the genus Gomphoides with Diastatomma infumata
Rambur as the type species, at the same time making D. obscura Ram-
bur the type of another new genus which he called Progomphus (De
Selys 1854). This arrangement was carried over to the Monographie
des Gomphines (De Selys 1858), the work to which authors are most
apt to refer for basic information in this group. The names Progomphus
and Gomphoides, in the Selysian sense, thus became well established in
the literature during the following years.
In his Catalogue of North American Odonata, Muttkowski uncov-
ered the above error and corrected it (Muttkowski 1910a:78). By


the code rules (Canon 42 A. O. U. Code and Art. 25 International
Code), the name Progomphus must be dropped in favor of Gomphoides
for the genus of which D. obscurus Rambur is the type, and the old
genus Gomphoides with D. infumata Rambur as the type must be
renamed. This new name was supplied by Muttkowski (1910a, 1910b)
as Negomphoides.
There is no doubt that the application of the rules of priority make it
necessary to discard the name Progomphus in favor of Gomphoides for
the genus under consideration in this paper. If the story ended here,
the situation would be relatively simple and easy to handle but it is
far more complicated, most of the complications being introduced from
the Negomphoides side of the case.

Cowley (1934: 244) states the facts as follows:
Muttkowski has already shown that Selys erected the genus Gomphoides so
that Progomphus Selys became a synonym of Gomphoides Selys, since the types
of the two genera are congeneric; consequently Gomphoides Selys (1854) ....
required a new name, for which Muttkowski proposed Negomphoides. Calvert
(1911 [Ent. News 22:81]) and Ris (1912 [Mem. Soc. Ent. Belg., 19:1011) objected
to this on the grounds that since the subgenera Gomphoides, Cyclophylla and
Aphylla had now been united, and as according to Art. 28 of the International
Rules, "a genus formed by the union of two or more genera or subgenera takes
the oldest valid generic or subgeneric name of its components," therefore, since
Gomphoides was invalid, Cyclophylla, which had page preference over Aphylla
should be used as the name for the genus; as, however, Muttkowski considered
Gomphoides, Cyclophylla and Aphylla as separate genera, he was justified in pro-
posing a new name for Gomphoides. Moreover, the exact application of Art. 28 is
not clear; Selys used the generic name Gomphoides to include his three subgenera;
if these subgeneric divisions are united ., should the generic name remain the
same, or should it be changed to the oldest (valid) subgeneric name? In the for-
mer case, the generic name Negomphoides, as the new name of Gomphoides, must
be used; in the latter case, Negomphoides cannot be used, since it is not the oldest
generic or subgeneric name, therefore the name Cyclophylla comes into considera-
tion; but is preoccupied by Cyclophylla Brandt (1837) in Coelenterata To pro-
pose a new name for Cyclophylla Selys would be to repeat the objections against
Negomphoides, so the oldest valid subgeneric name is Aphylla. I believe this con-
fusion may be resolved by applying the second sentence of Art. 28: "If the names
are of the same date, that selected by the first reviser shall stand." Calvert
(1905 [B. C. A.]) was the first author to unite these three subgenera; he was the
first reviser, and "selected" the name Gomphoides; .. then Negomphoides, as the
new name of Gomphoides, must be the valid name of the genus.

To further complicate the matter, Kirby in 1899 described a new
species of Gomphoides (now Negomphoides) which he called obscura.
There are, therefore, two separate species called obscura, both of which
have been, or will be, referred to as Gomphoides obscura.


The results of these nomenclatural difficulties in the Progomphus
series are to make the name Gomphoides so ambiguous and beset by
confusion that many authors, priority to the contrary notwithstand-
ing, have returned to the use of the names Progomphus and Gomphoides
as employed by De Selys in the Monograph of 1858.
Of some 37 titles published since 1910 involving reference to Ram-
bur's Diastatomma obscurus only 7 have employed the generic name
Gomphoides, the remainder Progomphus. Some authors have used both
names on different occasions (Byers 1930=Gomphoides; 1925, 1927,
1931, 1934a, 1934b=Progomphus). Calvert, Williamson, Needham,
Ris, Kennedy and others have, to the best of my knowledge, never used
Gomphoides in place of Progomphus, or Negomphoides in place of
Gomphoides. Ris (1911) argues against strict application of priority
when it leads to the upsetting of old established names. "Every Odon-
atologist knows what Progomphus means; he will have to look twice to
see which Gomphoides is meant" (Calvert, in a letter to the author Aug.
17, 1939). Confusion of names has caused one writer (Borror, 1937)
to use Negomphoides for Progomphus. Finally, the rule of priority has
been set aside by the International Commission when the strict applica-
tion of the rule will clearly result in greater confusion than in uniform-
ity (see decision 82. Suspension of Rules for Musca Linnaeus). The
Gomphoides-Progomphus case so far as I know has never been sub-
mitted in this light to the International Commission.
Kennedy (1929: 670), writing on, "The Theory of Nomenclature"
says in defense of the rule of priority:
If some of these names have been excepted from the rule of priority by special
rulings of the Commission, such names will always remain sore spots to be
reopened by those craving uniform rule. Following a uniform rule of priority will
eventually clear the situation as the present literature disappears into the past.
Making exceptions will always leave said exceptions for future arguments. We
have not removed the emotional from the code by making exceptions to the rule
of priority. Elimination of the emotional factors produces stability. The code
must be stabilized to become useful. [Kennedy in 1928 was still using the name
The present writer agrees with Dr. Kennedy the rules of priority
should be adhered to, but I shall use the name Progomphus, until
formal monographs for both Progomphus and Gomphoides may be writ-
ten and the consequent difficulties ironed-out.3
Genus PROGOMPHUS Selys 1854.
Genus GOMPHOIDES Selys 1850.
3A second reason for using Progomphus arises from the great amount of quoted
material in the present paper.


Type: Diastatomma obscurus Rambur 1842.
This genus may be characterized as follows:
Triangle and subtriangle usually4 with cross-veins (except P.
paucinervis and P. gracilis.) Supratriangle usually free of cross-
Triangles and subtriangles of fore and hind wings dissimilar,
triangle with outer side somewhat angulate, (4 sided).
Stigma with proximal base-vein.
Arculus at or proximal to 2nd Anq5; sectors of arculus distinctly
separated at their origin by an interval greater than the thickness
of either sector.
Usually one submedian cross-vein (Cuq) present (usually two
in the fore wing of P. phyllochromas).
Usually one basal subcostal cross-vein (Scq) present (except
P. gracilis, P. auriopictus and P. lepidus).
Thickened distal Anq usually the fifth.
Trigonal supplement well developed.
1-2 rows of cells between M1 and the next supplementary sector
below at the level of the distal end of the stigma.
2 rows of cells between Ml and M2 beginning more remote
than half-way from nodus to stigma.
2 rows of postanal cells in fore wing beginning proximal to the
level of the triangle (except P. pygmaeus, P. gracilis and P. pauci-
3 or more rows of cells between Cu2 and the hind wing margin.
Anal loop absent or indistinct.
Anal triangle of males not reaching backward to the anal angle
of the wing; usually three celled.
Legs short, hind femora hardly reaching to the apex of the first
abdominal segment, with short spines all of about equal length.
An occipital ridge present between the compound eyes.
Inferior appendages of the male bifid, the arms not divergent.
4"Usually" as here employed means "to be expected"; that the wing feature thus
designated is present in at least one of the wings (either fore or hind or both as
stated) but may be absent or modified in the other or others; generally it is pres-
ent in all of the wings indicated, (see analysis of wing venation for P. obscurus
page 42.) "Except" means that the feature thus described is usually lacking,
but it may be present in a small percentage of cases, seldom in all wings of the
same specimen.
51 am using the cross-vein abbreviations employed by Ris (1908, 1911, 1916):
Anq=antenodal cross-vein.
Pnq=postnodal cross-vein.
Scq=basal subcostal cross-vein.
Cuq=submedian cross-vein (other than the ones forming parts of the triangle).



The genus Progomphus is limited in its geographic distribution to
the Western Hemisphere and at present consists of 26 species including
the unnamed form described by Williamson (1920a:16-17).
These species may be listed as follows, in order of the date of descrip-
1. Progomphus obscurus Rambur. Described in 1842, the first
species of the genus to be described and probably today the best known.
Type locality given by Rambur-North America. (Rambur 1842).
2. Progomphus gracilis Hagen. 1854. Brazil (De Selys 1854).
3. Progomphus complicatus Selys. 1854. Brazil, Bahia (De Selys
4. Progomphus costalis Hagen. 1854. Brazil (De Selys 1854).
5. Progomphus zonatus Hagen. 1854. Mexico (De Selys 1854).
6. Progomphus intricatus Hagen. 1858. "The Amazon Country" (De
Selys 1858).
7. Progomphus paucinervis Selys. 1873. Quito [Ecuador] (De Selys
8. Progomphus borealis MacLachlan. 1873. Oregon [United States]
(De Selys 1873).
9. Progomphus pygmaeus Selys. 1873. Bogota [Colombia] (De Selys
1873 appendix).
10. Progomphus integer Hagen. 1878. Cuba (De Selys 1878).
11. Progomphus serenus Hagen. 1878. Haiti (De Selys 1878).
12. Progomphus polygonus Selys. 1879. Merida [Venezuela] (De
Selys 1879).
13. Progomphus clendoni Calvert. 1905. Mexico (Calvert 1901-
1908 B. C. A.)
14. Progomphus joergenseni Ris. 1908. Argentina. (Ris 1908).
15. Progomphus recticarinatus Calvert. 1909. Brazil (Calvert 1909).
16. Progomphus auriopictus Ris. 1911. "Territory Misiones" [Ar-
gentina]. (Ris 1911).
17. Progomphus lepidus Ris 1911. "Territory Misiones" [Argentina]
(Ris 1911).
18. Progomphus recurvatus Ris 1911. "Espirito Santo" [Brazil]
(Ris 1911).
19. Progomphus basistictus Ris. 1911. "Territory Misiones" [Ar-
gentina]. (Ris 1911).
20. Progomphus longistigma Ris. 1916. Costa Rica. (Ris 1916).


21. Progomphus phyllochromus Ris. 1916. Peru. (Ris 1916).
22. Progomphus perpusillus Ris. 1916. Peru. (Ris 1916).
23. Progomphus risi Williamson. 1920. Guatemala. (Williamson
24. Progomphus sp. 1920. Guatemala. The described but unnamed
species of Williamson. (Williamson 1920a).
25. Progomphus dorsopallidus Byers. Venezuela. (Byers 1934b).
26. Progomphus alachuensis n. sp. Florida [United States].
In general the type localities of the above listed species may be
grouped as follows: Brazil 6; Argentina 4; United States 3; Peru,
Venezuela, Guatemala, and Mexico each 2; Colombia, Ecuador, Costa
Rica, Haiti and Cuba each 1.
Thus the genus is primarily a South American group with the
greatest number of species occurring apparently in Brazil. The nymphs
of Progomphus are associated ecologically with moving water and
clean sand bottom habitats. No doubt the many small tributaries of
the Amazon furnish ideal habitats for them.
The following key has been compiled chiefly from published de-
scriptions (see literature references for each species as given in the fore-
going list of species) and has been checked against the available study
material (see list of material studied page 66). Specimen identifica-
tion should be checked with drawings of the male appendages and of
the thoracic pattern (Plates III, IV and V) when available.
1- N o basal Scq present .................................................... ........................... (2)
B asal Scq present ............................................................ ................................(4)
2-A diffuse golden yellow spot under the stigma of both wings; generally 2 Cuq
in fore wing; a light line on front of dorsal suture; superior appendages
of males small and acute; inferior appendages with no evident side branches;
stigma light reddish brown, 3 mm. long, covering 5 cells; abdomen of
male 29 mm, hind wing 24 mm. ...................................................... auriopictus.
-No yellow apical spot on wing; 1 Cuq in fore wing; dark lateral portion
of thoracic pattern well developed; no light line on dorsal suture; superior
appendages of male longer than 10th abdominal segment; inferiors with
side branches forked ........................................................................ ....... ... .. (3)
3-Male with pale humeral stripe broad, connected with collar, upper portion
wedge-shaped and tapering; female with humeral stripes well separated
from collar, reduced to narrow short lines; male superior appendages
very much expanded at base, abruptly narrowed at center with a small
angle on lateral margin; branches of inferior appendages very slim, dorsal
branch distinct from ventral view; stigma dark reddish brown, 3 mm. long,
covering 4-5 cells; abdomen 27-28 mm. long; hind wings 22-24 mm.
long ............. .................. ........... ......................... ..... ............... gra is.


-Male with pale humeral stripes narrow, sides parallel, united at lateral
edges to very reduced collar, (female unknown); male superior append-
ages parallel, lateral basal expansions very indistinct; branches of inferior
appendages thick, branches of dorsal process small, not noticeable when
viewed ventrally; stigma grey-brown. ............................... ................. epidus.
4-Dark brown markings present at base or along costa of wings................(5)
-No dark markings present on wings; colors, if present, limited to ochre,
yellow or grey. ..................................................... ................................... (15).
5-Wings with brown costal band from base to apex, limited by R and Ml;
size large, abdomen 34-36 mm., hind wing 27-31 mm. ......................costalis.
-Wings with no such wide costal brown band. ....................................................(6)
6-Stigma light yellow; no tubercle on the venter of abdominal segment 1.....(7)
-Stigma dark brown to black; tubercle usually present on venter of abdominal
segm ent 1. .............................................. ........... .................................... ......... (8)
7-Face brownish with a distinct black band; brown spot on nodus; size large,
abdomen 37 mm., hind wing 28 mm. ........................................joergenseni
-Face greenish grey, no distinct dark stripes or bands; no brown nodal spot;
size medium, abdomen 32 mm., hind wing 23 mm. ..............dorsopallidus.
8- Face with distinct brown markings.... ...............................................................(9)
-Face with diffuse or no brown markings. .............................................. (11).
9-Lower lip burnt yellow; upper lip deep brown, with the sides surrounding
a large oval milk-white spot, anterior border narrowly brown; rhinarium
white; nasus deep brown with lateral round spot and border next to rhin-
arium white; frons deep brown ahead, white above, the base in front
of ocelli with broad band of deep brown; stigma 4 mm. long sur-
m counting 7 cells. ........... .......................................... ....................... integer.
Face lacking w hite m arkings ...............................................................................(10).
10-Face with dark markings on inferior part of frons, on middle of nasus, and
middle of labrum; pale humeral stripe not confluent with the green or
yellow of the anterior mesothoracic margin; pale metepisternal stripe al-
most interrupted to form a superior spot and an inferior stripe; wings with
the slightest trace of brown at extreme base, not reaching half-way to
Scq; stigma 4.5 mm. long, surmounting 9 cells; abdomen 40-43 mm., hind
w ing 32-33 m m long. ...............................................................................zonatus.
-Face with lower lip indistinctly yellowish-brown, upper lip light olive, lower
part of frons olive, upper part diffuse brown; top of head dark brown;
pale humeral stripe similar to zonatus; pale metepisternal stripe entire, not
interrupted or broken; wings with very dark blackish-brown patches at the
base extending nearly to Anq 1; stigma 3.5 mm. long; abdomen 32 mm.,
hind w ing 25 m m long ........................................................................basistictus.
11-Face with obscure brownish markings on inferior part of frons, on middle of
nasus and middle of upper lip; pale humeral stripe not or barely (i e., by a
line) confluent with the green or yellow of the anterior mesothoracic
margin; pale metepisternal stripe similar to zonatus; abdomen 38-41 mm.,
hind wing 29-31 mm. long. Female unknown ..............................clendoni.
-Face without brown markings; pale humeral stripe broadly or very narrow-
ly confluent with the yellow-green of the mesothoracic margin (collar);


pale colors predominating on the mesepimeron and metapleuron, pale stripes
thereon not interrupted; wings with two short basal brown stripes........(12)
12-A single row of denticles on the inferior margin of male superior appendages,
the carina extending to the last denticle ........................ ................ .... (13)
-Denticulated carina on the inferior margin of male superior appendages
terminating apically in two or more rows of denticles, the carina not
extending to the last denticle. ...................................... ...................................(14)
13-Brown basal stripe in costal and subcostal spaces not reaching beyond basal
subcostal cross-vein; no brown traces at origin of sectors of arculus;
pale humeral stripe very broad and broadly joined to the pale area of the
mesothoracic collar; male superior appendages yellow; size large, ab-
domen 41-47 mm., hind wing 31-38 mm. long. .......................... borealis.
-Brown basal stripe in costal and subcostal spaces reaching to first antenodal
cross-vein or nearly so; a trace of brown between sectors of arculus at
their origin; pale humeral stripe narrow, sides serrated, not or very narrowly
joined to the restricted pale area of mesothoracic collar; male superior
appendages with tips yellow, basal third brown; size: abdomen 34-39
mm., hind wings 30-35 mm. long. .................. .........................alachuensis, n. sp.
14-Brown basal stripe of the costal and subcostal spaces reaching to first an-
tenodal cross-vein or nearly so; a trace of brown between the sectors of
the arculus at their origin; pale humeral stripe narrow, sides straight,
joined to pale area of mesothoracic collar; male superior appendages
yellow; size: abdomen 35-41 mm., hind wing 30-37 mm. long..........obscurus.
15-Size small; abdomen, male 18-25 mm., female 19-28 mm. long; hind wing, male
15-22 m m long. ....................................................... ................................(16)
-Size larger; abdomen, male 29-39 mm., female 30-40 mm. long; hind wing,
male 24-34 mm., female 25-36 mm. long. ................................................... (19)
16-Size very small, male abdomen 18 mm., hind wing 21 mm. Female un-
know n. .......................... ..... ............. ............... ................... perpu illus.
-Size larger, abdomen 22-28 mm., hind wing 19-25 mm. long. ......................(17)
17-Stigma dark-brown, 4-4.5 mm. long; thorax dark, pale humeral stripe broadly
wedged shape, tips attenuated, sides dark; abdomen 28 mm., hind wing 22-25
m m long ........................................ .......................... .................... longistigm a.
-Stigma dark-brown; thoracic pattern not as above. ..................................(18)
18-Anal field of fore wing one-cell wide throughout; 3 cells in proximal row
of postanal cells in hind wing; legs, especially third pair, relatively long and
slender; hind tarsus without claws, about two-thirds as long as hind
tibia; occipital hairs long (35-40 mm.); thoracic pattern distinct [PI.
IV fig. 1] ................................................................................................pygm aeus.
-Anal field of fore wing two-cells wide for short distance both distal and
proximal to the level of the triangle; 4 cells in proximal row of postanal
cells in hind wings; legs, especially third pair, short and stubby; hind
tarsus without claws, longer than the shortened second tibia, giving the
leg a malformed appearance; occipital hairs short (.25 mm); thoracic
pattern distinct [PI. IV fig. 81. ...... ................ ........ .......... .. i....................
19- Stigma yellow or gray-brown. ................................... ........ ................ (20)
-Stigma dark brown, red or black. .........................................................................(21)
20-2 Cuq in fore wings, 1-2 in hind wings; thorax with mesepisternal light


band (pale antehumeral stripe) lacking; pale humeral stripe very broad,
yellow ......................................................................................... phyllochrom us
-Usually 1 Cuq in all wings; thorax with pale antehumeral stripe present;
pale humeral stripe narrow and angulate at mesothoracic collar....recurvatus.
21-Triangles of wing nearly always free; fore wing with the anal field proximal
to the triangle one-cell deep; stigma 3 mm. long covering 5 cells, clear
brown; wings clear; face yellow with appearance of black on the frons;
abdomen 31 mm., hind wing 29 mm. long.................................. paucinrvis.
-Triangles 2-3 celled, seldom any free; fore wing with anal field proximal
to triangle with at least a few double cells................................... ............. (22)
22-Pnq 11-14; face light olive or white-green..................................................(23)
-Pnq 7-10; face dark brown or if light having dark bands...........................(24)
23-Triangle of fore wing with usually three cells; Anq. 15, Pnq 11-12; face white-
green, sides pale green ................................................ ...........................serenus.
-Triangle of fore wing with usually 2 cells; Anq. 16-17, Pnq 13-14; face olive
green slightly reddish. ...... ............................................................polygonus.
24-Size small, abdomen 30 mm., hind wing 25 mm.; all triangles 2 celled; Anq.
10-14, Pnq 7-8; stigma 4-5 mm. long; face, labium, nasus and frons rich
dark brown. .......................................... [Williamson's unnamed species. 1920a]
-Size larger, abdomen 30-39 mm., hind wing 23-36 mm., Anq 12-17, Pnq
8-10 .................................................................. .......... .......................... ...... (25 )
25-Face blackish-brown; size larger, abdomen 36-39 mm., hind wing 28-36 mm.
long. ......................................................................................................com plicatus.
-No blackish-brown on face; size smaller, abdomen 30-36 mm., hind wing 23-
30 mm.; yellow predominant on dorsum of abdominal segments 3-7. ....(26)
26-Occiput wide; superior frontal band pale brown; hindlobe of prothorax
blackish; dark color of thorax pale brown; distance on fore wing from
nodus to stigma 6-7 mm................................................................... intricatus
-Occiput narrow; superior frontal band blackish; hind lobe of prothorax yel-
low; dark color of thorax blackish-brown; distance on fore wing from
nodus to stigma 7-8 mm ...................... ......... ..........................recticarinatus.

It has been noted that Progomphus is primarily a South American
genus with the greatest number of its species occurring apparently in
Brazil; also, that there have been described two species from North
America north of Mexico-P. obscurus Rambur 1842 and P. borealis
MacLachlan 1873. The writer is describing a third species P. alachu-
ensis in this paper.

The two described North American species of Progomphus have re-
ceived a variety of treatment at the hands of various taxonomists.
P. borealis has been considered a good and distinct species by some,
a synonym of P. obscurus by others and a subspecies of P. obscurus by
still others. The history runs somewhat as follows:


1842. Rambur described obscurus, from North America, under the
generic name Diastatomma.
1850. De Selys made Ramur's obscurus the type of the genus Gom-
phoides (see discussion page 22.)
1854. De Selys wrote a description of adult female of P. obscurus
and stated, "Male unknown. Habitat: North America (Coll. Selys and
Berlin Museum)." In this same paper, De Selys erected the genus
Progomphus with Rambur's obscurus as the type.
1858. In the Monographie des Gomphines, De Selys repeated the
description and notes from the Synopsis (1854 above) and adds, "Hab-
itat North America after the female specimen in which the body is
nearly destroyed, and which served as a type for M. Rambur and
which is to be found in the Serville Collection coming from that of
Latreille. The other specimen is in the Museum of Vienna, and its origin
is not known." Some notes are included on the resemblance of P.
obscurus to other species of Progomplus and related genera.
1861. Hagen, under the name P. obscurus, rewrote the material
from De Selys 1858 and noted, "Habitat: North America (Vienna
Museum; collection of de Selys Longchamps)."
1872. Cabot assigned a nymph collected near Wareham, Massa-
chusetts to P. obscurus on supposition.
1873. De Selys, in the 3rd addition to the Synopsis of the Gom-
phines, included a description of a male gomphine under the name
Progomphus borealis MacLachlan, stated that the female was unknown,
and wrote, "Habitat: Oregon. One male from Lord Walsingham.
(Communicated by M. MacLachlan) N. B. Differs from P. costalis
(of Brazil) by larger size .... it approaches obscurus and zonatus in
size. It is easily separated from the first by the absence of an obscure
mark at the origin of the sectors of the arculus, the stigma more abrupt
not surmounting 5 cells, and the costa yellow externally."
To this point in the chronology, except for the unfortunate generic
mix-up made by De Selys in 1850 and 1854, events have taken place
smoothly enough. There have been described two species of North
American Progomphus-obscurus and borealis. The former represent-
ed by two female specimens, one in the Museum of Vienna, the other
in De Selys' hands, both specimens recorded simply from North
America by both De Selys and Hagen; the latter species represented
by a single male from Oregon, the specimen apparently in the hands
of MacLachlan. The nymph has been described from Massachusetts
by Cabot. The next year, however, Hagen published a paper that
greatly complicated the situation.
1874. Hagen described a male Progomphus from an adult speci-
men and "Abbott's drawings" in the British Museum, the specimen


having been taken in Georgia. He assigned the name Progomphus
borealis MacLachlan to this specimen and wrote (1874: 356), "The
discovery of this genus, hitherto not represented in the fauna of
North America, is very interesting." This statement is hard to explain
in view of the De Selys (1858) and Hagen (1861) records of female
Progomphus obscurus "Habitat: North America." Yet, that Hagen was
saying what he wanted to say is indicated by a statement in the intro-
duction to this same (1874) paper, "I was able directly to recognize
[from drawings of wing venation] a genus which I had never seen from
North America before, and of which only a short time ago a species was
described from Oregon by Baron de Selys Longchamps. As I have had
the pleasure since to receive from Mr. Uhler a specimen from Georgia,
the verification of this interesting drawing was confirmed." [page
350]. Thinking that the specimen and drawing that he had was P.
borealis, perhaps what Hagen wanted to say was: The discovery of this
species (not genus) from eastern North America is very interesting. He
continued his discussion under P. borealis [page 356] as follows, "The
only male described by de Selys is from Oregon, and I have not seen
it, but the description agrees; nevertheless, as the specific differences
in Progomphus species are rather obscure, and as the two localities,
Oregon and Georgia, are very distant, perhaps the species from the lat-
ter State may prove distinct." Why Hagen did not think to compare
his Georgia material with P. obscurus is hard to say unless he asso-
ciated P. obscurus with the Progomphus fauna of Mexico and Central
America; thus leaving "North America" in this paper to represent a
more circumscribed area than in his 1861 publication. Hagen continues
his discussion, "All that I can say now, is that I am unable to find
differences between P. borealis and Abbott's species. It is worthy of
notice, that Mr. Cabot in his monograph of the immature state of the
Gomphina, p. 6 [Cabot 1872] has described nymphae from Wareham,
Mass., supposed to belong to a species of Progomphus." Dr. Hagen
follows these remarks with a complete description of a male Pro-
gomphus from Georgia, the specimen being "in Mr. J. Ridings collec-
tion." This description indicates that this specimen was P. obscurus.
In Hagen's mind, at this date, therefore, there was only one species
of Progomphus in North America north of Mexico, namely P. borealis
from Oregon and Georgia with the possibility of larvae from Massa-
The next year, in his Synopsis of the Odonata of America, Hagen
listed (1875: 48) both Progomphus obscurus and Progomphus borealis.
Under the former he said. "Hab.: North America. A female from
Boston, Mass., July; may belong to P. obscurus." However under P.
borealis he wrote, "Hab.: Oregon; Georgia; Dallas, Texas. I have seen


two males from Georgia and Texas and the figures of male and female
by Abbott." Note that the specimens that Hagen has seen have all
been from the east, i. e., within the range of P. obscurus as we know
it today (Dallas is on the western boundary of this range). Hagen
continued, "MacLachlan states that the two small teeth before the tip
of the superior appendage are present there is no further doubt
about the identity [synonymy] of the males from Oregon and Georgia."
In 1878, when De Selys issued his 4th addition to the Synopsis of the
Gomphines, he stated that, "Hagen has written to me recommending
that borealis be made identical with obscurus of Rambur. After a de-
tailed examination of the material in my collection (type with head and
abdomen missing), I am tempted to believe that the two species named
should be united. At the time M. MacLachlan described borealis we
had in front of us only two specimens of young males, in each of which
the brown mark of the origin of the sectors of the arculus was not
distinct. However, we now have full adults in which this condition
varies." De Selys includes a description of a female in this paper and
gives the habitat of the species P. obscurus (synonym P. borealis) as
Georgia, Dallas (Texas), Boston and Oregon. So at this date we still
have only one species of Progomphus north of Mexico but its name is
now P. obscurus-P. borealis has been relegated to synonymy. When
and how Hagen "discovered" P. obscurus is not in the literature.
The tendency of writers after 1878 was to accept De Selys' judg-
ment and to refer to all specimens of Progomphus taken north of
Mexico as P. obscurus. However, in the following years P. borealis
gradually worked back into the literature as a good species. It is of
significance to note that in all probability Hagen and perhaps also
De Selys, never saw a specimen of the real P. borealis until circa 1880.
In 1885, Hagen discussed the nymphs of Progomphus in his paper on
the immature form of the Odonata. In his work he includes the diag-
nostic characters of the Progomphus nymph, a description of the
nymph of P. obscurus from Wareham, Mass., and notes on two other
Progomphus nymphs designated as Nos. 31 and 32 with no species
name attached. Of the P. obscurus material he says, "As P. obscurus
has now been collected near Boston, by Mr. Morrison, there can scarcely
be a doubt that the nymphae belong to this species." It is his note
under "32. Progomphus species" that is of special interest, "Six
nymphae in alcohol young and some probably full grown from Santiago,
Cala., . Length 20-27 mm., breadth 7 mm. These nymphae are
so similar to P. obscurus that it is difficult to separate them: but all
have lateral spines on segments 3 to 9 instead of 5 to 9. The black
spots above the abdomen are larger; the ventral segments 6-10 have
transversal black bands, enlarged to the lateral margins; the abdomen


a little broader. Perhaps the nymphae belong to P. zonatus or to P.
meridionalis n. sp. from Arizona, as a similar nymph 24 mm. long was
collected by R. Crotch, San Diego, Cala.'
The nymphs that Hagen described above (1885, No. 32) are un-
doubtedly those of P. borealis for he has included in his description
most of the special characters of borealis as worked out by Kennedy
(1917b). Also his remark that, "Perhaps these nymphae belong to
P. meridionalis n. sp., from Arizona" indicates that Hagen, on encoun-
tering his first specimen of P. borealis thought it was a new species and
named it P. meridionalis. Hagen never published a description of P.
meridionalis. However, Calvert (1901 B. C. A.) says, "I doubt whether
Hagen recognized the true borealis, especially since he labelled the
above listed material [of P. borealis as determined by Calvert] from
Arizona in the M. C. Z. meridionalis."
The next few years saw no important changes in this situation.
However, in 1905, Calvert, dealing with the genus in the Biologia
Central Americana, wrote, "Although Selys and Hagen, in their latest
paper, regarded obscurus and borealis as identical, I have ventured to
separate them again, the differential characters drawn from the carina
of the male superior appendages being herewith presented for the
first time." Dr. Calvert continued with the remarks regarding P. merid-
ionalis quoted above and gave diagnostic descriptions of both ob-
scurus and borealis. However, in separating these two species, Cal-
vert went only part way; he made subspecies-Progomphus obscurus
obscurus and Progomphus obscurus borealis.
In 1910, Muttkowski, Catalogue of the Odonata of North America,
follows Calvert and lists borealis as a subspecies of obscurus. During
these years however, authors generally followed De Selys and spoke of
Progomphus obscurus as the only member of the genus in North Amer-
ica north of Mexico; however, Williamson (1914) used P. borealis for
the western form.
C. H. Kennedy (1917b), writing on the Dragonflies of California
and Nevada, stated, "I believe the Pacific coast form (of P. obscurus)
to be a distinct species, so it would have to be called Progomphus
borealis MacLachlan. It differs in the imago in being larger than ob-
scurus and in having a single row of denticles on the inferior side of the
male superior appendage. In obscurus, as pointed out by Calvert, the
posterior end of this row is double or treble." Dr. Kennedy goes on to
show that, "The differences between the two species are more con-
spicuous in the nymph than in the adult."
Williamson (1920a), in listing the described species of Progomphus
says, "The elevation of borealis to its proper specific rank makes the
number twelve . "


Most of the authors writing between 1905-1929 were dealing with the
eastern species and so used P. obscurus correctly as a species name.
Kennedy (1921) used P. borealis, Seeman (1927) used P. obscurus in
referring to the western species.
In 1929, Needham and Haywood placed P. borealis, once again, as
a synonym of P. obscurus but added in a footnote, "Perhaps the
west coast G. borealis [sic] of MacLachlan should rank as a distinct
species. Kennedy thought so and gave some good reasons." From this
date on authors dealing with the western species have used the name
that they prefer.
A curious departure in naming arose when Munchberg (1932) used
the name P. dorsalis to refer to P. borealis. Where the name came
from I do not know.
The present writer, after working for some time on the taxonomic
characters of the various species of Progomphus, the results of which
are contained in this paper, has reached the same conclusion that
Kennedy (1917b) and Williamson (1920a) arrived at, namely that
P. borealis and P. obscurus are two separate and distinct species and
should be regarded as such. Furthermore, the foregoing historical survey
makes it apparent that these two species would never have been syn-
onymized if it had not been for the unfortunate misconceptions of
Hagen (1874, 1875, 1885) and the dearth of material in the hands
of De Selys (1878).
The distribution of the three species of Progomphus occurring in
North America north of Mexico, with limiting-dates of capture, are
given below; adult and nymphal records have not been separately indi-
cated. The data have been derived from two sources, (1) literature-
reference to which is given after each set of state records, (2) material
studied for the preparation of this paper, including records from ma-
terial in the Museum of Zoology, University of Michigan, not studied
in detail by the author-see material studied (page 66) for these
records. There is some overlapping of data as derived from these
sources, as much of the material in the Michigan collection has been
previously published on; also, as authors tend to pass records from
one to another, year following year, through the literature.
PROGolPHUS oBscrRus.-The eastern North American species.
Distributed from central Florida north along the Atlantic seaboard
to the region of Boston, Mass., west through northern Michigan to
Iowa and Kansas, south to eastern Texas; on the wing from February
(in Florida) to August. Detailed distributional data as follows:
FLORIDA.-Alachua, Jackson, Liberty Cos.-Feb. 2-July 12. Lit.


ref.: Byers (1930, 1934a); Needham (1897) this record is in all prob-
ability one for P. alachuensis n. sp.; Calvert (1901) after Needham.
GEoRGIA.-Johnson, Wayne Cos., June. Lit. ref: Hagen (1874),
"A specimen from Mr. Uhler" under name P. borealis; De Selys (1878)
after Hagen; Calvert (1901) after Hagen; Byers (1931); Kennedy
ALABAMA.-MObile Co. (Naveco),-February (nymphs). Lit. ref.:
Wright (1939).
LOUISIANA.-Covington (Bogue Chitto River, Sulphur Springs), Pol-
lock (Big Creek).-March-July (nymphs). Lit. ref.: Wright (1939).
NORTH CAROLINA.-Walnut Creek, Pool's Bridge, Lake Ellis, South-
ern Pines, Pine Bluff, Fuquay Springs-May-July. Lit. ref.: Brimley
(1903, 1908, 1918, 1938): Calvert (1901) after Brimley. Brimley
(1938) gives the distribution as, "Raleigh, Southern Pines and east-
TENNESSEE.-Jellico (Cumberland River), Parker's Lake.-June-
July. Lit. ref.: Wilson (1912): Wright (1938) after Wilson.
KENTUCKY.-Livingston, Walthersville (Powell Co.), Pineville
(Straight Creek), Parker's Lake-July-August. Lit. ref.: Williamson
(1905); Calvert (1901) after Williamson; Garman, H. (1924).
NEW YoRK.-Wading River, Deep Pond-June-August. Lit. ref.:
Needham (1901) says, "The genus Progomphus has not been found in
New York state, but it will probably be eventually. It ranges from
Massachusetts southward and westward across the continent, a single
P. obscurus Seyls [sic], having been taken in the whole of northeastern
United States."; Davis (1913); Needham (1926) after Davis.
NEW JERSEY.-Lakehurst, Brown's Mills, Bamber-May-July. Lit.
ref.: Calvert (1910); Davis (1913); Montgomery (1933). Calvert
(1923) states (in relation to this species), ". of limited occurrence
in New Jersey."
NEW ENGLAND.-P. obscurus is apparently distributed along the At-
lantic seaboard in New York and New Jersey (no records for inland
New York State) and reaches its northern limit around Boston, the
records from the Boston area being the only ones for this species from
the whole of New England. For the New England area, Howe (1917)
records, "Very rare-June 5 to Aug. 9."; Garman, P. (1927), "New
England-June 5-Aug. 9."; Howe (1921) includes P. obscurus as part
of the "Cape Fauna" and says, "The following thirteen typically south-
ern species [including P. obscurus] occur only along the coast as far
as Provincetown, and rarely Cape Ann;" for obscurus he adds "Trans-
continental, summer,-largely confined to the outwash ponds of the
MASSACHUSETTS.-Boston, Wareham, Chatham-South Orleans-
June-July. Lit. ref.: Hagen (1875, 1885); Calvert (1901, 1905); Howe


Howe notes, "South Orleans ... Recorded only from Boston and Ware-
ham before, this being the third station in New England."
PENNSYLVANIA.-Allegheny Co., Ohio Pyle-May-June. Lit. ref.:
Williamson (1899, 1902); Calvert (1901) after Williamson. Note:
These records are from the southwestern part of the state not far from
the Ohio line. P. obscurus has not been recorded from central or eastern
Omo.-Gallia Co. (Vinton), Lawrence Co. (Ironton)-May 12-Aug-
ust 11. Lit. ref.: Hine (1900, 1913); Williamson (1899); Calvert
(1901); Borror (1935, 1937). Notes: These records are from the lower
south-east tip of Ohio near the West Virginia-Kentucky line. The
literature records seem to be all based on the early work of Hine.
INDIANA.-Allen, Cass, Dearborn, Elkhart, Gibson, Huntington,
Jasper, Kosciusko, Lagrange, Miami, Noble, Parke, Posey, Putnam,
Steuben, Warren, Wells, Whitley Cos.-June 2-August 26. Lit. ref.:
Williamson (1899, 1901, 1917, 1920c); Kennedy (1902, 1928); Need-
ham and Hart (1901); Calvert (1901); Montgomery (1925: Pts. 1,
3, 4, 7, 8, 9). Note: These counties are well distributed over the state
and some of them have been intensively worked (Wells Co. for ex-
MIcHIGAN.-Cheboygan, Emmet, Lake, Washtenaw-June 28-Aug-
ust 1. Lit. ref.: O'Brien (1911); Evans (1915); Byers (1925, 1927).
Note: The Douglas Lake records (Cheboygan and Emmet Cos.) rep-
resent the farthest point north that P. obscurus has been taken.
WISCONSIN.-No records. Muttkowski (1908) predicts its occur-
rence in the southern part of the state.
ILLINoIS.-Effingham (Little Wabash River), Spoon River-June-
July. Lit. ref.: Hagen (1885); Williamson (1899); Needham and
Hart (1901); Calvert (1901). Needham and Hart state, "The nymph
of this species .... has not been found in the Illinois or its adjacent
lakes, but seems common in the sand of the smaller rivers, such as the
Spoon River and upper Sangamon . The adults are rare in col-
lections although the species seems widely distributed. The adult has
been taken in Illinois June 29 to July 13."
IOwA.-Henry Co. Hoffman (1924).
MissouRI.-Carter, Stoddard (Dexter Cos.)-June-July. Lit. ref.:
Williamson (1932).
ARKANSAS.-Lincoln Co.-July. No literature references.
KANSAS.-Clark, Hamilton, Harper, Logan, Ness, Osborne, Rooks,
Russell, Trego Cos.-June-July. Lit. ref.: Calvert (1901); Kennedy
(1917a). Kennedy writes, "Eastern species which cross Kansas and
reach their western limit somewhere on the plains of eastern Colorado."
Again in the same paper he says, "An eastern species, found only on


streams whose beds are loose, flowing sand, in which its highly special-
ized nymph burrows. All the Kansas records except Harper county
(south central) are from the western end of the state, where sandy
streams abound."
OKLAHOMA.-Alfalfa, Beckham, Canadian, Cleveland, Comanche,
Cotton, Greer, Hughes, Kiowa, Latimer, Love, Major, Oklahoma, Pon-
totoc, Pushmataha, Tillman, Woods Cos. lit. ref.: Bird (1932).
TEXAS.-Dallas, Victoria (Colleto Creek.)-July. Lit. ref.: Hagen
(1875); De Selys (1878); Calvert (1901); Kennedy (1921). Ken-
nedy remarks, "It is interesting to those studying distribution to find
that these obscurus naiads from Texas, where both obscurus and bore-
alis are found, are in no wise intermediate in character between the two
species but are true obscurus naiads." Note: The above locations are in
eastern Texas.
PROGOMPHUS BOREALIS.-The western North American species. Dis-
tributed from northwestern Mexico and Baja (Lower) California
north to Oregon and east to Colorado, New Mexico and western Texas;
on the wing from April to October. Detailed distributional data as
TEXAS.-Presidio Co. (Shafters, Marfa), Jeff Davis Co. (Phantom
Lake)-June-September. Lit. ref.: Kennedy (1921); Tinkham (1934).
Note: These two counties are in the extreme southwestern corner of
NEW MEXIco.-Albuquerque.-April 18, 1903 (a general female
specimen in University of Michigan collection).
COLORADO.-No definite record for P. borealis. A. D. Hess, in a
Master's Thesis from Colorado College, 1933, writing on the "Dragon-
flies of the Pikes Peak Region" records P. obscurus from the "Foothill
Zone, 5,800-8,000 feet." From the locality this is probably P. borealis,
though Kennedy (1917a) states that the true P. obscurus reaches its
western limit ". somewhere on the plains of eastern Colorado."
UTAH.-Washington Co. (St. George)-June 6, 1919. Lit. ref.:
Brown (1934).
ARIZONA.-Catalina Mts. (Pima Co.); Granite Reef Dam, Mesa,
Salt River, Gila Bend (Maricopa Co.); Hot Springs, Jerome, Pres-
cott (Yavapai Co.); Florence (Pinal Co.); Ft. Grant, Graham Mts.
(Graham Co.); Bright Angel, Colorado Canyon-April-September.
Lit. ref.: Currie (1903); Calvert (1901); Williamson (1914); Tink-
ham (1934).
CALIFORNIA.-Los Angeles, Santiago, Santa Ana River, Palm Springs,
Chico, Oroville, San Jose-June. Lit. ref.: Calvert (1895, 1901); Ken-
nedy (1917b); Seeman (1927); Needham and Heywood (1929).


OREGON.-The type locality for the species. Lit. ref.: MacLachlan
(De Selys 1873); Hagen (1874, 1875) after De Selys; Calvert (1895,
1901) after De Selys. As nearly as I can determine, no specimen of P.
borealis has recorded from Oregon since MacLachlan's type was sent
by Lord Walsingham (De Selys 1873).
MExIco.-Baja California (Purissima); Sonora (Hermosillo); Chi-
huahua (Santa Rosalia); Jalisco (Guadalajara); Michoacan (Uru-
apan)-April-October. Lit. ref.: Calvert (1895, 1901); Williamson and
Williamson (1930).
PROGOMPHUS ALACHUENSIs, n. sp.-The Florida species. Distribu-
tion restricted to central Florida; on the wing from February to May.
FLORIDA.-Clay, Alachua, Orange, Volusia, Lake Cos.-Feb.-May.
Lit. ref.: (under name P. obscurus) Byers (1930 in part); Davis and
Fluno (1938); Needham (1897), in all probability Needham's speci-
men from Orange Co. is of this species.
Florida.-The ranges of P. obscurus and P. alachuensis overlap in
Florida. In Alachua County-the northern extent for alachuensis and the
southern for obscurus-both species are part of the fauna, alachuensis
occupying the lakes and obscurus the cooler streams (see discussion
page 56). It is also possible that P. integer, from Cuba, may turn up
some time in the southern part of the state.
Texas.-The ranges of P. obscurus and P. borealis meet in Texas
(Kennedy 1921) and, in all probability, in Colorado also (Kennedy
1917a). Progomphus has been reported from the latter state but which
species the writer is not certain.
Mexico.-Mexico has a sizable Progomphus fauna in addition to P.
borealis which has been reported as far south as Uruapan. The ranges
of P. zonotus and P. clendoni overlap that of P. borealis along its south-
ern limit. P. pygmaeus and P. risi occur farther south (Guatemala).
A study of the foregoing records indicates that the flying season of
the North American Progomphus varies considerably. Over most
of the territory, these dragonflies may be considered as part of the sum-
mer fauna-June to early August with the peak in mid-July. How-
ever, in Florida both P. obscurus and P. alachuensis are spring species
-February to June (occasionally July) with the peak in May. Farther
north (Georgia and Carolinas) the season apparently moves forward
somewhat-May to July with the peak in June. On the other hand,
P. borealis may emerge as early as April (Mexico, New Mexico, Ari-
zona) and remain on the wing as late as September (Arizona) or even
October (Mexico), thus becoming part of the autumn fauna in the


SYNONYMY. The following authors have used the following names in
referring to this species:
Diastatomma obscura Rambur.-Rambur (1842).
Gomphoides obscura (Rambur)-Brimley (1938); Byers (1930);
Cowley (1934); Davis (1913); Hine (1913); Muttkowski
(1910a); Wilson (1912); Selys (1850).
Progomphus obscurus (Rambur)-Bird (1932); Borror (1935);
Brimley (1903, 1908, 1918); Beutenmuller (1890); Byers
(1925, 1927, 1931, 1934a, 1934b); Butler (1904); Cabot
(1872); Calvert (1901, 1905, 1910, 1923); Evans (1915); Gar-
man, H. (1924); Garman, P. (1927); Hagen (1861, 1875,
1885); Hine (1900); Hoffman (1924); Howe (1917, 1919,
1920, 1921); Karsch (1890); Kennedy (1902, 1917a, 1917b,
1921, 1928); Kirby (1890); Lew (1934); Montgomery (1925,
1933); Muttkowski (1908); Needham (1901, 1903a, 1903b,
1926); Needham and Hart (1901); Needham and Anthony
(1903); Needham and Heywood (1929); O'Brien (1911); Ris
(1908, 1911); Selys (1854, 1858, 1878 pars); Williamson (1899,
1901, 1902, 1905, 1917, 1920c, 1932); Wright (1938, 1939);
St. Quentin (1938).
Progomphus obscurus Selys.-Needham (1901).
Progomphus obscurus obscurus (Rambur).-Calvert (1901).
Progomphus borealis MacLachlan.-Hagen (1874, 1875).
Negomphoides obscura Rambur.-Borror (1937).
DESCRIPTIONS. Satisfactory descriptions have been written by the fol-
lowing authors:
De Selys (1858: 202-205). Gives earliest detailed description, fe-
male only.
Hagen (1874: 356-358). Complete description of male, from Georgia
specimen and Abbott's drawings, under name of P. borealis.
Williamson (1899: 283). Brief description of male and female.
Calvert (1901: 148-149). Presents diagnostic and "key" characters,
some of them new.
P. Garman (1927: 126-127). Compact description.
Needham and Heywood (1929: 62-63). Brief description, key char-
acters and notes (drawings are of P. borealis).
Byers (1930: 48-50). Description of male, female and nymph.
Figures-The following have published figures of this species:
Wings: P. Garman, (1927: P1. X, fig. 1); Needham (1903a: P1. 34,
fig. 2).


Head: Williamson (1899: P1. IV, figs. 3-4).
General: Needham and Anthony (1903: Thorax); Butler (1904:
labium); Lew (1934: head); St. Quentin (1938: legs); Kennedy
(1917a: P1. I, fig. 14-18), have made morphological studies of the
Odonata including this group.
Head (fig 1. P1. I).-Top of frons semi-angular with a median in-
dentation, 3.8-4.0 mm. in length; hind margin of occiput concave,
depth moderate.
Thorax (fig. 3. P1. III).-Pale humeral stripe broad, joined to the
pale area of the mesotharacic collar, edges straight and parallel, dorsal
area between dark and medium brown (not light). Pale antehumeral
stripe variable (see discussion p. 43) usually present as a fine, but dis-
tinct, stripe separated from the pale spot above the antealar carina.
Abdomen.-Obscure reddish-brown. Paler dorsal pattern on seg-
ments 3-7 reduced and not conspicuous. Segments 8-10 usually solid
brown, occasionally with pale yellow wash along sides; no distinct
yellow spots or bands. No tubercle present on the venter of segment
1; a median spine present in the males. Vulvar lamina of females
(fig. 1. P1. II) with a V-shaped central notch.
Appendages male (fig. 4. P1. II).-Superior abdominal appendages
short (2-2.5 mm.) blade-like, lower margin curving to the tips in
lateral view (fig. 4. P1. I), inferior margin with two or more rows
of denticles apically, the carina not extending to the last denticle;
color bright yellow. Inferior appendages black, comparatively short and
Wings.-Brown basal stripe of costal and subcostal spaces reaching
to the first antenodal cross-vein or nearly so; usually a trace of brown
between the sectors of the arculus at their origin (may be absent in
a small number of specimens-usually western ones). Stigma reddish-
brown, 4 mm. long in males, 5 mm. long in females. For variation
in wing venation see analysis given below.
Size.-Medium large, abdomen 36-38 mm., hind wings 31-34 mm.
A study of variation, based on the material examined (see page 66),
gave the following results:
Wing venation.-Venation, based on 128 fore and hind wings and
12 9 fore and hind wings (64 a and 12 9 specimens) varied as follows:



_______ ~ 2$ 8 $
Anq of fore wings 13-19 15-18 15.30 16.62
Anq of hind wings 5-14 11-15 11.45 12.13
Pnq of fore wings 6-12 9-13 9.49 10.42
Pnq of hind wings 7-13 9-13 9.91 10.62
Scq in all wings 1 except left fore wing and right and left hind
wing of one male which had none.
Cuq in all wings 1 except 6 $ fore wings, 2 S hind wings, 1 9 hind
wing in all of which 2 were present.
.Distal thickened Anq 2nd in 1 wing 5th in 255 wings
3rd in 2 wings 6th in 37 wings
4th in 8 wings 7th in 1 wing

Fore- Hind- Fore- Hind-
wing wing wing wing Fore wings Hind wings
1 celled 0 0 0 0
2 celled 0 98 1 11 43 8 5
3 celled 126 27 23 12 62 119 6 49
4 celled 1 3 0 1
1 celled 1 28 0 2 8 1
2 celled 124 98 23 18 60 119 42 S 7 9
3 celled 2 2 1 4
1 celled 127 123 24 22 63 S 12 61 S 119
2 celled 0 5 0 2 __ 2 2 19

Stigma covers cells: Male fore wing 4-8 (aver. 5.99); hind wing 4-8
(aver. 6.01). Female fore wing 6-9 (aver. 7.09); hind wing 5-9
(aver. 6.99).
Anal field of fore wing proximal to the triangle 2 cells wide for a
distance of from 4-6 cells (average approximately 5), these may
be so irregularly arranged as to make counting difficult.
Anal triangle of male: 2 celled in 1 wing; 3 celled in 108 wings;
4 celled in 19 wings.
MEASUREMENTS6 (in millimeters).-Abdomen (including append-
ages) male 35-41 (aver. 36.92); female 35-41 (aver. 38.77). Length

6Montgomery (1933) collected four males near Brown's Mills, N. J. of which
he states, "These specimens which measure 46, 47, 47, and 48 mm. in length,
are considerably smaller than other specimens of this species which I have


of hind wing male 30-33 (aver. 31.38); female 32-37 (aver. 34.28).
Stigma length male 4-5 (aver. 4.15); female 5-5.5 (aver. 5.04).
VARIATION IN GEOGRAPHIC. GROUPS.-For the purpose of studying
regional variation within the species, the above data and the specimens
upon which they were based were broken down into three geographic
Group 1= Florida, Georgia, Alabama (25 specimens)
Group 2=Kentucky, Indiana, Michigan (10 specimens)
Group 3=Arkansas, Missouri (41 specimens)
In general, there were no constant differences between groups
sufficiently marked to warrant geographic races. However, certain
tendencies were noted:
Group 1. The only group in which triangles of 4 cells appear.
Also, the group in which the anal triangle of the male is most apt to
show the 4-celled condition.
Groups 1 and 2. Have somewhat smaller bodies, larger wings and,
in general, darker color than group 3. These features are more marked
in group 1 than in group 2.
Group 3. The western specimens are paler brown than the eastern;
the brown dorsal area on the thorax between the pale humeral stripes
markedly so. In this group there is a strong tendency for the
subtriangle of the hind wing to be open (1 cell-no cross-veins). On the
whole, the western specimens have larger bodies and slightly smaller
wings than have the eastern ones.
is a marked and graded variation in the degree of development of
the dorsal thoracic pattern (pale humeral and antehumeral stripes
and the surrounding dark areas).7
This graded variation is most noticeable in observing the pale ante-
humeral stripe throughout the series. The behavior of this stripe
may be likened to the development of a photographic negative-
at first the plate is uniform and then slowly the picture appears,
the high-lights first. The series studied thus may be arranged, on
the basis of the antehumeral stripe, into an arbitary set of six grades
as follows:
Grade 1. Pale antehumeral stripe absent; pale antealar spots faint-
ly indicated (fig. 2. P1. III).
Grade 2. Pale antehumeral stripes present but very faint; pale
antealar spots present, separated from antehumeral stripe.
7Also, this holds true when a series of specimens of P. borealis and P. alachu-
ensis are studied along with P. obscurus.


Grade 3. Pale antehumeral stripe present, thin, distinct, separ-
ated from pale antealar spots (fig. 4. P1. III).
Grade 4. Pale antehumeral stripe present, fairly wide, distinct, ap-
proaching union with the pale antealar spots.
Grade 5. Pale antehumeral stripe wide (0.5 mm.), distinctly joined
with the pale antealar spots (fig. 3. P1. III).
Grade 6. Pale antehumeral stripe quite wide (0.7 mm.), broadly
joined with the pale antealer spots (fig. 5. P1. III).
The following table (including P. alachuensis, P. borealis and the
three geographic groups of P. obscurus given above), based on the
six pattern grades, shows the amount and nature of this antehumeral

P. obscurus-group 1 25 3 6 11 5 0 0
P. obscurus-group 2 10 0 0 6 1 3 0
P. obscurus--group 3 41 0 3 31 3 4 0
P. borealis 12 0 0 0 0 0 12
Total all 115 27 12 48 9 7 12
Total P. obscurus 76 3 9 48 9 7 0
The remainder of the thoracic pattern, while showing some varia-
tion, is not as definite as the above. There is a gradual tendency,
already noted, for the colors to become lighter from southeast to
west and for the humeral stripe to become broader and more dis-
tinctly joined to the pale area of the mesothoracic collar.
ECOLOGICAL NOTES:-Information on the habits, frequency and
special peculiarities of P. obscurus may be gathered from a reading of
published material covering various portions of the species' range.
Brimley (1903: North Carolina) states, "Fairly common in summer
in sunny spots along the banks of Walnut Creek usually resting,
when not in flight, on bare patches of sand or mud close to the water's
edge . This species was also observed at Poole's bridge .
resting, when not on the wing, on bare rocks projecting above the
surface of the water."
Byers (1925: Michigan) writes, "Common along sandy beaches of
Douglas and Burt Lakes." However, for Florida, Byers (1930) notes,
"Very little is known of the adult habits of this species in Florida.
Certainly it is a very shy, retiring, woods-loving form. In spite
of the very common occurrence of the nymphs, only four adults
have appeared in collections examined." I may add that, at the pres-
ent date, more Florida material is available (see material studied


p. 66); also, that the adults are not so rare if looked for at the right
time and place-central Florida lake shores in May.
Davis (1913: New Jersey) states, "Found on several occasions in
May and June flying up and down a shaded ditch by the side of the
railroad track at Lakehurst, N. J." Needham and Heywood (1929)
reprint Prof. Davis' observation. Montgomery (1933), also collecting
in New Jersey, reports specimens from, "Along a stream in the pine
barrens near Brown's Mills."
Howe (1919: Massachusetts) writes, "Not uncommon in oak scrub
Mr. Bangs tells me."; again (1917), in the Manual of the Odonata
of New England, he notes, "Very rare-scrub oak uplands, rivers and
lakes"; later in the Manual, Howe adds "ponds" to the foregoing
Kennedy (1917a: Kansas) notes, "Progomphus obscurus occurs
wherever in the east there are sand bedded streams, as its nymph
is adapted to rapid burrowing in flowing sand; consequently it is
abundant throughout western Kansas."; again in the same paper,
"An eastern species, found only on streams whose beds are loose
flowing sand .."
Both Williamson and Montgomery have recorded observations of
P. obscurus in Indiana. Williamson (1899) states, "Not rare at the
ripples [St. Mary's River, Ft. Wayne?], a pair taken resting on a
boulder in mid-stream." In later years, Williamson (1920c) writes
of an interesting case as follows, "Salamonie River, Huntington Co.,
.... several caught and others seen, many had wings darkly stained
with crude oil; Wabash River, Bluffton, Wells Co . only one
seen, a male with oil stained wings. At this time there was no crude
oil along the Wabash River near Bluffton, and I think the Wabash
specimen was undoubtedly a stray from the Salamonie about twenty
miles away." Montgomery (1925: Pt. 1) writes, "A general female
[taken near Vincennes] was resting on the tip of a dead weed a
few feet from a gravel pit in the manner of Celethemis elisa." Again,
(1925: Pt. 3) Montgomery quotes Williamson (unpublished), "A
male taken along stream south of Goshen . Stream 12-20 feet
wide, swift and deep. Several seen, resting on logs in stream. This
individual flew along stream and lighted on vertical twig in a dead
cornus bush-only time I ever saw obscurus on a twig."
Wilson (1912: Tennessee) states, "This species was very common
on the Clear Fork of the Cumberland River at Jellico . a dozen
of each sex were obtained . Both sexes rest much of the time
upon the sand of the river bank in bright sunshine. They are easy to
approach and capture at such times and are not as swift flyers as
many of the gomphines." The present writer's experience in collecting


P. obscurus in Wayne and Johnson Cos., Georgia tends to confirm
Wilson's observations in Tennessee.
Williamson (1932: Missouri) notes, "Progomphus sat on twigs,
weed stems, and, preferably, flat on sand bars [stream west of
Dexter, Stoddard Co.]".
One adult habit, that I have noticed in both P. obscurus and P.
alachuensis, has not been mentioned by any of the foregoing authors.
That is, when the adult is at rest, with the feet on a flat surface, the
body assumes a position in which the thorax and abdomen are tilted
away from the plane of the resting surface at a marked angle (45-60)
making the insect appear as if it were standing on its head. Calvert
(1901) and Williamson (1914) have observed this habit in P. borealis.
In regard to the habits of P. obscurus it may be deduced from the
above material that:
1) The species, over most of its range, is not rare and may fre-
quently occur in numbers large enough to deserve "not uncommon."
2) It is most intimately associated with sand bottomed streams,
lakes or even pools where its larva may find burrowing conditions.
3). It usually rests on some low projecting object (rock, etc.),
or simply on the ground close to the water's edge; but may, on oc-
casion, alight on twigs, low bushes, etc. When resting on the ground
it assumes the characteristic tilted position.
4) The species tends to be associated with woods (oak scrub, pine
barrens, hammocks) where it flies in the shade or rests in open sunny
spots near water.
5) It is shy, but not a fast flier or unusually difficult to capture.
SYNONYMY. The following authors have used the following names
in referring to this species:
Gomphoides borealis (McLachlan).-Muttkowski (1910a); Need-
ham and Heywood (1929: foot-note).
Progomphus borealis (MacLachlan).- Brown (1934); Byers
(1934b); Hagen (1875); Kennedy (1917b, 1921); Selys (1873-
MacLachlan); Williamson (1914, 1920a); Williamson and
Williamson (1930).
Progomphus obscurus (Rambur).-Calvert (1895); Currie
(1903); Kirby (1890); Needham and Heywood (1929);
Selys (1878); Seeman (1927); Tinkham (1934).
Progomphus obscurus borealis MacLachlan.-Calvert (1901).
Progomphus dorsalis.-Munchberg (1932).
Progomphus meridionalis [Nomen nudum of Hagen].-Hagen
(1885); Calvert (1901).


Progomphus No. 32.-Hagen (1885).
DESCRIPTIONS. Satisfactory descriptions have been written by the
following authors:
De Selys (1873: 764-765). The original description of the male
sent to De Selys by MacLachlan.
De Selys (1878: 471). Description of female (in part) and note
on synonymy.
Calvert (1895: 499-502). Detailed description of male and female.
Calvert (1901: 148-149). Presents diagnostic and "key" characters.
Kennedy (1917b: 524-529). Good description.
Figures-The following have published figures of this species:
Calvert (1895: P1. XVI, figs. 74-79). Male genitalia (3 figs.),
female vulva, triangle and adjacent parts of right fore wing.
Kennedy (1917b: figs. 148-154, page 524). Color pattern, seg-
ments nine and ten of female, segment two of male, segment ten
and appendages of male (3 figs.).
Needham and Heywood (1929: page 63). Male genitalia, female
vulva (under name P. obscurus; same figures published by Byers-
1930); also, (page 58), drawing of the basal area of hind wing
(reprinted from Seeman-1927).
Head (fig. 2. P1. I).-Top of frons angular, 4.2 mm. in length; hind
margin of occiput straight with slight median convexity, thick.
Thorax. (fig. 5. P1. III). Pale humeral stripe very broad, joined
broadly to the pale area of the mesothoracic collar, edges straight and
parallel, dorsal area between light in color. Pale antehumeral stripe
narrow, distinct, joined with the pale spot above the antealar carina.
Abdomen.-Segments 3-7 black with dorsal yellow area composed
of a broad basal spot narrowly connected with a more rounded apical
spot which does not reach the apical margin. Segment 8 black with
two basal cuneiform yellow spots on dorsum; sides with a yellow
spot near apical margin. Segment 9 dorsum black, two lateral yellow
spots-one basal (which may be lacking) the other apical. Segment
10 black with a yellow spot below the base of each superior abdom-
inal appendage ( ), or with entire apical half or more yellow (9).8
No tubercule present on venter of segment 1; a median spine present
in males. Vulvar lamina (fig. 2. P1. II) with a broad U-shaped
central notch.

8In one female specimen examined, all of segment 10 was yellow except
for a reduced black median basal line.


Appendages male (fig. 5. Pl. II).-Superior abdominal appendages
long (3 mm.), ends truncate in lateral view (fig. 5. P1. I), a single
row of denticles on the inferior margin, the carina extending to the
last denticle; color bright yellow.
Wings.-Brown basal stripe in costal and subcostal space reduced,
not reaching beyond the basal subcostal (Scq) cross-vein; no trace
of brown at the origin of the sectors of the arculus (may also be
absent in a small percentage of specimens of P. obscurus). Stigma
black, 4 mm. long in males, 5-5.5 mm. long in females. Wings in
general with dense venation (see venation analysis below).
Size.-Large (average length of abdomen 44 mm.), proportions slen-
der. In general, P. borealis is a large, slender, pale colored Progomphus.
A study of variation in wing venation and measurements, based on
the material examined (see page 66), gave the following results. The
tables are based on the study of 18 8 fore and hind wings and 69
fore and hind wings (9 S and 3 9 specimens).
__9 ~ 9
Anq of fore wings 14-17 15-18 14.88 16.83
Anq of hind wings 10-12 11-15 10.88 12.16
Pnq of fore wings 7-10 8-11 8.70 9.60
Pnq of hind wings 8-11 8-12 9.83 9.05
Scq in all wings 1 in all counted except the left fore wing of 1 9
in which 2 were present.
Cuq in all wings 1 in all counted except the right fore wing of 1 S
in which 2 were present.
Distal thickened Anq: 4th in 2 wings; 5th in 42 wings; 6th in 4 wings.
Fore Hind Fore Hind Fore wings Hind wings
wing wing Iwing I wing _____ ______
1 celled (open) 0 0 0 0
2 celled 0 5 0 0 2$
3 celled 18 13 2 4 9 19 6 29
4 celled 0 0 4 1 29
5 celled 0 0 0 1
1 celled 0 6 0 0 2$
2 celled 17 12 1 6 8$ 58 39
3 celled 1 0 5 0 29
1 celled 18 17 6 6 9 39 8 39
2 celled 0 1 0 0


Stigma covers cells: Male fore wing 4-7 (aver. 5.50); hind wing
5-6 (aver. 5.61). Female fore wing 6-8 (aver. 6.75); hind wing 5-8
(aver. 5.83).
Anal field of fore wing proximal to the triangle 2 cells wide for
a distance of from 5-7 cells (approximate average 5), these may
be so irregularly arranged as to make counting difficult.
Anal triangle of male: 3 cells in 16 wings; 4 cells in 2 wings.
MEASUREMENTS (in millimeters).-Abdomen (including appendages)
male 41-47 (aver. 44.18); female 42-47 (aver. 44.00). Length of hind
wing, male 31-35 (aver. 32.88); female 35-38 (aver. 36.00). Length
of stigma male 4 (aver. 4.00); female 5-5.5 (aver. 5.16).
EcOLOGICAL NoTEs.-Information on the habits, frequency and
special peculiarities of P. borealis may be gathered from a reading
of published material covering various portions of the species' range.
Probably P. borealis has been most thoroughly- observed in Cali-
fornia. Writing on the habits of the adult in this State, Kennedy
(1917b) says:
This western Progomphus is apparently widely distributed throughout Cali-
fornia, but during the summer's collecting I have seen not more than ten or
twelve specimens on the wing. Perhaps the expression "widely distributed" should
be used with some qualification as under it I have included the evidence from
the wide distribution of the nymphs . Apparently this species has a long
season on the wing as Grinnell's records are from May to August. On the
Chico River the few individuals I saw were flying about a quiet pool at the
head of a long rapids and when not in motion were seated on snags in the
water or on the gravelly shore. Their habits seem much like Ophiogomphus bison
with which they were associated, except that they were much more wary and
difficult to capture. Their large size, slender abdomen and peculiar brown thorax
readily distinguish them in the field from any other western gomphine.
In this same paper, Kennedy records P. borealis from "An eleva-
tion of 2,500 feet", he also repeats (p. 541) observing it in associ-
ation with Ophiogomphus bison but adds Gomphus sobrinus, and (p.
605) notes again that it is difficult to capture.
Needham (Needham and Heywood, 1929), writes, "In May the
adults were seen along the upper reaches of the Santa Margarita
River [California] They flew swiftly from one resting place
on the bare sand to another and were very inconspicuous when at
rest. They were not especially difficult to approach by stalking,
or to capture when at rest." Writing from this same region, Seeman
(1927) records, "The adult is usually seen flying about the borders
of streams. It may be most easily captured by clapping a net down


over it as it rests on the sand; but one must look closely, for it is a
clear winged creature and inconspicuous when quiet, although it is
easily recognized when caught. .. ."
Tinkham (1934) in writing of this species (he uses the name P.
obscurus) in southwestern Texas states, "The Cibolo creek at Shafter
supported Progomphus obscurus as its commonest species. There
were no adults in the air at this time in late May." To a locality
record in southern Arizona (10 miles east of Gila Bend), Tinkham,
in this same paper, appends, "Very rare and miles from any water."
Williamson (1914) writing about September Dragonflies About
Mesa Arizona, says in regard to P. borealis, "All were taken away
from water 10-40 feet; were resting on the tips of twigs, often
with the abdomen elevated, and none were seen resting on the
ground." Currie (1903) records one male of P. borealis as having
been taken in July at Bright Angel, Colorado Canyon, Arizona, at
an elevation of 2,700 feet.
Notes on P. borealis in Mexico are sparse. Calvert (1901) writes,
"This species and Erpetogomphus crotalinus were found on sand, or
mud-banks, hardly above water-level, along the sides of an irrigation
ditch into which water from the baths of Santa Rosalia empties.
On alighting on these banks, P. obscurus borealis held its abdomen
slanting upward to form an angle of 45-600 with the bank, while E.
crotalinus held it nearly horizontal." In addition to the above, Wil-
liamson and Williamson (1930) mention P. borealis as an associate,
"along this stream [Rio Purissima], of a new Mexican species-
Erpetogomphus natrix.
SYNONYMY. The following authors have applied the name Progom-
phus obscurus or Gomphoides obscurus to this species.
Gomphoides obscurus (Rambur).-Byers (1930, in part): Davis
and Fluno (1938).
Progomphus obscurus (Rambur).-Byers (1934a, in part); Need-
ham (1897, probably).
Holotype Male. Rear of head deep brown above to tan below; two
light tan spots at outer margin of compound eyes. Face and mouth-
parts pale greenish-yellow to buff; mandibles, base of labrum and front
of frons deeper brown. No distinct stripes, bands or spots on face.
Frons above long (4.5 mm.) and narrow (fig. 3, P1. I), light greenish
yellow with a median posterior, ill defined oval, darker brown area.
Compound eyes dark brown. Vertex and antennae chiefly dark brown,


lighter toward inner angles of compound eyes; postocellary ridge
present as a double tubercle behind lateral ocelli. Occiput yellow, deeply
furrowed at base between projecting angles of compound eyes, hind
margin straight, edged with fine hairs (fig. 3 P1. I).
Prothorax dark brown, a large pale irregular spot on lateral mar-
gins of dorsal plate; sides light brown to tan.
Thorax (synthorax) predominantly dark brown, especially on the
dorsum; colors ranging from dark brown to greenish-yellow and gray.
Thoracic pattern (fig. 2. P1. III) as follows: mesothoracic collar yellow,
divided at mid-dorsal carina which is edged with yellow. Pale thoracic
humerall) stripe narrowly separated from pale area of mesothoracic
collar; narrow with jagged edges and tapering ends, color yellow.
Pale antehumeral stripe absent, represented by a faint spot near the
antealar carina. Mesepimeral stripe pale, complete, about 1 mm. wide;
remainder of mesepimeron dark brown. Metepisternum brown becoming
lighter near the second lateral suture. Metepimeron pale greenish-yel-
low except for a brown stripe along the second lateral suture. Mes-
infraepisternum golden brown, lighter near base of coxae. Metinfraepis-
ternum light greenish yellow with brown sutures.
Legs dark to light brown with coxae and knees yellow; front femora
light yellow below, next pair darker, hind femora dark brown below.
Wings clear with a brown basal stripe in costal and subcostal
areas reaching as far as the first antenodal cross-vein or nearly so;
a trace of brown between the sectors of the arculus at their origin;
anal crossing (Cuq) slightly infuscated. Costa yellow; remaining veins
dark brown. Stigma light yellow-brown, 5 mm. long, covering 6-7
cells. Antenodal cross-veins (Anq) of fore wing 15-16; hind wing
10-11. Distal thickened antenodal the fifth. Postnodal cross-veins
(Pnq) of fore wing 8-9; hind wing 9-10. A single basal antenodal
cross-vein of the second series (Scq) present in all four wings. Anal
field of fore wing proximal to the triangle two cells wide for a dis-
tance of five cells. Anal triangle with three cells. Supratriangle open-
no cross-veins. Cells of triangle (T) and subtriangle (Ti) vary in the
four wings as follows:
T 3:3 Ti. 2:2
2:2 2:0
Abdomen, in general, black to brown on sides, yellow markings
on dorsum which become obscure in dried specimens. Segments 1-2,
sides pale green to yellow, dorsum brownish with large pale area on
segment 2; auricles yellow. Segments 3-7, sides black to dark brown,
dorsum with a broad basal yellow area narrowly joined to a more re-
duced subapical yellow spot. Segment 8, sides golden brown, dor-
sum somewhat darker, no distinct pattern. Segments 9-10, practically


uniform dark brown, no yellow spots. Venter of segment 1 with a small
median basal spine; no ventral tubercle. Accessory genitalia or sec-
ond abdominal segment 2 (fig. 7. Pl. I) yellow to dark brown.
Superior abdominal appendages (fig. 6. Pl. I) long (2.8 mm.),
slender and pointed; a single row of denticles on inferior margin
(fig. 6. P1. II) carina extending to the last denticle; tips light yellow,
basal third brown. Inferior appendages and accessories (fig. 6. PI. II)
dark brown to black.
Measurements: Total length 55 mm. Abdomen, including append-
ages, 40 mm. Fore-wing 34 mm. Hind wing 32 mm.
Material: Holotype male. Alachua Co., Florida. Newnan's Lake
(4 miles east of Gainesville). May 18, 1930. Catalogue No. 17;
specimen No. 15. C. F. Byers collection.
Allotype Female. Similar in most respects to the male; head pat-
tern essentially the same. Top of frons oval, 5 mm. long; hind mar-
gin of occiput straight. Thoracic pattern the same; pale humeral
stripe narrowly joined to the pale area of the mesothoracic collar.
Legs similar to male.
Wings colored as in male; an additional reduced brown stripe in
cubital area not reaching to the anal crossing (Cuq), an extra cross-
vein in this area of the left hind wing. Stigma 6 mm. long, covering
7-8 cells. Antenodal cross-veins (Anq) of fore wing 17; hind wing
13-14. Distal thickened antenodal the sixth. Postnodal cross-veins
(Pnq) of fore wing 8-9; hind wing 9-11. A single antenodal cross-
vein of the second series (Scq) present in right fore and hind wing,
lacking in left fore and hind wing. Anal field of fore wing proximal
to the triangle with double cells irregularly arranged. Supratriangle
open-no cross-veins. Cells of triangle (T) and subtriangle (Ti) in
the four wings as follows:
T _3:3 Ti 2:2
T= 3-3 Ti =2
2:2 2:2
Abdomen more robust than in male. Color pattern essentially similar.
Basal pale area and subapical spot on dorsum of segments 3-7 larger,
covering more of the top of the segments. Lateral apical corner of
segment 9 with reduced yellow spot; dorsum of segment 9 and all
of segment 10 dark brown. Appendages conical, 1.5 mm. long-slightly
longer than the dorsum of segment 10, base brown, tips yellow. Vulvar
lamina (fig. 3. P1. II) with deep U-shaped notch, sides of notch
Measurements: Total length 54 mm. Abdomen, including append-
ages, 39 mm. Fore wing 36 mm. Hind wing 35 mm.


Material: Allotype female. Alachua Co., Florida. Newnan's Lake.
May 18, 1930. Catalogue No. 17; specimen No. 19. C. F. Byers
Paratypes. From the material examined the following are desig-
nated as paratypes. All are from Florida. Alachua County: Newnan's
Lake, Gainesville, May 18, 1930, 3 8, C. F. Byers (Byers collection),
Clay County: Gold Head Branch State Park, February 1939, 19,
reared by J. G. Needham (Cornell); Kingsley Lake, May 1935, 6S
3 reared by C. F. Byers (Byers collection). Orange County: Win-
ter Park, 1 3, E. M. Davis (Byers collection). Volusia County: Ben-
son Springs [formerly Enterprise], April 22, 1921, 1 J. H. William-
son (University of Michigan), April 26, 1921, 2 S, J. H. Williamson
(University of Michigan). Lake County: Lake Harris, August 5, 1938,
1 3, Hubbell and Friauf (University of Michigan).
Head (fig. 3. P1. 1).-Top of frons long (4.5 mm.), oval; hind mar-
gin of occiput straight, mediumally thick.
Thorax (fig. 2, Pl. III).-Pale humeral stripe usually separated from
the pale area of the mesothoracic collar, edge jagged, tapering to a
point, dorsal area between dark brown. Pale antehumeral stripe usually
absent, occasionally present as a very faint line. Pale spots above ante-
alar carina present.
Abdomen.-Pale dorsal pattern on segments 3-7 extensive, usually dis-
tinct. Segment 8, sides golden brown, dorsum somewhat darker, no
distinct pattern. Segments 9-10 practically uniform dark brown-no
yellow spots. No tubercles present on venter of segment 1; a median
spine present in males. Vulvar lamina of female (fig. 3. P1. II) with
deep U-shaped central notch.
Appendages male (fig. 6. P1. II).-Superior abdominal appendages
long (2.8 mm.), slender, somewhat pointed apically (fig. 6. P1. 1),
a single row of denticles on inferior margin, the carina extending to
the last denticle; color, base brown, tips yellow. Inferior appendages
slender and pointed.
Wings.-Brown basal stripe in costal and subcostal spaces reaching
to the first antenodal cross-vein or nearly so; a trace of brown be-
tween the sectors of the arculus at their origin. Stigma brown, 5 mm.
long in males, 5-6 mm. long in females. Wing venation not as dense
as in P. borealis (see analysis tables).
Size.-Medium (average length of abdomen 38 mm.), proportions
slender. In general, P. alachuensis is a medium large, slender, dark


VARIATION. A study of the paratypes and the other material exam-
ined (see page 67) indicated the following variations:
In the thoracic pattern, the sides of the humeral stripe exhibit
degrees of serration from those that are quite jagged (fig. 2. Pl. III)
to the condition found in four males in which the sides of this pale
stripe are simply sinuate. In these same four males and an additional
female the pale humeral stripe is narrowly and faintly joined to the
pale area of the mesothoracic collar. In a few specimens studied
(.76%) there is a very faint suggestion of a fine antehumeral stripe
in addition to the faint spot above the antealar carina. In the abdominal
pattern the dorsal areas show some variation in the extent of the
pale markings which is difficult to measure; the females, in general,
have the more extensive light areas.
Venation based on 36 fore and hind wings and 169 fore and hind
wings (18 and 8 9 specimens), varies as follows:
___F I I e 1 I 19
Anq of fore wings 14-16 14-17 14.97 16.00 15-16 17
Anq of hind wings 10-13 11-14 11.44 12.30 10-11 13-14
Pnq of fore wings 7-10 7-9 8.11 8.14 8-9 8-9
Pnq of hind wings 8-10 7-11 9.03 8.60 9-10 9-11
Scq in all wings 1 in all counted except in left fore wing and hind wing
of allotype 9 and left hind wing of 1 S in which it
is absent.
Scq in all wings 1 present in 87 wings; 2 present in 17 wings.
Distal thickened Anq the 5th in 99 wings, 6th in 5 wings (4.5%).

Fore Hind Fore Hind Fore Hind
wing wing wing wing wing wing
1 celled
(open) 0 0 0 0
2 celled 5 21* 0 7* 1 95* 29*
3 celled 31* 15 16* 9 6S 39
1 celled 0 11* 0 6 3 29
2 celled 36* 25* 16* 10* 185* 89* 10 49*
3 called 0 0 0 0
Supratriangle: _
1 celled 36* 1 36* j 16* 16* T all males and females
*includes holotype cd and allotype 9


Stigma covers cells: Male fore wing 5-7 (aver. 8.11); hind wing
5-8 (aver. 6.20). Female fore wing 6-8 (aver. 7.70); hind wing 6-7
(aver. 7.20).
Anal field of fore wing proximal to triangle 2 cells wide for a dis-
tance of from 3 to 6 cells (approximate average 5); these may be so
irregularly arranged as to make counting difficult.
MEASUREMENTS (in millimeters).-Abdomen (including appendages)
male 38-40 (aver. 38.82); female 37-40 (aver. 38). Length of hind wing
male 30-32 (aver. 31.33); female 31-35 (aver. 32.25). Length of
stigma male 5 (aver. 5); female 5-6 (aver. 5.6).
ECOLOGICAL NOTES.-P. alachuensis is a new species of Progom-
phus restricted, as far as our records indicate, to the central group of
Florida counties (Clay and Alachua on the north to Orange, Volusia
and Lake on the south). South of Alachua County it seems to be
the only Progomphus in the dragonfly fauna; while in north and
west Florida P. obscurus alone is present. Thus, the new species over-
laps with P. obscurus in Alachua and Clay Counties at least.
All available records for P. alachuensis, adult, nymph or reared, in-
dicate a lake-pond habitat, chiefly lake. Adults are comparatively rare
and are to be found flying along the lake shore and low among the
plants of the shore zone or pond margin. Collection of adults has been
made from April to August; the peak of emergence seemingly coming
in May. In most respects the adults behave like obscurus.
J. H. Williamson notes on the envelope of a male specimen (No. 21
U. of Mich. coll.) taken April 22, 1921, at Enterprise (now Benson
Springs), "New Gomphine. Only one seen. Was flying nervously
about a small open space among small pine trees near Figure 8
pond and just as I approached to size him up, lit on a dried bit
of vegetation (like clover tip)."
The nymphs are found burrowing in the white sand bottom of lake
shores in fairly shallow water (up to about 3 feet) where they act
in a manner identical with those of P. obscurus (Byers 1930: 247. The
lake inhabiting nymphs referred to are those of P. alachuensis.)
Nymphs are common and have been collected in Newnans and Wau-
berg Lakes, Alachua County, along with the rarer "long-tailed" nymph
-Gomphoides williamsoni Gloyd.
Needham (1897) received a collection of Odonata from Orange Co.,
Florida including a Progomphus. Though the present author has not
seen this material, it is his belief, based on collector's data from
Orange County, that the species is alachuensis rather than obscurus.
Davis and Fluno (1938) state, "Gomphoides (Progomphus) obscura
(Rambur). Found always on sandy shores of local lakes [Winter
Park, Orange Co.], fairly common Apr. to Aug."


P. alachuensis is ecologically distinct from P. obscurus as far as the
writer's experience may indicate. In the region studied, chiefly Alachua
County, P. alachuensis is a lake-pond species; P. obscurus is an inhabit-
ant of sand bottom streams. The situation is of interest, not only within
itself, but also, when compared with that exhibited by the dragonflies
of the genus Neurocordulia (Byers 1937). In Neurocordulia the north-
ern species inhabit the lakes and streams where the nymphs can find
the required rocky conditions; the southern species occur only in
similar habitats in streams. Temperature may be the factor involved-
the southern lakes being too warm. The northern Progomphus also may
be found in lakes or streams (generally lakes-as in many regions the
streams have rock or gravel bottoms); in the south, the northern P.
obscurus is an inhabitant of the cooler streams, while the new species
(P. alachuensis) has come in to occupy the sand bottom habitat of the
warmer lakes. In the warmer southern part of Florida, P. obscurus
seems to be absent.
P. alachuensis was named after the type locality-Alachua County,
The nymphs of the genus Progomphus are best known for their abil-
ity to burrow rapidly in the sands of river and lake bottoms. This
habit is correlated with marked structural adaptations. In general,
these nymphs may be separated from other North American Gomphinae
by the following set of characters:
Third segment of the antennae cylindrical, not greatly flattened;
the 4th segment from one-third to one-half as long as the third, apex
tapering and upturned; apex of lateral lobe rounded; middle coxae
more approximate than the fore or hind ones; abdomen long and
slender, 9th segment with a dorsal spine.
A more detailed description of the nymph based on reared Florida
material (see material studied p. 67) of P. obscurus in the author's
collection follows:9
Head heart-shaped, depressed, sloping anteriorly, broadly notched
behind; hind angles rounded. Antennae inserted into cylindroid ele-
vations on the front, depressed and incurved; two basal joints very
short; terminal segment (4th) long and decidedly upturned; third

9The italics indicate special P. obscurus characters; non-italicised portions are
applicable to the genus as far as the nymphs of the various species are known.
lOFor figures of the nymph see references under the various species given on
pages 60 and 63.


segment 3-5 times longer than the terminal one, slightly flattened and
hairy. Labium rather small, reaching when folded to the base of the
middle legs; median lobe short and square, its cephalic border convex
and fringed with one series of short hairs overlying small denticles;
lateral lobes short, nearly straight, apex rounded, margins entire, mov-
able hooks stout and slightly incurved.
Thorax sloping to the head and the bases of the legs. Wing cases
strongly divergent. Legs conspicuously fossorial; fore legs approxi-
mate to the sides of the head, bearing shields of stiff hairs behind
which the middle legs may be brought forward; middle legs with coxae
closer together on the ventral surface than those of the fore and
hind legs, rotated downward and extended horizontally close under
the fore legs; hind legs longer and more nearly normal; tarsi and
tarsal claws modified for digging; each femur with a distal anterior
process which rests against and supports the tibia when moved back-
Abdomen long and narrow, segments about equal, 9th a little longer
than the others; dorsal spines on segments 1-9 varying in length on the
different segments; lateral spines on 5-9; appendages slender, tapering;
superior and inferiors about equal length, laterals about half as long.
Length 29mm.; length of abdomen 20mm.; width of abdomen 6mm.;
length of hind femora 3mm. General color light buff with brownish
spots on either side of the abdomen.
The first contribution made to our knowledge of the immature forms
of Progomphus was that of Cabot (1872: 6-16); he correctly assigned
certain nymphs from Wareham, Massachusetts, to this genus and pub-
lished the first figures.
Hagen (1885:48) published a detailed description of the nymph
of P. obscurus and in addition stated, "As P. obscurus has now been
collected near Boston, by Mr. Morrison, there can scarcely be a doubt
that the nymphae belong to this species." The material consisted of:
"Three nymphae in alcohol, male and female, probably full grown, from
Wareham, Mass., May, 1863 [Cabot's material], by Prof. L. Agassiz;
one in alcohol from Little Wabash River, Effingham, Ill., June, 1878,
Prof. S. A. Forbes." In this same paper (1885) Hagen lists under
the name, 32. Progomphus species" six nymphs from Santiago, Cali-
fornia, which in view of our present knowledge, were undoubtedly those
of P. borealis. This is the first published record of the nymphs of this
western species.
In 1890, Wm. Beutenmuller published a catalogue of the immature
forms of Odonata, he included both the Cabot and Hagen references
given above.


Needham (1897) received from Orange County, Florida, some
Odonata material collected by Mr. Adolph Hempel, April 21, 1897.
Needham writes, "Mr. Hempel took a Progomphus obscurus Ramb.,
with its skin in transformation. While the nymph was known by fair
supposition it appears not to have been reared before." It is the
opinion of the present writer that this specimen was one belonging to
the species P. alachuensis.
Needham and Hart (1901) described and figured the nymph of
P. obscurus from Illinois and noted, "The nymph was described and
by supposition connected with this species by Hagen (1885) and the
supposition was verified by Mr. Needham (1897)."
Kennedy (1917 b) first described and figured the nymphs of P. bo-
realis in sufficient detail to assign them to the correct species. Ken-
nedy's work is still the most important in our knowledge of this species
though he did not work from reared material.
Seeman (1927) indicated that P. borealis (she used the name P. ob-
scurus), was reared in the laboratory at Pomona College."
Other contributions to our knowledge of Progomphus nymphs have
been made by authors from time to time (see references under the
various species treated on pages 40 and 46).

The determination of species of Progomphus in the immature stages
is quite difficult; only borealis and obscurus are separable with any
degree of ease. The following key attempts to separate the four
known species-three of these have been reared, the fourth, P. integer
is included on supposition. Hagen (1885), under the heading "31.
Progomphus species", lists six nymphs from various regions of Bra-
zil, the specific identity of which was then, and still is, impossible to
establish, (See Cabot 1872 for a note on these specimens).

1-Lateral spines on abdominal segments 3-9. Dorsal spines low and rounded
on segments 4-9; high and well developed on segments 1-3. Fourth an-
tennal segment nearly half as long as third. Length of hind femora
(fully grown nymphs) 4mm. Body short and broad (length of abdo-
men 18-20mm., width 7mm.). Large black spots on dorsum of abdo-
men, especially on segment 7. Inhabit the shifting white sand beds of
the rivers of western United States (western Texas and Colorado to
northern M exico and the Pacific)..................................................... P. borealis.
-Lateral spines on abdominal segments 5-9. Dorsal spines on abdominal seg-
ments 1-9 moderately or well developed. Fourth antennal segment from


one-half to one-third as long as the third antennal segment11. Length of
hind femora (fully grown) 3mm. Body long and slender (length of
abdomen 20-23 mm., width 5-6mm.). Markings on dorsum of abdomen
brown, generally restricted, with isolated pale areas. Inhabit sand bottom
regions of streams, lakes and rivers in eastern United States and Cuba-.....(2)
2-Dorsal spines on abdominal segments 4-9 long and sharp, that on segment
9 about one-half as long as the dorsum of segment 10. Abdominal ap-
pendages relatively short (measured on ventral surface, about two-thirds
the length of segment 10). Nymphs small and slender. Cuba.....P. integer.
-Dorsal spines on abdominal segments 4-9 short and pointed, that on segment
9 from one-fourth to one-third as long as the dorsum of segment 10.
Abdominal appendages relatively long and pointed (measured on ventral
surface, slightly longer than the length of segment 10). Nymphs long
and slender. Eastern United States (Atlantic coast to eastern Texas and
C olorado. ....................................................................... ............................ (3)

-Nymphs inhabiting streams in central Florida, lakes or streams elsewhere in
eastern United States ................. ....................... ............. ..... P. obscurus.


After a careful and prolonged study of the various possible specific
characters for the separation of the nymphs of P. obscurus and P. ala-
chuensis, the conclusion has been reached that none exist. The nymphs
of these two species are practically identical as far as it is possible to
indicate differences in words. After long experience in handling ma-
terial, the worker feels that slight undescribable differences do exist,
but that they defy measurement or description.
Probably the dorsal spines of the four keyed species offer the best
set of characters for judging degrees of similarity and difference-
morphologically speaking. These spines are present on abdominal seg-
ments 1-9 and assume different degree of sharpness, length and height
not only on the various abdominal segments of the same species, but
in the different species as well. Some of these specific differences
have been utilized in the above key; they are not sufficiently distinct,
however, between obscurus and alachuensis to be so employed. To il-
lustrate this condition, the accompanying graph has been drawn. The
graph has been based on units of length of the dorsal spine (a unit is

111n nymphs of obscurus and alachuensis, at least, there is a sexual dimorph-
ism exhibited in the comparative lengths of antennal segments 3 and 4. In
males the ratio is 2.5: 1.0; in females it is 2.5: 1.5.


equal to a little less than a millimeter) shown in the vertical column;
the horizontal base-line gives the number of the abdominal segment
bearing the spine.

/o 9 8 6 5 f 3 a I

NYMPH OF Progomphus obscures
DESCRIPTION AND FIGURES.-Satisfactory descriptions, or diagnostic
characters, and figures have been published by the following authors:
Byers (1930: 50). Description.
Cabot (1872: 6-16). Description. Figure (P1. II, fig. 3).
P. Garman (1927: 126). Description. Figure (labium, Pl. V,
fig. 7).
Hagen (1885: 274). Description.
Howe (1917: 127). Pictoral keys. Figure (labium, Pt. II).
Needham and Hart (1901: 55-56). Description. Figure (nymph,
Pl. I, fig. 3).
Needham and Heywood (1929: 62). Diagnostic characters. Fig-
ure (nymph).



Needham and Needham (1927). Diagnostic characters. Figure
(nymph, P1. IV, fig. 8).
ECOLOGICAL NOTES.-Probably more has been written on the habits
of this species than on any other phase of the study of Progomphus,
no doubt because of the unusual burrowing habits of the nymph.
Brimley (1903) writes, "A number of nymphs were observed trans-
forming, from about 10 A. M. to 1 P. M. on June 5, 1900, resting on
bare patches of sand or on the bare clay banks, close to the water's edge,
but not climbing up plants."
Byers (1925) records the nymphs as, "Emerging in numbers in early
July" along the sandy shores of Douglas Lake, Michigan. The same
author (Byers 1930) records the nymphs as of "common occurrence"
in Florida and continues to say:
The nymphs are of much interest because of their ability as true sand bur-
rowers. The fore and middle legs are especially fitted for this process. So effi-
ciently do these legs operate that their possessor can become completely buried
within a few seconds after being placed on a bed of clean sand Small white-
sand bottom creeks, flowing through woods are their favorite abode As ob-
served they always occur under moving water conditions. In the laboratory,
nymphs thrive better under moving water conditions, even though the sand has
been previously cleaned. Two nymphs were reared in the laboratory. Each
molted once and on May 14 and 15 adults appeared. At this time in May cast
skins began to appear out-of-doors. To emerge, the nymph crawls up the shore
but a little way above the water line and continues the process directly from
that position. One cast skin was found on a small sand shoal temporarily raised
into a mid-stream island no longer than a dollar bill. The nymph had emerged
so near the water line that its feet were practically wet during the process, al-
though the skin was dry and in its customary position. The food [of the nymph]
consists mainly of small annelid worms found in large numbers buried in the
sand with them. These Odonata larvae may be considered as an ecological
index to a habitat of clean sand underlying moving water.
Howe (1917-Pt. II Larva) states, "Larva [of P. obscurus] in-
habit sandy ponds and river bottoms. Larval life occupies probably
three years."
Kennedy (1917a) in discussing the distribution of P. obscurus in
Kansas notes that the species, occurs wherever there are
sand-bedded streams, as its nymph is adapted to rapid burrowing in
flowing sand; consequently it is abundant throughout western Kansas."
In the same paper, Kennedy further notes that P. obscurus is
". found only on streams whose beds are loose, flowing sand, in
which the highly specialized nymph burrows".
Kennedy (1921) writing about some dragonfly nymphs sent to him
from eastern Texas (Colleto Creek, Victoria, July 20, 1920) states,


"The material consists of 35 naiads. The largest two would
undoubtedly have emerged during the season of 1920, as the Progom-
phus season is a long one. Whether the numerous smaller examples would
have emerged the same season or one or two seasons later is impossible
to say, as so little is known concerning the life history of Progomphus.
Burrowing insects in general are slow feeders and consequently slow
Needham and Hart (1901) write, "The nymph of this species .
represents an extreme specialization for life as a burrower in the bed of
running streams it is common in the sand of smaller rivers .
one specimen was found at the lower edge of a sand-bar in the Spoon
River [Illinois]). Perhaps the peculiar habits of the nymph may be
correlated with a relatively long period of nymphal life and a short
period of life as an imago." Needham, in several papers, records the
nymph of obscurus as an inhabitant of "rapid streams", "lotic waters",
etc. (see 1901, 1903b, 1927, 1929), as do other authors after him.
Williamson (1901) observes that "A half dozen exuviae were gath-
ered on the sand beaches near the water's edge [Chapman Lake, Indi-
ana, June 30, 1901]."
In summarizing the foregoing, we may say that the nymphs of Pro-
gomphus obscurus are:
1-Highly adapted for burrowing in the clean sand-bottoms of
streams, rivers and lakes, (P1. VI, fig. 2).
2-Associated under these conditions with more or less rapidly
moving water.
3-Found emerging near the water's edge directly from bare patches
of sand, river banks, lake shores, etc. Not found emerging from vege-
tation, the sides of wharves, far from water, etc.
4-Generally common in their proper habitat.
5-Slow feeders, slow growers, with long larval life.
Dr. Kennedy (1921) made a phylogenetic study of Progomphus
nymphs based on material (already noted) that he received from east-
ern Texas. His P. obscurus specimens-35 in all-ranged in length
from 7mm. to 28mm., the two largest being 25 and 28mm.; some 20
of these specimens were broken, thus making a table of measurements
impossible. Regarding this collection he writes:
The small specimens are the smallest Progomphus naiads the writer has seen,
and are interesting in that they show, by the way the mature naiads differ from
them, the direction in which the latter are specialized .The specimens 7mm.
long differ from the mature naiads [25-28mm. long] as follows:
1. The wing pads are minute triangular flaps.
2. The dorsal spines on segments 2 and 3 are lacking.


3. The superior pair of anal appendages are mere tubercles.
4. The fourth segment of the antennae is only one-third as long as the third.
(In mature naiads it is nearly one-half as long as segment 3).
5. Labium with two triangular teeth on the middle lobe besides the usual row
of truncate teeth. (In the adult [sic] there are not noticeable triangular teeth).
6. Lateral lobe with several broad though shallow teeth.
It can be legitimately inferred that these ontogenetic changes point out the
phylogeny of this most specialized of burrowing Odonate naiads. They must
have developed through a Gomphus-like form that had the characteristics of the
very young naiads. It is interesting to note that these specializations have devel-
oped more in the naiads of P. borealis, and that on the naiadal characters alone
obscurus is more generalized than borealis. This pair of species agrees with
Amphiagrion and a number of other pairs of closely related species of Odonata in
which the western form is more specialized than the eastern. It is interesting to
those studying distribution to find that these obscurus naiads from Texas, where
both obscurus and borealis are found, are in no wise intermediate in character be-
tween the two species but are true obscurus naiads.

DESCRIPTION AND FIGUREs-Satisfactory descriptions, or diagnostic
characters, and figures have been published by the following authors:
Hagen (1885: 275). Diagnostic characters: species not named.
Kennedy (1917b: 527-529). Good description. Figures (nymph,
first leg, mentum detail, head, labium, segments 9-10 of male and
female, figs. 155-163, page 528).
ECOLOGICAL NOTES.-The most important, and virtually the only,
contribution made to our knowledge of the nymphs of P. borealis
was that of Kennedy (1917b). Regarding their habits in southern
California he writes:
These nymphs seem to occur in the drifting beds of all the sand-bottomed
streams of central and southern California where there is a permanent flow of
water . The nymph of this species is more highly specialized for burrowing
than any other odonata nymph with which I am familiar. It is the most frequently
occurring form of large insect life found in the sand beds of the rivers of central
and southern California. All of the rivers of California originate in the moun-
tains as swift, clear streams, but nearly all before reaching the sea have almost
level lower courses where they lose their swiftness, become warm and meander
over beds of shifting white sands. These sand beds are so unstable that they
actually flow with the water though at a much slower rate . in such a
stream the bottom inch of water is turbid with moving sand. It is in such an
unstable environment that Progomphus flourishes. Only once or twice have I
actually found nymphs in the flowing streams, but in any side channel where
the water has ceased to flow the sand is marked in all directions by the curving
burrows of these wandering nymphs. They burrow in the loose upper stratum
of the sand and just deep enough to cover the back. Usually the tenth segment
protrudes. I have traced burrow tracks that were from ten to fifty feet long. If one


arrived at the fresh end of a burrow before some water fowl had found it the
nymph would be found spasmodically burrowing forward. These nymphs could
burrow as rapidly as the average nymph can .walk. Only such vigor and speed
could save an insect of this size from being buried in the moving sand.
The following authors have added to or substantiated Dr. Ken-
nedy's work on P. borealis:
Seeman (1927) writes, "The nymph lives in the shifting sand
bars of streams. In the Santa Ana River, in the sandy shallows near
the shore, a pint jar might have been filled in a short time so abund-
ant were they. They burrow out of sight so rapidly that when once un-
covered, one must work quickly or Progomphus has buried himself
in the sand and is out of sight like a flash."
Needham and Heywood (1929) note, "The senior author found
nymphs of this species exceedingly abundant in the sandy bed of the
Santa Ana River below Riverside, California, in April. A dozen or
more could be taken at each sweep of the sieve-net on the bottom."
Tinkham (1934) writes, "The Cibolo Creek at Shafter [southwest
Texas] supported Progomphus as the commonest species. At Wilson's
Ranch, 25 miles southeast of Marfa [Texas], a number of Gomphoid
naiads were found burrowing in the sand of the Alamito Creek bed
leaving long meandering trails behind them. These were probably
Progomphus . but there were no adults in the air at this time
in late May."
From the foregoing accounts, it will be seen that the habits of
P. borealis are very similar to those of P. obscurus.. However, several
points of difference may be noted:
1) No mention is made of finding P. borealis in lakes. They seem to
be restricted to the quieter portions of rivers and streams.
2) The production of meandering burrows visible and traceable
on the sand bottom surface. No author has recorded, nor have I seen,
burrows of this nature for P. obscurus.
3) The more specialized burrowing adaptation in P. borealis. It
would be interesting to time the burrowing speed of both P. borealis
and P. obscurus respectively to see which was the more rapid.
The nymphs of P. alachuensis are structurally (see key page 58)
and ecologically similar to those of P. obscurus with the notable ex-
ception that, in the Florida area where the ranges of the two species
overlap, alachuensis is a lake-dweller and obscurus is found restricted
to streams.
The morphological similarity of the nymphs of these two species
(alachuensis and obscurus) together with their geographic distribution


and ecological peculiarities opens some interesting lines of speculation.
The following facts and hypothesis may be utilized in an attempted
1). P. obscurus and P. borealis are both widely distributed species
and the morphological differences between them in both nymphal and
adult stages are marked.
2). Of these two species obscuruss and borealis), P. obscurus ap-
pears to be the most primitive (Kennedy 1921). Therefore, we may
assume that borealis was derived from obscurus at some time suffici-
ently long ago to permit its wide spread distribution, distinct charac-
ters, and absence of intermediates in the region (Texas) where it con-
tacts the range of obscurus.
3). Associated with obscurus, on the basis of nymphal similarities
(see key page 58), are alachuensis and integer. In the adult stage
these are distinct enough and are undoubtedly good species. However,
in the nymphal stages they converge in such a manner that the
nymphs of integer are quite similar to those of obscurus, but with a
marked set of minor characters (see graph of dorsal spines page 60);
while the nymphs of alachuensis are practically identical with those of
4). Both P. integer of Cuba and P. alachuensis of central Florida are
limited in distribution. The geological history of the Antillean region
indicates that both Cuba and an area of central Florida are or were
at one time islands. This Central Florida area just about delimits
the present range of alachuensis.
5). Applying the concept of isolation in the formation of species and
the role that islands play in this process, we may perhaps assume
that originally integer and alachuensis were isolated reminants of an
eastern North American Progomphus stock, and that subsequent
speciation resulted from mutation and isolation. Thus, three species
resulted, P. obscurus on the mainland, P. alachuensis on the Central
Florida Island, and P. integer on Cuba.
6). The reunion of the Central Florida Island (Pleistocene) with
the mainland of North America brought the ranges of P. obscurus and
P. alachuensis in contact with each other. The separation, however,
remained long enough for adult specific characters to be produced
but not long enough for nymphal ones to arise.
7). On the reunion of the two land masses, P. obscurus invaded
north and north central Florida. In north central Florida the invad-
ing species overlapped the range of the endemic one but remained
ecologically distinct, inhabiting the rivers and streams, leaving the
lakes for the species already established.


8). The isolation of P. integer has continued with consequent accumu-
lation of specific differences extending to the nymphal stage.
There are other explanations for the P. obscurus-P. alachuensis
relationship-perhaps simpler ones. For example, P. alachuensis may
be a far more modern species than is assumed in the above hypo-
thesis, and have been separated from P. obscurus through ecological
isolation exclusively; isolation, in the sense that the Florida endemic
species is adapted to somewhat warmer lakes and P. obscurus can
establish itself only in the cooler streams of north and north cen-
tral Florida.
The following listed material was studied or examined by the author
in the preparation of this paper:

Progomphus obscurus:
Florida: Alachua Co., Gainesville, April 2, 1923, 1 T. H. Hubbell (Byers
coll.), May 14, 1928, 1 May 30, 1929, 1$, May, 1933, 1, July 7, 1935
(Hog Town Creek) 2 July 12, 1935, 19; Hawthorne (Magnesia Springs),
May 16, 1932, 2 6, May 25, 1935, 1 3, C. F. Byers (Byers coll.). Jackson Co.,
Chipola River, April 13, 1935, 19, Bellamy (Byers coll.). Liberty Co.,
Camp Torreya, May 30, 1924, 19, May 31, 1924, 1 19, June 1, 1924,
19, C. F. Byers (Byers Coll.). Georgia: Johnson Co., Keysville, June
28, 1930, 4 C. F. Byers (Univ. of Mich. coll.). Wayne Co., Jesup, June
22, 1930, 16, June 23, 1930, 4 C. F. Byers (Univ. of Mich. coll.).
Kentucky: Powell Co., Walthersville, Red River, Aug. 1, 1929, 2 Aug. 2,
1929, 3 Jean Lowry, Thornton Scott, E. B. Williamson, (Univ. of Mich.
coll.). Indiana: Allen Co., Devils Hollow, July 9, 1922, 19, E. B. Williamson
(Univ. of Mich. coll.). Wells Co., Bluffton, June 22, 1921, 1 19, E. B. Wil-
liamson (Univ. of Mich. coll.). Michigan: Cheboygan Co., Douglas Lake, July
8, 1924, 1 19, C. F. Byers (Byers coll.). Arkansas: Lincoln Co., July 5, 1931,
4 July 9, 1931, 6 L. H. Bridwell (Univ. of Mich. coll.). Missouri: Carter
Co., June 21, 1930, 3 o, E. B. Williamson, et al. (Univ. of Mich. coll.). Stoddard
Co., Dexter, July 17, 1930, 24 a 49, E. B. Williamson, et. al. (Univ. of Mich.

Progomphus borealis:
Arizona: Granite Reef Dam, Sept. 22, 1912, 1 Sept. 15, 1912, 19, B. J.
Rainey (Univ. of Mich. coll.). Oslar, Salt River, April 12, 1902, 1 19, (Univ.
of Mich. coll.). Pima Co., Catalina Mts., Aug. 6, 1930, 2 L. K. Gloyd (Univ.
of Mich. coll.). California: Chico, June 17, 1914, 1 C. H. Kennedy, (Univ.
of Mich. coll.). Oroville, June 13, 1914, 1 &, C H. Kennedy (Univ. of Mich.
coll.). Utah: St. George, June 6, 1919, 1 Tom Spalding (Univ. of Mich.
coll.). New Mexico: Albuquerque, April 18, 1903, 19, Oslar? (Univ. of Mich.
coll.). Texas: Davis Mountains, Fort Davis Quad., Phantom Lake, June 1,


1916, 2 19, June 9, 1916, 5 $ 29, July 13, 1916, 1 F. M. Gaige (University
of Mich. coll.). Mexico-Baja California: Purissima, Oct. 13, 1923, 19, J. H.
Williamson (Univ. of Mich. coll.). Mexico-Sonora: Hermosillo, Sept. 25, 1923,
1 1 9, J. H. Williamson (Univ. of Mich. coll.).

Progomphus alachuensis: (In addition to types and paratypes).
Florida: Alachua Co., Newnan's Lake, Gainesville, May 15, 1931, 1S, C.
F. Byers (Byers coll.). Clay Co., Kingsley Lake, May 1935, 2 8 2 9, reared
by C. F. Byers (Byers coll.). Volusia Co., Benson Springs (Enterprise) April
21, 1921, 19, J. H. Williamson (Univ. of Mich. coll.), April 26, 1921, 1S, J.
H. Williamson (Univ. of Mich. coll.).

Progomphus clendoni:12
Guatemala: Agua Caliente, June 1, 1909, 1 S, E. B. Williamson, Amatitlan, June
8, 1909, 1 E. B. Williamson. El Fiscal, June 6, 1909, 1 E. B. Williamson
Los Amates, June 19 1909, 1 E. B. Williamson. Puerto Barrios, May 28-30,
1909, 3 a, E. B. Williamson. Mexico: Hacienda de San Marcos, near Villegas,
State of Jalisco, Nov. 22, 1923, 1 J. H. Williamson. Acaponeta, State of
Nayarit, Nov. 2, 1923, 1 S, J. H Williamson.

Progomphus intricatus:
Brazil: Chapada, December, 1 S, H. H. Smith.

Progomphus joergenseni:
Argentina: Chacras de Coria, March 5, 1907, 2 ; March 9, 1907, 19.

Progomphus phyllochromus:
Colombia: Vista Nieve, Mt. San Lorenzo, Santa Marta Mts., June 15, 1920,
48 19, F. M. Gaige.
Progomphus pygmaeus:
Colombia: Bolivar, Dec. 24-25, 1916, 10 $ 3 9, E. B. and J. H. Williamson.

Progomphus risi:
Guatemala: Gualan, June 16, 1909, 2 S 19 (Types and paratype), E. B.

Progomphus dorsopallidus:
Venezuela: San Esteban, Feb. 5-6, 1920, 4 e (type and paratypes), J. H. and
E. B. Williamson, W. H. Ditzler.
Progomphus obscurus:
Florida: Alachua Co., Gainesville, Rattlesnake Branch, April 10, 1925, 1, T.
H. Hubbell; "C-creek", May 2, 1928, 15, February 21, 1930, 7, March 1,
12All of the following specimens for South American and Central American
localities are in the University of Michigan collection.


1930, 4; Santa Fe River, November 10, 1931, 10; city environs, May 14, 1928, 2
(reared $s), C F. Byers (Byers coll.). Michigan: Washtenaw Co., Whitmore
Lake, October 1923, 8; Ann Arbor, 1925, 15 exuviae, C. F. Byers (Byers Coll.).
Cheboygan Co., Douglas Lake, July 8, 1924, 1 (reared &) C. F. Byers (Byers
coll.). Oklahoma: Camp Boulder, Wichita Mts., June 12, 1926, 1, T. H. Hubbell
(Byers coll.).

Progomphus borealis:
Arizona: Tucson, 2, J. G. Needham (Cornell coll.).

Progomphus alachuensis:
Florida: Alachua Co., Gainesville, Wauberg Lake, April 29, 1933, 1 (reared ),
R. Trogdon (Byers coll.). Clay Co., Gold Head Branch State Park, February
1939, 1, (reared?), J. G. Needham (Cornell coll.). Kingsley Lake, May 1935, 13
(85 9 reared), C. F. Byers (Byers coll.).

Progomphus integer:
Cuba: Solidad, 5, J. G. Needham (Cornell coll.).

1890. "Preliminary Catalogue of the Described Transformations of the Odonata
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1932. "Dragonflies of Oklahoma," Univ. of Okla. Publications, Vol. 4, pp. 50-57.

1935. "New Records of Ohio Dragonflies," Ohio Journ. Sci., Vol. 35, pp. 451-456.
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1934. "Notes on Some Generic Names of Odonata," Ent. Monthly Mag.,
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1903. "The Odonata Collected by Messrs. Schwarz and Barber in Arizona and
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DAVIS, E. M., and J. A. FLUNO.
1938. "Odonata at Winter Park, Florida," Ent. News, Vol. 49, pp. 44-47.

1913. "Dragonflies of the Vicinity of New York City with a Description of
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1850. "Revue Des Odonates ou Libellules d'Europe," Mem. Soc. Sci. Liege,
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1915. "Dragonflies of the Douglas Lake Region, Michigan," Mich. Geol. and
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1929. "A Revision of the Fissilabioidae," 2 Parts. Mem. Indian Mus., Vol.
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1924. "Odonata from Kentucky," Ent. News, Vol. 35, pp. 285-288.

1927. "Guide to the Insects of Connecticut. Part V. The Odonata or Drag-
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1861. "Synopsis of the Neuroptera of North America," Washington: Smith-
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1874. "The Odonata Fauna of Georgia," Proc. Boston Soc. Nat. Hist., Vol. 16,
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1885. "Monograph of the Earlier Stages of the Odonata," Trans. Amer. Ent.
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1900. "Additions and Corrections to the 'Odonata of Ohio'," Ohio State Univ.
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1924. "The Odonata of Henry County (Iowa)," Iowa Acad. Sci., Vol. 31,
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IOWE, R. H., JR.
1917. "Manual of the Odonata of New England," Mem. Thoreau Mus. Nat.
Hist., II, pp. 1-138; Mem. David Mason Little Memorial Mus. Nat.
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1919. "A List of the Dragonflies Collected at Wareham, Mass., During the
Years 1911 to 1913 by Mr. Outram Bangs," Psyche, Vol. 26, pp. 65-68.
1920. "Odonata of Chatham, Massachusetts," Psyche, Vol. 27, pp. 57-60.
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1890. "Ober Gomphiden." Entomologische Nachrichten, Vol. 16, pp. 370-382.
1902. "A List of the Dragonflies of Winona Lake," Proc. Indiana Acad.
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1917a. "Studies in Kansas Insects. The Dragonflies of Kansas," Bull. Univ.
of Kansas Biol. Ser., Vol. 18, pp. 127-145.
1917b "Notes on the Life History and Ecology of the Dragonflies (Odonata)
of Central California and Nevada," Proc. U. S. National Mus., Vol.
52, pp. 483-635.
1921. "Some Interesting Dragonfly Naiads from Texas," Proc. U. S. National
Mus., Vol. 59, pp. 595-598.
1928. "Evolutionary Level in Relation to Geographic, Seasonal and Diurnal
Distribution of Insects," Ecology, Vol. 9, pp. 367-379.
1929. "The Theory of Nomenclature," International Congress Ent. IV., Vol.
2, pp. 665-670.
1890. "A Synonymic Catalogue of Neuroptera Odonata or Dragokflies,"
London, pp. III-IX; 1-202.
1934. "Head Characters of the Odonata," Entomologica Americana, Vol. 14,
pp. 41-97.
1925. "Records of Indiana Dragonflies," Proc. Indiana Acad. Sci., Pt. I-
Vol. 34, pp. 383-388. Pt. II-Vol. 36, pp. 387-291. Pt. III-Vol. 38,
pp. 335-343. Pt. IV-Vol. 39, pp. 309-314. Pt. V-Vol. 40, pp. 347-
349. Pt. VI-Vol. 41, pp. 499-454. Pt. VII-Vol. 43. pp. 211-217, Pt.
VIII-Vol. 44, pp. 231-235. Pt. IX-Vol. 46, pp. 203-210. 1925-1937.
1933. "Notes on New Jersey Dragonflies," Ent. News, Vol. 44, pp. 40-44.
1932. "Beitrage zur Kenntnis der Biologie der Odonatenfamilie der Gomphidae
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.MTTrKowsKI, R. A.
1908. "Review of the Dragonflies of Wisconsin," Bull. Wise. Nat. Hist. Soc.,
Vol. 6, pp. 57-127.


1910a. "Catalogue of the Odonata of North America," Bull. Public Mus. Mil-
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1897 "Preliminary Studies of North American Gomphinae," Can. Ent.,
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1903. "The Skewness of the Thorax in Odonata," Journ. N. Y. Ent. Soc., VoL
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and Gomphidae," Bull. Ill. State Lab. Nat. Hist., Vol. 6, pp. 194.

1929. "A Handbook of the Dragonflies of North America," Baltimore: Charles
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1927. "A Guide to the Study of Fresh-water Biology," New York: American
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1911. "Odonata Collected at Douglas Lake, Michigan, in the Summer of 1910,"
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1842. "Histoire Naturelle des Insects. NeuroptBres," Suites a Buffon, Paris.

Ris, F.
1908. "Beitrag zur Odonatenfauna von Argentina," Deutsch. Ent. Zeitschr.,
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1927. "Dragonflies, Mayflies, and Stoneflies of Southern California," Journ.
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1938. "Die Tibialleiste der Odonaten," Zoologischer Anseiger, 1. 3. Band 121,
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1934. "The Dragonfly Fauna of Presidio and Jeff Davis Counties of the
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1899. "The Dragonflies of Indiana," 24th Ann. Rept. Dept. Geol. Nat. Res.
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FIos. 1-3.-Top of head showing diagnostic characters of frons and occiput.
FIG. 1.-Progomphus obscurus, $, Dexter, Missouri.
FIo. 2.-Progomphus borealis, 6, Baja California.
Fio. 3.-Progomphus alachuensis, 9, Alachua County, Florida.

FIGS. 4-6.-Male abdominal appendages, lateral view.
FIG. 4.-Progomphus obscurus, Dexter, Missouri.
FIG. 5.-Progomphus borealis, Pima County, Arizona.
FIG. 6.-Progomphus alachuensis, Alachua County, Florida.

FIG. 7. -Male genitalia of abdominal segment No. 2, ventral view. Progomphus
alachuensis, Alachua County, Florida.




FIGs. 1-3.-Ventral view of abdominal segment No. 9 of female showing vulvar
FIG. l.-Progomphus obscurus, Dexter, Missouri.
FIG. 2.-Progomphus borealis, Baja, California.
FIG. 3.-Progomphus alachuensis, Alachua County, Florida.

FIGs. 4-6.-Male abdominal appendages, ventral view.
FIG. 4.-Progomphus obscurus, Dexter, Missouri.
FIG. S.-Progomphus borealis, Pima County, Arizona.
FIG. 6.-Progomphus alachuensis, Alachua County, Florida.




FiG. 1. -Diagram of Progomphus thorax (synthorax).

FIGs. 2-5.-Thoracic patterns of North American Progomphus:
FIG. 2.-Progomphus alachuensis, Alachua County, Florida.
FIG. 3.-Progomphus obscurus, Powell County, Kentucky. Light.
FIG. 4.-Progomphus obscurus, Dexter, Missouri. Average or normal
FIo. 5.-Progomphus borealis, Pima County, Arizona.


*1 1







FIcs. 1-8.-Thoracic pattern of South and Central American Progomphus. Re-
drawn mostly from literature as cited.

FIG. 1.-Progomphus pygmaeus. Ris (1916:140. Fig. 83).
FIG. 2.-Progomphus phyllochromus. Ris (1916:140. Fig. 85).
FIG. 3.-Progomphus longistigma. Ris (1916:140. Fig. 79).
FIG. 4.-Progomphus gracilis, 8. Ris (1911:111. Fig. 7).
FIG. 5.-Progomphus gracilis, 9. Ris (1911:111. Fig. 8).
FIG. 6.-Progomphus lepidus, &. Ris (1911:112. Fig. 9).
FIG. 7.-Progomphus sp.? Williamson (1920a. P1. I, Fig. 3).
FIG. 8.-Progomphus risi, &. Williamson (1920a. P1. I, Fig. 1).





FIcs. 1-7.-Thoracic pattern of South and Central American Progomphus. Re-
drawn from literature as cited.

FIG. 1.-Progomphus costalis. Ris (1911:116. Fig. 16).
FIc. 2.-Progomphus recurvatus. Ris. (1911:114. Fig. 13).
FIG. 3.-Progomphus basistictus. Ris (1911:118. Fig. 18).
FIc. 4.-Progomphus complicatus. Espirito Santo. Ris (1911:114,
Fig. 11).
FIG. 5.-Progomphus complicates, San Bernardino. Ris (1911:114.
Fig. 12).
FIG. 6.-Progomphus auriopictus. Ris (1911:109. Fig. 4).
FrG. 7.-Progomphus dorsopallidus. Byers (1934b. P1. II, Fig. 1).






FIGS. 1-2-Progomphus habitats in Florida.

FIG. 1.-Newnan's Lake, Alachua County, Florida. Habitat of Pro-
gomphus alachuensis.
FIG. 2.-Hog Town Creek near Gainesville, Florida. Habitat of Pro-
gomphus obscurus.


i. F;



Peninsular Oil and Refining Company, Tampa

French geologists have been making paleogeographic maps since
the latter part of the nineteenth century and Charles Schuchert of Yale
University has since 1910 made himself the American pontiff of this
specialized branch of the science of geology. Such maps have been of
great service in the study of intercontinental relationships in the class-
room and in the last decade the petroleum geologist has found such maps
of much help in his search for new fields. Since the discovery of the
East Texas Field the trend in exploration for oil has been in search
of buried shore lines, and as structure hunting today is done largely
with geophysical instruments, the burden on the geologist is the study
of buried conditions in sedimentary basins to give some idea as to
source and accumulation beds.
This paper is here presented for the threefold purpose of (1) call-
ing to the attention of the Academy the importance of the Florida area
in the interplay of forces between the North American continent and
the Antillean region, (2) showing closer control on paleogeographic con-
ditions previously postulated, and (3) presenting data on the subsurface
of the state which are as yet unpublished.
Unless otherwise noted the paleontologic information here used
comes from Esther Richards Applin of Fort Worth, Texas, who ex-
amined cuttings for the Peninsular Oil and Refining Company from
its own well and other wells drilled in the area discussed. Apprecia-
tion is acknowledged to those operators who furnished well samples
for study. The literature used is from the works of Schuchert1,

*Awarded the Achievement Medal of the Academy for 1939.
1Schuchert, Chas., Historical Geology of the Antillean-Caribbean Region (New
York: John Wiley & Sons, 1935)


Stephenson2, Cooke and Mossom3 and others. Schuchert's work on
the Antillean Region and Stephenson's on the Gulf Coastal Plain have
furnished most of the information from beyond the limits of Florida.
Acknowledgment is made of the kindness of Dr. Roy E. Dickerson of
Philadelphia and Dr. W. A. Ver Wiebe of the University of Wichita
for reading the rough draft of this paper and making some suggestions.
The maps presented are not radical departures from those previously
published, but represent a closer control on land and sea limits. Em-
phatically these maps indicate only the general limits of the seas and
should not be regarded as mapping the location of ancient shorelines.

The geologic record in the literature is comprised of various phe-
nomena. The presence of marine, continental, coastal, or estuarine beds,
studied with particular attention to lithologic and paleontologic faces
and their variation, indicate the distribution of the lands and seas.
The presence of unconformities indicates areas where the above
evidences may have once been present but now have been removed
by erosion. The nature of the unconformity may indicate the nature of
the forces removing the sedimentary record.
The record in the well reports is almost entirely confined to the
presence or absence of deposits of various ages. Naturally the pres-
ence of marine strata of a particular age indicates the former pres-
ence of the sea at that point. Its absence might indicate that the
sea had never been present there or that the deposits had been
later removed by erosion. Here of course the judgment of the paleo-
cartographer is involved.

2Stephenson, Lloyd W., "Major Marine Transgressions and Regressions and
Structural Features of the Gulf Coastal Plain," American Journal of Science, 5th
Series, Vol. XVI., No. 94 (1928), pp. 281-298.
--- "Correlation of the Upper Cretaceous or Gulf Series of the Gulf
Coastal Plain," American Journal of Science, 5th Series, Vol. XVI, No. 96 (1928),
pp. 485-496.
--, and Reeside, John B., Jr., "Comparison of Upper Cretaceous Deposits
of the Gulf Region and Western Interior Region," Bulletin American Association
of Petroleum Geologists, Vol. 22, No. 12 (1938), pp. 1629-1638.
--, and Monroe, Watson H., "Stratigraphy of Upper Cretaceous Series in
Mississippi and Alabama," Bulletin American Association of Petroleum Geologists,
Vol. 22, No. 12 (1938), pp. 1639-1657.
3Cooke, C. Wythe and Mossom, Stuart, "Geology of Florida," Florida Geological
Survey, Twentieth Annual Report, (1927-1928), pp. 29-227.


Thomas Mann in his recent account of Joseph and his circle opens
with a philosophic overture which reads, "Very deep is the well of
the past. Should we not call it bottomless?" Had his past included the
geologic past as did that of H. G. Wells in his famous Outline he
would have seen that his "well of the past" was over one hundred
miles deep, for the post-Proterozoic stratigraphic section has been esti-
mated as 350,000 feet thick4, and Dana5 and others have estimated
that this represents only about half of geologic time. This section
in the "well of the past" is divided into comparative eras, Arche-
ozoic, Paleozoic, Mesozoic, and Cenozoic, with many subdivisions,
but avoiding any assignment of absolute age in terms of years. True,
attempts have been made to arrive at an absolute age by estimating
rates of sedimentation, denudation, or the increasing salinity of the
sea6, but at best they are only springboards for guesses. More recently
however, the physicist7 has furnished a better technique by showing
that an absolute age may be arrived at by comparing the amount of
helium and lead yielded by uranium, with unaltered uranium. This
method of approach has given an approximate age for the oldest
known rocks as one billion eight hundred million (1,800,000,000)
years. The method has the merit of checking with the comparative
method but Fenner8 has pointed out that errors usually tolerated in
analyses made of radioactive material may lead to grievous errors
when applied to earth age determinations. A recent textbook gives
the following table for such eras:9
4Field, R. M., "Chronology" in Van Nostrand's Scientific Encyclopedia
(New York: D. Van Nostrand Company, 1938), p. 263.
5Schuchert, Chas., The Evolution of the Earth (New Haven: Yale University
Press, 1923), pp. 45-81.
6Lane, Alfred C., "The Earth's Age by Sodium Accumulation," American
Journal of Science, 5th Series, Vol. XVII. No. 100 (1929), pp. 342-346.
7Kovarik, Alois F., "Basis for Computing the Age of a Radioactive Mineral
from the Lead Content," American Journal of Science, 5th Series, Vol. XX, No.
116 (1930), pp. 81-100.
8Fenner, Clarence N., "The Analytical Determination of Uranium, Thorium
and Lead as a Basis for Age-Calculations," American Journal of Science, 5th
Series, Vol. XVI, No. 95 (1928), pp. 369-381.
Fenner, Clarence N., "Radioactive Minerals from Divina de Uba, Brazil,"
American Journal of Science, 5th series, Vol. XVI, No. 95 (1928), pp. 382-391.
SMiller, William J., An introduction to Historical Geology (4th ed., New
York: D. Van Nostrand, 1937), p. 31.


Quaternary: 0 to 1,000,000 years ago.
Tertiary: 1,000,000 to 50,000,000 years ago.
Mesozoic: 50,000,000 to 200,000,000 years ago.
Paleozoic: 200,000,000 to 500,000,000 years ago.
Proterozoic: 500,000,000 to 1,000,000,00 years ago.
Archeozoic: 1,000,000,000 to 1,800,000,000 years ago.
But other than permitting the teacher to awe his pupils with
the magnitude of the study, and helping the lecturer dull the senses
of his hearers so that they are impressed with the futility of question-
ing later statements, there is little use for this absolute geologic cal-
endar. Happy in realizing that he does not have to squeeze all geol-
ogic time into Archibishop Ussher's 4000 years, the geologist returns
to his comparative scale.
Peninsular Florida's subsurface is best shown in cross section
(Fig. 2). It is based on the findings in nine wells (Fig. 1) and
shows the subsurface distribution of the geological formations in a
north-south direction the length of the Florida peninsula. It should
be kept in mind that the diagram is very much foreshortened
making the dips appear at much steeper angles than is actually the
case. The distortion will be easily understood if it is kept in mind
that this north-south distance represents over 500 miles, while the
vertical section represents less than 3 miles.
Adams McCaskill Wells.-In the last year and a half there have
been two wells drilled in Pierce County, Georgia, that have revealed
much of interest in the study of the subsurface of the southeastern
United States. The first was drilled by the Pan-American Production
Company on the McCaskill farm. After drilling 600 feet of undif-
ferentiated beds, the Ocala (Eocene) limestone was encountered
and persisted to a depth of 2000 feet, where a brown and green
marl with Midway fossils occurred.
At 2700 feet was found brown glauconitic shale with Cretaceous
fossils continuing to 4000 feet, then succeeded by coarse gray sands
whose character suggests Eutaw. The last 30 feet of section, to a
total depth of 4370 feet, is described in the samples as pink and
grayish altered granite, but reports from the well indicate that the
drilling terminated in hard, unweathered granite.
Not satisfied with this the group that had assembled the acreage
drilled another well, the Donald Clark No. 1, about three miles


away, apparently in the hope of finding some oil on a postulated
shore around this "granite island." The results in general were the
same, a similar section being penetrated. The samples saved were
taken at 30 foot intervals, so contact points cannot be given with
any assurance, but all paleontologists whose reports are available are
agreed on the section drilled through-a Tertiary and Upper Cre-
taceous section normal to the outcrop areas of Georgia and Alabama,
underlaid by granite.
St. Mary's River Oil Corporation Well.-At present there is drill-
ing in Nassau County, Florida, a well that has unexpectedly stumbled
on a black shale, which, though it shows no fossils, has been tentatively
regarded as Pennsylvanian or older by a number of workers who have
examined the samples. This well, known as the "Brown well" after
the president of the operating company, is located in the northwest
corner of Nassau County, a short distance from the Georgia line, and
started drilling in a formation mapped as Pleistocene but soon (at
40 feet) went into beds containing Elphidium gunteri var., E. incertum,
Rotalia beccarii var. parkinsoniani, Bolivina pulchella var. primitive,
Noniori cf. depressulum. This fauna is suggestive of the Choctaw-
hatchee formation of the Upper Miocene. From 400 feet to 500
feet the cuttings showed a granular, cream-colored lime without fos-
sils but tentatively referred to the Tampa. At 500 feet the cuttings
revealed the top of the Eocene, with characteristic Ocala fossils ap-
pearing for the first time.
The Tertiary formations as seen in the outcrop areas in Georgia
are not found in wells on the peninsula of Florida, the entire Eocene
here being limestone. In wells the section below the Ocala has two
generally accepted divisions, the Dictyoconus gunteri zone and an
undescribed Borelis zone. These formations were found in the Brown
well at 945 feet and 2215 feet respectively. The first marks the
top of the Middle Eocene and as such might be regarded as the
equivalent of the Claiborne and the second considered to represent
the combined Wilcox-Midway. It should be kept in mind however
that they are equivalents only in that they are in the same part of
the geologic section. There is no lithologic or paleontologic corre-
lation, and the names Claiborne, Wilcox and Midway should not be
used, at least for the present, in connection with the Eocene for-
mations of peninsular Florida.
The Upper Cretaceous, recognized by the presence of Lepidorbi-
toides and Camerina cf. dickersoni (common in the Cuban Cretaceous),
was encountered at 2985 feet, and at 3280 feet the bit entered a
chalky lime characterized by many Inoceramus prisms that appar-
ently represents the Selma chalk of the outcrop area. The Eutaw


Fig. 1

Sketch Map



Showing location of
geologic section and
deep wells upon which
it is based.

., .;0

3 Ir41


formation was described at the surface in Alabama on its lithologic
character making its recognition in the Florida subsurface diffi-
cult, but on a basis of faunal groupings its top has been put in
this well at 3748 feet. A non-fossiliferous sandy formation entered
at 4547 feet is assigned to the Tuscaloosa, then at 4640 feet the
black shale mentioned above was encountered. This has been de-
scribed as Pennsylvanian or older and a note by the writer appear-
ing in the current Bulletin of the American Association of Petroleum
Geologists suggests its correlation with the Chattanooga shale (lower
Mississipian).lo There are no contained fossils but the writer thinks
that general considerations of paleogeography make it seem the
most likely formation of the Paleozoic to be encountered here. The
publication of this note should provoke a discussion from which
some definite correlation may be determined.11
Cosden Well (Sec. 25, T. 3 S., R. 20 E.) Marion County.-This well,
J. S. Cosden's Lawson No. 1, was drilled in 1928 to a depth of 4434
feet. The Eocene was divided as follows: Ocala 100 feet, Dictyoconus
905 feet, Borelis 1730 feet. The Upper Cretaceous was entered at
2450 feet in beds determined as Maestrichtian in age, and the Tus-
caloosa logged at 3570 feet. Mrs. Applin's report considers all beds
from that depth to the bottom of the hole as Tuscaloosa but the
correlation with the York well ten miles to the south throws some
doubt on this. This will be developed after the discussion of the
York well.
"The York Well," Ocala Oil Corporation, Clark-Ray-Johnson No. 1,
(Sec. 10, T. 16 S., R. 20 E.) Marion County.-The York well, despite
its incomplete record, is probably the most interesting well yet drilled
in the state. The only correlation points available are the Eocene
Dictyoconus at 920 feet and Borelis zone at 2140 feet. Samples
were not furnished throughout the interval where the top of the
Upper Cretaceous should have been found but cuttings from for-
mations of that age were found at lower depths. However at 4100
feet schist and quartzite were found and the bit continued in
quartzite to a total depth of 6180 feet. The Florida Geological Sur-
vey and the United States Geological Survey have tentatively assign-

loCampbell, Robert B., "Paleozoic Under Florida?" American Association of
Petroleum Geologists Bulletin, Vol. 23, No. 11 (November 1930), pp. 1712-13.
11More recently Dr. R. S. Bassler of the National Museum has identified
some ostracods in this shale as a form close to the genus Amphissites and feels
confident of the Chattanooga age of the beds.-Written communication from L.
W. Stephenson.


ed these basement rocks to the pre-Cambrian because of a similarity
to the Piedmont area of the Atlantic Coast States.
A correlation of the Cosden and York wells indicates that both
holes terminated in the same formation, but Mrs. Applin refused
to accept any "basement" determination for the bottom-most rocks
claiming for the Cosden well quartzitic etched sandstone with in-
dividual etched, and grains visible, rather than quartzites and micaceous
shales rather than schists. In this she was supported by some nationally
known petrographers. The writer of the present paper searched the
slush pits of both wells and found pieces of cuttings which he called
quartzite and on the basis of similar lithology it is here correlated with
the Pifiar or San Cayetano of Cuba, a formation made up of a
series of phyllitic shales and sandstones, the phyllitization varying
considerably in short distances, the shales and slightly sericitic shales
alternating with the true phyllites. Its sandstones grade into quartz-
ites with individual grains still visible in some coarser varieties.
This variation within short distances probably explains the difference
in the samples from the Marion County wells. These formations
in Cuba have recently been identified by Dickerson12 as Oxfordian
(Upper Jurassic) in age and this determination is by extension applied
in this paper to the "basement rocks" under the Ocala Uplift. Dr.
Dickerson13 recognizes the possibility of this correlation.
South Lake Well.-In southern Lake County-the Oil Development
Company's J. Ray Arnold No. 1 (Sec. 17, T. 24 S., R. 25 W.), carried
as inactive in current reports, reached a depth of 6129 feet after
penetrating probably all of the Eocene (in which diagnostic fossils were
so altered as to render satisfactory subdivision impossible), entering
the Upper Cretaceous at 3510 feet and a miliolid limestone, here
referred to the Middle Cretaceous, at 5383 feet. At 6050 feet some
red clays, gravels and sands were encountered by the bit. Reports
were widespread that granite had been found, and some samples
circulated as having come from this well were actually found to be
granite, but Mr. Arnold14 denies that his well entered such rocks
and quotes some well-known authorities in support of his statement.
Mr. Arnold has not yet released his samples but they will be looked
for with a great deal of interest as this well is the best sampled

12Dickerson, Roy E., "Cuban Jurassic," Bulletin American Association of Pe-
troleum Geologists, Vol. XIX. No. 1 (1935), pp. 116-118.
--- "The Lower Cretaceous as a Possible Source of Oil in Cuba," Mining
and Metallurgy, September 1937, pp. 418-421.
13--- Oral communication.
14Arnold, J. Ray. Personal communication.


in the state, cores having been taken practically the entire length of
the boring. But granite or not, the miliolid limestones at 5383 feet,
some beds of anhydrite, and the red sands and gravel are regarded
in this paper as representing Middle Cretaceous rocks, and the well
is assumed to be in Jurassic at the bottom of the hole. In this
connection it might be mentioned that Schuchert regards granites in
central Cuba as having been intruded at the end of the Jurassic.
Dickerson also notes the association of granite with his Oxfordian.
In this paper the Cretaceous is divided into three parts conforming
to the practice in Mexico and here called the Eo-Cretaceous, Middle
Cretaceous and Upper Cretaceous. In Texas the division is twofold,
Lower (or Comanchean) and Upper Cretaceous, representing the up-
per two-thirds of the Mexican section. The beds exposed in western
Cuba belong to the Lower Cretaceous of Mexico but the miliolid
limestones below the Upper Cretaceous chalk in Florida wells are
correlated with the Lower Cretaceous of Texas. To avoid confusion
the term "Lower Cretaceous" is dropped from this paper and the
above subdivision made for Florida's subsurface, namely Eo-Cretaceous,
Middle Cretaceous and Upper Cretaceous.
Kissingen Springs Well (Sec. 27, T. 30 S., R. 25E.) in Polk County.
-The Hecksher-Yarnell No. 1, after drilling several hundred feet
of Miocene and Oligocene beds, encountered the Ocala at 275 feet.
The interesting feature of this well is its very thick Eocene section,
especially in the Borelis zone, to the bottom of the hole at 4540 feet.

Peninsular Oil and Refining Company's Cory No. 1 in northeastern
Monroe County (Sec. 6, T. 55 S., R. 34 E.)-The Cory No. 1, a dry
hole completed May 19th of this year at a total depth of 10,006 feet,
is by far the deepest well drilled in the state. The boring commenced
in the Pliocene, entered certain Miocene about 500 feet (though some
beds above that point were questionably referred to that epoch)
and Oligocene at 900 feet. For the Eocene, tops are Ocala 1200 feet,
Dictyoconus at 1800 feet and Borelis at 3300 feet. The Upper Cre-
taceous was encountered about 5700 feet being entirely chalk and
chalky lime to 7676 feet where the formations were slightly shaly
for a little over 100 feet. About 8000 feet the soft limestone was fol-
lowed by a harder white lime with alternations of anhydrite and
gypsum. The limestone was characterized by many Miliolidae and
small fragments of Gryphaea, and is correlated with the Fredericksburg
division of the Comanchean of Texas. One worker found an isolated
Orbitolina walnutensis in the bottom-most core correlating the bottom
of the hole with the Walnut Clay of Texas.

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